Endangered and Threatened Wildlife; 12-Month Finding on a Petition To Identify the Northwest Atlantic Leatherback Turtle as a Distinct Population Segment and List It as Threatened Under the Endangered Species Act, 48332-48421 [2020-16277]
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Federal Register / Vol. 85, No. 154 / Monday, August 10, 2020 / Rules and Regulations
This finding was made on
August 10, 2020.
ADDRESSES: The Status Review Report
are available on NMFS’ website at
https://www.fisheries.noaa.gov/species/
leatherback-turtle.
FOR FURTHER INFORMATION CONTACT:
Jennifer Schultz, NMFS Office of
Protected Resources, (301) 427–8443,
jennifer.schultz@noaa.gov. Persons who
use a Telecommunications Device for
the Deaf (TDD) may call the Federal
Information Relay Service (FIRS) at
1–800–877–8339, 24 hours a day and 7
days a week.
SUPPLEMENTARY INFORMATION:
DATES:
DEPARTMENT OF INTERIOR
Fish and Wildlife Service
50 Part 17
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Parts 223 and 224
[Docket No. 200717–0190]
RIN 0648–XF748
Endangered and Threatened Wildlife;
12-Month Finding on a Petition To
Identify the Northwest Atlantic
Leatherback Turtle as a Distinct
Population Segment and List It as
Threatened Under the Endangered
Species Act
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce; U.S. Fish and Wildlife
Service (USFWS), Interior.
ACTION: Notification of 12-month
petition finding.
AGENCY:
We, NMFS and USFWS,
announce a 12-month finding on a
petition to identify the Northwest
Atlantic population of the leatherback
turtle (Dermochelys coriacea) as a
distinct population segment (DPS) and
list it as threatened under the
Endangered Species Act (ESA). In
response to the petition, we completed
a comprehensive status review of the
species, which also constitutes the 5year review of the species, to determine
potential DPSs following the Policy
Regarding the Recognition of Distinct
Vertebrate Population Segments Under
the ESA and to perform extinction risk
analyses. Based on the best scientific
and commercial data available,
including the Status Review Report, and
after taking into account efforts made to
protect the species, we conclude that
seven populations would meet the
discreteness and significance criteria for
recognition as DPSs, including the
Northwest Atlantic population.
However, even if we were to list them
separately, all seven DPSs would meet
the definition for endangered species
(i.e., they are in danger of extinction
throughout all or a significant portion of
their range). The species is already
listed as endangered throughout its
range. We have determined that the
listing of DPSs is not warranted, and
therefore we do not propose any
changes to the existing global listing.
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SUMMARY:
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Background
The leatherback turtle species as a
whole was listed as an endangered
species (one determined to be
threatened with worldwide extinction)
(35 FR 8491; June 2, 1970), under the
Endangered Species Conservation Act of
1969, the precursor statute to the ESA
(16 U.S.C. 1531 et seq.). When the ESA
was enacted in 1973, it specifically
provided for continuity with the lists
previously in effect under the
Endangered Species Conservation Act.
Section 4(c)(3) of the ESA directed that
species on the lists of endangered
foreign or native wildlife at the time the
ESA took effect would be deemed
‘‘endangered species’’ under the ESA
without interruption. See 39 FR 1444
(January 9, 1974) (explaining transition
provisions); 39 FR 1158, 1172 (January
4, 1974) (setting out the final list of
‘‘endangered foreign wildlife,’’
including ‘‘Turtle, Leatherback’’ at 50
CFR 17.11).
On September 20, 2017, the Blue
Water Fishermen’s Association
petitioned NMFS and USFWS (together,
the Services) to identify the Northwest
(NW) Atlantic leatherback turtle
population as a DPS and to list it as
threatened under the ESA. On December
6, 2017, NMFS published a ‘‘positive’’
90-day finding in the Federal Register
(82 FR 57565) announcing the
determination that the petition
presented substantial information
indicating that the petitioned action
may be warranted. At that time, NMFS
also solicited information on
leatherback turtles and announced that
it would commence, jointly with
USFWS, a status review of the entire
listed species, pursuant to ESA section
4(b)(3)(A) and 50 CFR 424.14. The
resulting Status Review Report includes
all information used to evaluate the
petitioned actions and explains the
process followed by the Status Review
Team (i.e., the Team). The following
summarizes that information; for
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additional details, please see the Status
Review Report (see ADDRESSES).
ESA Statutory, Regulatory, and Policy
Provisions and Evaluation Framework
Under the ESA, the term ‘‘species’’
includes any subspecies of fish or
wildlife or plants, and any DPS of any
vertebrate fish or wildlife which
interbreeds when mature (16 U.S.C.
1532(16)). The Services adopted a joint
policy clarifying their interpretation of
the phrase ‘‘distinct population
segment’’ for the purposes of listing,
delisting, and reclassifying a species
under the ESA (‘‘Policy Regarding the
Recognition of Distinct Vertebrate
Population Segments Under the
Endangered Species Act,’’ 61 FR 4722
(Feb. 7, 1996; ‘‘DPS Policy’’). The DPS
Policy stipulates two elements that must
be considered: (1) Discreteness of the
population segment in relation to the
remainder of the species to which it
belongs; and (2) the significance of the
population segment to the species to
which it belongs.
Section 3 of the ESA defines an
endangered species as any species
which is in danger of extinction
throughout all or a significant portion of
its range and a threatened species as one
which is likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range (16
U.S.C. 1532(6) and (20)). Thus, we
interpret an ‘‘endangered species’’ to be
one that is presently in danger of
extinction. A ‘‘threatened species,’’ on
the other hand, is not presently in
danger of extinction, but is likely to
become so within the foreseeable future
(that is, within a specified later time). In
other words, the primary statutory
difference between a threatened and
endangered species is the timing of
when a species may be in danger of
extinction, either presently
(endangered) or within the foreseeable
future (threatened). The ESA uses the
term ‘‘foreseeable future’’ to refer to the
time over which identified threats are
likely to impact the biological status of
the species. The duration of the
‘‘foreseeable future’’ in any
circumstance is inherently fact-specific
and depends on the particular kinds of
threats, the life-history characteristics,
and the specific habitat requirements for
the species under consideration. The
existence of threats to a species and the
species’ response to such threats are not,
in general, equally predictable or
foreseeable. Hence, in some cases, the
ability to foresee a threat to a species is
greater than the ability to foresee the
species’ exact response, or the
timeframe of such a response, to that
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threat. For purposes of making this 12month finding, the relevant
consideration is whether the species’
population response (i.e., abundance,
productivity, spatial distribution,
diversity) is foreseeable, not merely
whether the emergence of a threat is
foreseeable. The foreseeable future
extends only as far as we are able to
reliably predict the species’ population
response to threats.
Pursuant to the ESA and our
implementing regulations, we determine
whether a species is threatened or
endangered based on any one or a
combination of the following ESA
section 4(a)(1) factors or threats (16
U.S.C. 1533(a)(1), 50 CFR 424.11(c)):
1. The present or threatened
destruction, modification, or
curtailment of its habitat or range;
2. Overutilization for commercial,
recreational, scientific, or educational
purposes;
3. Disease or predation;
4. Inadequacy of existing regulatory
mechanisms; or
5. Other natural or manmade factors
affecting its continued existence, which
could include but are not limited to:
Fisheries bycatch; vessel strikes;
pollution (including marine debris and
plastics, contaminants, oil and gas
activities, and derelict fishing gear);
natural disasters; climate change; and
oceanographic regime shifts.
Section 4(b)(1)(A) of the ESA requires
us to make listing determinations based
solely on the best scientific and
commercial data available after
conducting a review of the status of the
species and after taking into account
efforts being made by any State or
foreign nation or political subdivision
thereof to protect the species’ existence
(16 U.S.C. 1533(b)(1)(A)).
Approach to the Status Review
The Services convened a team of
NMFS and USFWS biologists (i.e., the
Team) to gather and review the best
available scientific and commercial data
on the leatherback turtle, assess the
discreteness and significance of
populations by applying the DPS Policy,
evaluate the extinction risk of any
population segments that meet the DPS
criteria, and document all findings in a
report (i.e., the Status Review Report).
Although the petitioner requested
evaluation only of the NW Atlantic
leatherback population, we instructed
the Team to perform a comprehensive
status review to identify and evaluate
the status of all potential DPSs.
The Team compiled information on
leatherback turtle life history, biology,
ecology, demographic factors, and
threats. This included the information
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received in the petition and in response
to the Federal Register request
associated with the 90-day finding (82
FR 57565; December 6, 2017). The Team
also requested leatherback nesting data
from beach monitoring programs. To
evaluate recent abundance and trends,
unpublished nesting beach monitoring
datasets were often the best available
data (i.e., most recent and relevant). The
Team assessed these data in terms of
standardization (i.e., the use of
standardized methodology), consistency
(i.e., consecutive seasonal data
collection), and duration of data
collection (i.e., the number of years that
data were collected). When evaluating
threats, peer-reviewed information,
specifically primary research with large
sample sizes and long-term sampling
duration, was often the best available
data. In some locations, reports from
governments or non-governmental
organizations and expert opinion
constituted the best available
information. The Team also addressed
the source and magnitude of any
uncertainty and the impact on its
conclusions.
The Team evaluated the discreteness
and significance of each population and
provided their evaluation of whether
each population would meet the criteria
of the DPS Policy. The DPS Policy states
that a population of a vertebrate species
may be considered discrete if it satisfies
one of the following conditions: (1) It is
markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors
(quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation); or
(2) it is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the ESA (61 FR 4722,
February 7, 1996). While the Team used
the term ‘‘DPS’’ in describing and
discussing populations that they
concluded meet the requirements of
discreteness and significance, it is
important to note that the DPS term is
used throughout the Status Review
Report for ease of reference only. A DPS
is formally recognized under the ESA
only upon a listing action by the
Services, and the Services cannot
delegate authority to take formal listing
actions to status review teams. The
information compiled by the Team must
be reviewed by the Services, which
retain responsibility for making the
listing determination after complying
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with all the requirements of Section 4 of
the ESA and considering agency
policies. Because we ultimately
conclude for the reasons discussed in
this finding that it would not be
appropriate to disaggregate the existing
global listing into DPSs, references in
the Status Review Report (and in this
finding when we are reviewing the
information presented by the Team)
must be understood as references to
potential or hypothetical DPSs only.
The Team evaluated significance in
terms of the importance of the
population segment to the overall
welfare of the species, such as: (1)
Persistence of the population segment in
an unusual or unique ecological setting;
(2) evidence that loss of the population
segment would result in a significant
gap in the range of the taxon; (3)
evidence that the DPS represents the
only surviving natural occurrence of a
taxon that may be more abundant
elsewhere as an introduced population
outside its historic range; or (4)
evidence that the population segment
differs markedly from other populations
of the species in its genetic
characteristics.
For each population segment that the
Team determined would meet the
criteria of the DPS Policy (which the
Team and we refer to as a ‘‘DPS’’ for
ease of reference), the Team performed
an extinction risk analysis, which
involved the evaluation of demographic
factors and threats. Demographic factors
reflect the impact that operative threats
have had on the species. In some cases
those threats or the impacts from the
threats are continuing in nature. The
demographic factors included
abundance, productivity, spatial
distribution, and diversity. Because sea
turtles spend the majority of their lives
at sea, where they are spread across vast
distances, it is difficult to estimate total
abundance. However, the number of
nesting females can be counted directly,
or estimated indirectly by counting the
number of nests on beaches, during a
nesting season. Females nest more than
once in a season (i.e., clutch frequency,
which is the average number of nests
per season) and do not nest every season
(i.e., remigration interval, which is the
average number of years between
successive nesting seasons). To
calculate the index of nesting female
abundance at a nesting beach, the Team
summed the total number of nests over
the most recent remigration interval
(i.e., a run-sum) and divided this
number by the clutch frequency. The
Team performed these calculations only
if available data were recent (i.e., last
year of the remigration interval occurred
in 2014 or more recently), consistent
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(i.e., seasonal data collected for each
year of the remigration interval), and
collected in a standardized manner (i.e.,
data collection methods remained the
same over the remigration interval), as
further detailed in the Status Review
Report. To provide a total index of
nesting female abundance for each DPS,
we summed the indices of nesting
female abundance for all monitored
beaches used by that DPS. The total
index of nesting female abundance for
each DPS is an index (rather than a
census) because not all nesting beaches
met these criteria. However, the nesting
beaches that were not included were
generally unmonitored or not recently
monitored because they host few
nesting females. Even where data were
not sufficient to allow for a calculation
of the index of nesting female
abundance, the Team provided all
available data to ensure the analysis
would be as robust as possible.
The Team evaluated the productivity
for each DPS by evaluating nesting
trends (through trend analyses or bar
graphs) and productivity metrics. Where
available data allowed it, they estimated
the long-term trend for individual
beaches using a Bayesian state-space
model of stochastic exponential
population growth (Boyd et al. 2017),
where the rate parameter describes the
annual percent change in observed nest
counts (or female counts where
applicable) over the period of data
collection. This is further explained in
the Status Review Report. To reflect
current trends over approximately three
remigration intervals, the criteria for
trend analyses were as follows: Nesting
data (i.e., nest or nesting female counts)
consistently collected over nine or more
years in a standardized manner (for that
site), with the most recent data
collection in 2014 or later and with a
minimum average number of nests of 50
annually. The Team reported the
median trend, along with the standard
deviation (sd), 95 percent credible
interval (CI), and an ‘‘f statistic’’ which
is the proportion of the posterior
distribution with the same sign as the
median (i.e., the confidence that the
trend is positive or negative). When the
data did not meet the criteria for
performing trend analyses, the Team
provided bar graphs and/or historical
data in the Status Review Report. Based
on the trend analysis (where possible)
and the best available historical data,
the Team characterized the nesting
trend for each DPS as decreasing, stable,
or increasing. The Team also evaluated
the following productivity metrics (if
available): Average size of nesting
female; nesting female survivorship;
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remigration interval; clutch size; clutch
frequency; internesting interval;
incubation period; hatching success (the
proportion of eggs in a nest that produce
live hatchlings); and sex ratio. Each of
these metrics contributes to the growth
rate, or reproductive potential, of the
population.
For each DPS, the Team evaluated
spatial distribution, which included the
number and location of nesting beaches
and foraging areas, as well as spatial
structure (i.e., whether the DPS exists as
a single population or several
subpopulations connected by
metapopulation dynamics). The Team
also evaluated diversity, which like
spatial distribution, is a measure of
resilience. In general, diverse
populations with broad spatial
distributions and metapopulation
dynamics are more resilient to threats
and environmental changes than less
diverse populations with narrow
distributions.
For each DPS, the Team next
evaluated each of the ESA Section
4(a)(1) factors (or ‘‘threats’’) as listed
above (16 U.S.C. 1533(a)(1), 50 CFR
424.11(c)). For each threat, the Team
used the best available information to
describe the threat, identify which life
stages are affected, and describe the
impact to the DPS with as much
specificity as the best available
information allowed to link the threat to
the demographic factor it affected. The
best available data often allow only for
qualitative assessment. For each DPS,
the Team identified the primary
threat(s) to its continued existence, as
well as other threats. The Team
considered the impact of each threat
individually, with the primary threat(s)
given the greatest weight, and all threats
cumulatively, to determine the
extinction risk. To assess confidence in
the extinction risk determination, the
Team identified any sources of
uncertainty and the impact of
uncertainty on the conclusions. They
analyzed all threats assuming the DPS
had lost ESA protections going forward
because a DPS would not receive such
protections if it was not listed under the
ESA. For example, a DPS would not
have benefits of section 9 take
prohibitions or section 7 consultations
on actions that may affect the DPS.
The Team performed an extinction
risk assessment for each of the seven
DPSs by evaluating the demographic
factors and threats, as described above.
Then, the Team voted, based on the best
available data, on whether the
extinction risk of each DPS was high,
moderate, or low, following the
definitions included in NMFS’ internal
guidance document, ‘‘Guidance on
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Responding to Petitions and Conducting
Status Reviews under the Endangered
Species Act, Section II’’ (i.e., NMFS’
Guidance; November 9, 2017) and in the
Status Review Report.
After the Team completed its draft
Status Review Report, the Services met
to review and discuss that document
and conservation efforts. The Services
based our status determinations of the
DPSs on the best scientific and
commercial data available (as compiled
and reflected in the Status Review
Report) and after taking into account
efforts by States and foreign nation, or
any political subdivision thereof, to
protect the species as mandated by the
statute.
DPS Analysis
The following is a summary of the
DPS analysis conducted by the Team.
For a detailed description of the Team’s
analyses of discreteness and
significance, please see the Status
Review Report. As a starting point, the
Team considered seven leatherback
populations that were previously
identified as regional management units
(RMUs) by Wallace et al. (2010) and
recognized as subpopulations under the
International Union for Conservation of
Nature (IUCN) Red List (https://
www.iucnredlist.org/species/6494/
43526147). The Team found that seven
leatherback populations met the
discreteness and significance criteria
per the DPS Policy and identified the
following potential DPSs: Northwest
(NW) Atlantic; Southwest (SW) Atlantic;
Southeast (SE) Atlantic; SW Indian;
Northeast (NE) Indian; West Pacific; and
East Pacific.
Discreteness
The Team evaluated all populations
for discreteness and determined that
each showed marked separation from
the others as a consequence of
behavioral and physical factors.
Behavioral factors, especially returning
to waters off a turtle’s natal beach to
breed, have prevented interbreeding,
resulting in reproductive isolation, as
indicated by genetic discontinuity.
Although some populations use the
same foraging areas, tagging and
telemetry studies also demonstrate the
discreteness of the populations at
nesting beaches. Physical factors, such
as land masses, ocean currents, and
other oceanographic features, have
established and reinforced barriers to
gene flow among the seven populations.
Genetic data provide the most
compelling evidence for discreteness
among the seven populations. The most
recent and comprehensive global
analysis of published and unpublished
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mitochondrial deoxynucleic acid
(mtDNA) sequence data (i.e., 28
haplotypes, which are unique sequences
of mtDNA) evaluated samples collected
from 21 nesting sites representing key
regions from all ocean basins (Dutton et
al. 2007; Dutton et al. 2013; Shanker et
al. 2011; Dutton and Shanker 2015);
analyzing the evolutionary relationship
of these data revealed three distinct
haplogroups (i.e., similar haplotypes
that cluster together, relative to other
haplotypes) that are geographically
segregated across the Atlantic, Indian,
and Pacific Oceans (Dutton,
unpublished data; NMFS and USFWS
2020). Early mtDNA analyses indicated
strong genetic discontinuity, globally
(FST = 0.415, P <0.001) and within ocean
basins (FST = 0.203 to 0.253, P <0.001;
Dutton et al. 1999). Wallace et al. (2010)
combined these and other genetic data
with nesting, flipper tagging, and
satellite telemetry data to identify seven
leatherback RMUs, which provided the
starting point for our identification of
discrete populations.
From this starting point, the Team
then evaluated more recent genetic data.
Subsequent genetic analyses confirmed
genetic discontinuity among the NW,
SW, and SE Atlantic populations
(Wallace et al. 2010; Dutton et al. 2013;
Carreras et al. 2013; Molfetti et al. 2013;
Vargas et al. 2017). Elevated genetic
differentiation at nuclear DNA (FST =
0.211¥0.86) indicates that males, like
females, likely return to the waters off
their natal beaches to mate and that
male-mediated gene flow may not be as
pronounced as previously thought
(Dutton et al. 2013; see Jensen et al.
2013). Nuclear (FST >0.126, P <0.001;
Dutton et al. 2013) and mtDNA (FST
>0.061, P = 0.05¥0.001; Dutton et al.
2013; FST >0.061, P <0.01; Vargas et al.
2017) analyses indicate genetic
discontinuity between the Atlantic
populations and the SW Indian
population. Preliminary mtDNA results
for leatherback turtles nesting at Little
Andaman Island, India (Shanker et al.
2011; Dutton and Shanker 2015),
indicate that this population is closely
related to the extinct Malaysian
population, with which it shares
common haplotypes. It is markedly
different from the South African nesting
population, as well as those in the West
Pacific population (Dutton et al. 2007,
2013 and unpublished). Samples from
extant and extirpated nesting
aggregations of the NE Indian
population (Shanker et al. 2011; Dutton
and Shanker 2015; Dutton et al.
unpublished data) are genetically
differentiated from the SW Indian
population (FST = 0.415, P <0.003;
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Dutton et al. 1999) and the West Pacific
population (X2 = 49.346, P = 0.002;
Dutton et al. 2007). There is genetic
discontinuity between the West and
East Pacific populations, as
demonstrated by significant genetic
differentiation between the samples
from Solomon Islands in the western
Pacific and Mexico or Costa Rica in the
eastern Pacific (FST = 0.270 and 0.331,
P <0.001; Dutton et al. 1999). Genetic
discontinuity among all seven
populations provides evidence for
marked separation from the others and
thus discreteness of each population.
Tagging and telemetry studies confirm
marked separation of the seven
populations because nesting sites
remain distant and isolated. Nesting
females of one population have not been
tracked to, or observed on, beaches used
by another population, even though
telemetry data indicate shared use of
foraging areas by different populations.
Telemetry studies demonstrate that
females nesting on NW Atlantic beaches
move throughout most of the North
Atlantic from the Equator to about 50°
N latitude (Ferraroli et al. 2004; Hays et
al. 2004; James et al. 2005a; James et al.
2005b; 2005c; Eckert 2006a; Eckert et al.
2006b; Hays et al. 2006; Doyle et al.
2008; Evans 2008; Dodge et al. 2014;
Fossette et al. 2014; Aleksa 2017; Aleksa
et al. 2018). Turtles originating from
beaches of the NW Atlantic appear to
mix at foraging areas throughout the
North Atlantic Ocean (Fossette et al.
2014), but their movements rarely
extend into waters south of the Equator.
Tagging studies further support the
connectivity within and among nesting
beaches and foraging areas of the North
Atlantic Ocean (Troe¨ng et al. 2004;
Bra¨utigam and Eckert 2006; Chaco´nChaverri and Eckert 2007; Turtle Expert
Working Group (TEWG) 2007; So¨nmez
et al. 2008; Dutton et al. 2013b;
Horrocks et al. 2016), but turtles tagged
in the North Atlantic Ocean have never
been found on nesting beaches in Brazil
(SW Atlantic population) or Africa (SE
Atlantic population). In the South
Atlantic Ocean, post-nesting females
tracked from nesting beaches in Gabon
and Brazil use the same foraging areas,
including waters off SW Africa, in the
south equatorial Atlantic and off SE
Brazil and Uruguay (Almeida et al.
2011; Witt et al. 2011). Turtles
incidentally captured in fisheries off
South America (Billes et al. 2006,
Lo´pez-Mendilaharsu et al. 2009) also
demonstrate that turtles originating from
the SW and SE Atlantic Ocean beaches
share foraging areas. Despite such
mixing at foraging areas, there is no
evidence for the shared use of nesting
beaches. Genetic data indicate that
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turtles return to their natal beaches to
nest on opposite sides of the Atlantic
Ocean (Dutton et al. 2013; Vargas et al.
2017), and no tag recoveries contradict
these data.
In the Indian Ocean, telemetry studies
have been conducted at South African
nesting beaches in the SW Indian Ocean
(Hughes et al. 1998; Luschi et al. 2006;
Robinson et al. 2016) and at Andaman
Islands nesting beaches in the NE Indian
Ocean (Namboothri et al. 2012;
Swaminathan et al. 2019). South
African nesting females showed diverse
movements that were highly influenced
by complex oceanographic currents and
features that lead them to foraging
destinations in the South Atlantic
Ocean, SW Indian Ocean, and
Mozambique Channel (Hughes et al.
1998, Luschi et al. 2006, Robinson et al.
2016). About half of the 10 post-nesting
females tagged at the Andaman Islands
moved westward: Two individuals
reached the Mozambique Channel; the
other half moved southeastward, past
the Indonesian islands of Sumatra and
Java, with one leatherback reaching an
apparent foraging ground off NW
Australia before transmissions stopped
(Namboothri et al. 2012; Swaminathan
et al. 2019). Despite overlap in one
foraging area (i.e., reaching the
Mozambique Channel), tagging data do
not indicate movement between the
distant nesting beaches.
Within the Pacific Ocean, nearly all
turtles tracked from East Pacific nesting
beaches moved southward across the
Equator to forage in open-ocean waters
of the SE Pacific Ocean or in the coastal
waters of Central America, Peru, and
Chile. The movements of post-nesting
females from the West Pacific Ocean are
dependent on the season in which they
nest, with winter-nesting females
predominantly tracked into the
Southern Hemisphere and summernesting females foraging in diverse
coastal and oceanic ecosystems
throughout the northern Indo-Pacific
region (Benson et al. 2011). Telemetry
data indicate little or no overlap with
foraging destinations utilized by nesting
females of the East and West Pacific
populations (Bailey et al. 2012; Benson
et al. 2011). However, a genetic study of
bycaught turtles off the coast of Chile
and Peru indicated that 15 percent of
leatherback turtles originated from West
Pacific nesting beaches (Donoso and
Dutton 2010), suggesting that foraging
overlap may be more prevalent than
estimated by telemetry data. Still, there
is no genetic evidence for contemporary
interbreeding between the two
populations (Dutton et al. 2007), and
telemetry and tagging data do not
indicate movement between the distant
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nesting beaches. Thus, flipper tagging
and satellite telemetry data support the
marked separation, and thus
discreteness, of the seven populations at
their nesting beaches.
Physical factors likely shape and
reinforce the behavior patterns that
result in reproductive isolation. Though
the species has a global range, with
foraging areas extending into high
latitudes, nesting mainly occurs on
tropical or subtropical beaches. Posthatchling dispersal is determined by the
ocean currents they encounter off
nesting beaches. While adults move
throughout tropical and temperate
waters irrespective of ocean currents,
both males and females return to the
waters off their natal nesting beach to
mate. This natal homing is somewhat
flexible, (Dutton et al. 2013; Jensen et al.
2013), creating reproductive isolation
only among distant nesting sites, which
may also be physically separated from
one another by land masses and
oceanographic barriers to gene flow. For
example, leatherback turtles in the
Atlantic Ocean are physically separated
from those in the Pacific Ocean by the
Americas. Though leatherback turtles
have greater cold tolerance than other
sea turtles, they do not appear to
venture into latitudes greater than 47° S
or 71° N (Eggleston 1971; Eckert et al.
2012). Therefore, the low latitude and
cold waters of the Cape Horn Current
likely prevent movement between the
Atlantic and Pacific Oceans. Within
ocean basins, nesting beaches of the
discrete populations are separated by
long distances of uninterrupted deep
water (e.g., the East Pacific Barrier and
the mid-Atlantic Barrier). While
leatherback turtles clearly cross these
open-ocean barriers to reach distant
foraging areas, they do not appear to do
so for nesting and breeding, but rather
return to their natal region to breed and
nest (Barragan et al. 1998; Dutton et al.
1999; Barragan and Dutton 2000; Dutton
et al. 2013). Within ocean basins,
currents shape post-hatchlings’
movement patterns, which they may
retain as adults (e.g., Fossette et al.
2010; Benson et al. 2011). The NW
Atlantic leatherback population appears
to be physically separated from the SE
and SW Atlantic populations by the
current systems of the South and North
Atlantic Gyres, respectively. NW
Atlantic leatherback nesting beaches are
adjacent to northward moving currents
(e.g., Gulf Stream). Leatherback
hatchlings from these nesting beaches,
therefore, are transported northward,
remaining in the North Atlantic Ocean.
Those that survive return to their
nesting beaches as adults, completing
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their life stages within the North
Atlantic (Fossette et al. 2010; Chambault
et al. 2017). The SE and SW Atlantic
populations are similarly retained in the
South Atlantic Ocean by the South
Atlantic Gyre and the Benguela Current,
which flows northward along the SE
coast of Africa, restricting movement
into the Indian Ocean. Within the
Indian Ocean, the Somali Current runs
between the nesting beaches of the SW
and NE Indian populations. The NE
Indian and West Pacific populations
likely became isolated as a result of
exposed land barriers between
Indonesia, New Guinea, and the
Philippines as a result of low sea levels
within the past 6,000 years (Barber et al.
2000). Seasonal monsoons may also
play a contemporary role by altering
current directions and hatchling
dispersal patterns (Benson et al. 2011;
Gaspar et al. 2012). Thus, physical
factors have likely helped to shape, or
at least reinforce, the reproductive
isolation among distant nesting beaches.
Based on these data, the Team
concluded that the seven populations
demonstrate discreteness, or marked
separation from each other, due to
behavioral and physical factors. These
are the NW Atlantic, SW Atlantic, SE
Atlantic, SW Indian, NE Indian, West
Pacific, and East Pacific populations.
Significance
Each of the discrete populations is
significant to the species because the
loss of any one would result in a
significant gap (i.e., a half or quarter of
an ocean basin) in the range of the
species. Several populations also persist
in unique ecological settings. Each
population likely possesses unique
genetic characteristics and local
adaptations as a result of thousands of
years of reproductive isolation, but none
have yet been identified because all
genetic studies have involved neutral
markers. Therefore, the Team did not
rely on evidence of unique genetic
characteristics and local adaptations for
its significance finding.
A loss of the NW Atlantic population
would result in a gap (i.e., the entire
North Atlantic Ocean) of the nesting and
foraging range of the species. If the NW
Atlantic population were extirpated, it
is unlikely that leatherback turtles from
other populations would recolonize the
North Atlantic Ocean in an ecological
time frame (i.e., tens to hundreds of
years), leaving a significant gap in the
range of the species. Extirpation of this
population would also significantly
reduce the genetic diversity of the
species, as reflected by the possession of
several unique haplotypes. Leatherback
turtles of the NW Atlantic Ocean also
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occur in a unique ecological setting; this
is the only DPS that regularly forages at
high latitudes. Sightings have been
documented as far north as Norway and
Iceland (Brongersma 1972; Goff and
Lien 1988; Carriol and Vader 2002;
McMahon and Hayes 2006; Eckert et al.
2012). Such high latitude foraging is
likely facilitated by the warm Gulf
Stream, which meets cold water
currents to create highly productive
foraging areas. The Team concluded that
the NW Atlantic population is
biologically significant to the species.
In the SW Atlantic Ocean, leatherback
turtles only nest in a small area of the
coastline of Brazil. All other nesting in
South America occurs above the Equator
or on the Pacific Coast. Therefore, the
loss of this population would result in
a gap of the nesting range of the species
(i.e., the SW Atlantic coast). Although
SE Atlantic leatherback turtles forage off
the coasts of Brazil, Argentina, and
Uruguay, they do not breed there.
Rather, they return to the waters off
western Africa to mate (Vargas et al.
2017). Therefore, if the SW Atlantic
population were extirpated, it is
unlikely that leatherback turtles from
other populations would recolonize this
region, leaving a significant gap in the
nesting range of the species. The
extirpation of this population would
also significantly reduce the genetic
diversity of the species, as reflected by
the possession of unique haplotypes and
high genetic diversity, despite the small
population size (Vargas et al. 2017). The
SW Atlantic population is biologically
significant to the species.
Leatherback turtles of the SE Atlantic
population nest in West Africa and
forage in the South Atlantic Ocean. This
population is much more abundant than
the SW Atlantic population, which also
forages in the South Atlantic Ocean.
Therefore, the loss of this population
would result in a gap of the nesting
range of the species (i.e., western Africa)
and a significant reduction in the
abundance of leatherback turtles
foraging throughout the South Atlantic
Ocean. The extirpation of this
population would also significantly
reduce the genetic diversity of the
species, as reflected by the possession of
unique haplotypes. The Team
concluded that the SE Atlantic
population is biologically significant to
the species.
In the SW Indian Ocean, leatherback
turtles only nest in a small area along
the South African and Mozambican
coastlines. No other leatherback turtles
nest in eastern Africa or in other areas
throughout the entire western Indian
Ocean. Therefore, the loss of this
population would result in a gap of the
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nesting range of the species (i.e., the SW
Indian Ocean). The SW Indian
population also occurs in a unique
ecological setting: It is the only
population to nest on temperate
beaches. The warm Agulhas Current,
adjacent to the nesting beaches, likely
facilitates their high-latitude nesting.
The Team concluded that the SW Indian
population is biologically significant to
the species.
Leatherback turtles nest in small
numbers in the NE Indian Ocean. These
nesting sites are separated from other
Indian Ocean nesting sites by at least
5,000 km. Although western Pacific
nesting sites are closer, males and
females return to the waters off their
natal beaches to breed, preventing
interbreeding among NE Indian and
West Pacific populations. Therefore, the
loss of this population would result in
a gap of the nesting range of the species
(i.e., the NE Indian Ocean). The
extirpation of this population would
also significantly reduce the genetic
diversity of the species, as reflected by
the possession of unique haplotypes.
The Team concluded that the NE Indian
population is biologically significant to
the species.
West Pacific leatherback turtles nest
in small numbers primarily in four
nations of the West Pacific Ocean. These
nesting sites are separated from East
Pacific nesting sites by over 10,000 km.
Though NE Indian nesting sites are
closer in distance, male and female
philopatry prevents interbreeding.
Therefore, the loss of this population
would result in a gap of the nesting
range of the species (i.e., the West
Pacific Ocean). The loss of this
population would also result in a gap of
the foraging range of the species (i.e., the
North Pacific Ocean). The extirpation of
this population would also significantly
reduce the genetic diversity of the
species, as reflected by the possession of
unique haplotypes. The West Pacific
population is ecologically unique in two
ways: It is the only population to forage
in both hemispheres; and it nests yearround, with nesting peaks in the
summer and winter. The Team
concluded that the West Pacific
population is biologically significant to
the species.
Leatherback turtles nesting on eastern
Pacific coastlines also forage in the East
Pacific Ocean. A loss of this population
would result in a gap of the nesting
range of the species (i.e., the East Pacific
Ocean). Though West Pacific
leatherback turtles may forage off the
coasts of Peru and Chile, they do not
breed there (Donoso and Dutton 2010).
Therefore, if the East Pacific population
were extirpated, it is unlikely that
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leatherback turtles from other
populations would recolonize this
region, leaving a significant gap in the
nesting range of the species. The
extirpation of this population would
also significantly reduce the genetic
diversity of the species, as the
population possess several unique
haplotypes. The East Pacific population
is unique in having the smallest nesting
female size, clutch size, and egg size of
all populations, possibly reflecting
unique foraging conditions that are
subject to oceanographic regime shifts
(e.g., the El Nin˜o Southern Oscillation,
or ENSO). The Team concluded that the
East Pacific population is biologically
significant to the species.
DPS Summary
The Team found that seven
populations met the definition for
discreteness. These populations are
markedly separated as a result of the
behavioral factors of movement (as
demonstrated by satellite telemetry and
flipper tagging studies) and philopatry,
which has led to reproductive isolation
(as demonstrated by genetic
discontinuity). They are also physically
separated by land masses,
oceanographic features, and currents.
The Team found these seven
populations to be significant to the
species because the loss of any one of
them would result in a significant gap
in the range of the species as well as a
significant loss of genetic diversity,
reducing the evolutionary potential of
the species. Some populations also
occur in a unique ecological setting.
Thus, after reviewing the best available
information, the Team identified the
following populations as potential
DPSs: NW Atlantic, SW Atlantic, SE
Atlantic, SW Indian, NE Indian, West
Pacific, and East Pacific. The Team
defined the potential DPSs as
leatherback turtles originating from
nesting beaches within the boundaries
for each DPS. The range of each DPS,
which also includes foraging areas, thus
extends beyond the nesting boundaries
for most DPSs, and may overlap
extensively with the range of another
DPS. The boundaries are based on the
best available genetic, telemetry, and
observational data. When such data
were not available, the Team used
information on possible barriers to gene
flow, such as oceanographic features.
For ease of use, the Team applied
political boundaries when this did not
conflict with biological or
oceanographic data. Additional
information on the boundaries is
available in the following sections,
which summarize the extinction risk
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48337
analysis for each DPS, and in the Status
Review Report.
NW Atlantic DPS
The Team defined the NW Atlantic
DPS as leatherback turtles originating
from the NW Atlantic Ocean, south of
71° N, east of the Americas, and west of
Europe and northern Africa; the
southern boundary is a diagonal line
between 5.377° S, 35.321° W and
16.063° N, 16.51° W. The northern
boundary reflects a straight latitudinal
line based on the northernmost
documented occurrence of leatherback
turtles (Brongersma 1972; Goff and Lien
1988; Carriol and Vader 2002; McMahon
and Hayes 2006; Eckert et al. 2012). The
southern boundary is a diagonal line
between the elbow of Brazil, where the
Brazilian current begins and likely
restricts the nesting range of this DPS,
and the northern boundary of Senegal.
The boundary between Senegal and
Mauritania was chosen because the SE
Atlantic DPS does not appear to nest
above this boundary (Fretey et al. 2007).
The range of this DPS (i.e., all areas
of occurrence) extends throughout the
North Atlantic Ocean, including the
Caribbean Sea, Gulf of Mexico (GOM),
and Mediterranean Sea. Available data
indicate that the NW Atlantic DPS
occurs (at varying levels of frequency) in
the waters of the following nations or
territories: Albania, Algeria, Anguilla,
Antigua and Barbuda, Aruba, Azores,
Bahamas, Barbados, Belize, Bermuda,
Bonaire, Bosnia and Herzegovina,
Brazil, British Virgin Islands, Canada,
Cape Verde, Cayman Islands, Colombia,
Costa Rica, Croatia, Cuba, Curac
¸ao,
Cyprus, Denmark, Dominica, Dominican
Republic, Egypt, France, French Guiana,
Greece, Greenland, Grenada,
Guadeloupe, Guatemala, Guyana, Haiti,
Honduras, Iceland, Ireland, Israel, Italy,
Jamaica, Lebanon, Libya, Madeira,
Malta, Martinique, Mauritania, Mexico,
Montenegro, Montserrat, Morocco,
Netherlands Antilles, Nicaragua,
Norway, Panama, Portugal, Slovenia,
Spain, St. Barthelemy, St. Eustatius, St.
Kitts and Nevis, St. Lucia, St. Maarten,
St. Pierre and Miquelon, St. Martin, St.
Vincent and the Grenadines, Suriname,
Sweden, Syria, Trinidad and Tobago,
Tunisia, Turkey, Turks and Caicos
Islands, United Kingdom, United States
(including Puerto Rico and the U.S.
Virgin Islands (USVI), Venezuela, and
Western Sahara.
All nesting in this DPS occurs in the
NW Atlantic Ocean, concentrated from
the southeast United States throughout
the Wider Caribbean Region (Dow et al.
2007). Leatherback nesting in the NW
Atlantic can be grouped into several
broad geographical areas, including the
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U.S. mainland (primarily Florida),
North Caribbean (including USVI and
Puerto Rico), West Caribbean (Honduras
to Colombia), and Southern Caribbean/
Guianas (Venezuela to French Guiana;
TEWG 2007). The largest nesting
aggregations occur in Trinidad, French
Guiana, and Panama. The northern-most
confirmed nesting occurs in North
Carolina, but there has been a crawl
recorded as far north as Assateague
Island National Seashore, Maryland
(Rabon et al. 2003). No nesting occurs in
the Mediterranean Sea (Casale and
Margaritoulis 2010).
Nesting occurs on unobstructed, highenergy beaches with either a deep water
oceanic approach or a shallow water
approach with mud banks, but without
coral or rock formations (TEWG 2007).
The main characteristics of leatherback
nesting beaches include coarse-grained
sand; steep, sloping littoral zone;
obstacle-free approach; proximity to
deep water; and oceanic currents along
the coast (Hendrickson and Balasingam
1966 in Eckert et al. 2015). During the
nesting season, adult females and males
inhabit the waters off nesting beaches.
During a nesting season, females
generally stay within about 100 km of
their nesting beaches, remaining close to
the coast on the continental shelf, and
engaging in shallow dives (Eckert et al.
2012). Intra-seasonal movement of
greater than 100 km also occurs,
especially between French Guiana and
Suriname (Fossette et al. 2007; Georges
et al. 2007), Panama and Costa Rica
(Chaco´n-Chaverri and Eckert 2007), and
among Caribbean nesting beaches,
including those on Trinidad (Brautigam
and Eckert 2006; Georges et al. 2007;
Horrocks et al. 2016). Adult males
migrate from temperate foraging areas in
the North Atlantic Ocean to waters off
nesting beaches, typically arriving
before the nesting season and remaining
for the majority of the season (James et
al. 2005b; Doyle et al. 2008; Dodge et al.
2014).
Foraging areas of the NW Atlantic
DPS include coastal and pelagic waters
of the North Atlantic Ocean (Eckert et
al. 2012; Saba 2013; Shillinger and
Bailey 2015). These waters include the
GOM, North Central Atlantic Ocean,
northwestern Atlantic shelf waters of
the United States and Canada, waters
along the southeastern U.S. coast, the
Mediterranean Sea, and the northeastern
Atlantic shelf waters of Europe and
northwestern Africa (TEWG 2007).
Some post-nesting females also remain
in tropical waters to forage (Fossette et
al. 2010). This DPS is mostly commonly
associated with open-ocean and coastal
shelf foraging areas off Nova Scotia
(Canada), northeastern United States,
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GOM, northwestern Europe, and
northwestern Africa (James et al. 2005a,
2006b, 2007; Eckert 2006; Eckert et al.
2006; Fossette et al. 2010a; Fossette et
al. 2010b; Dodge et al. 2014; Stewart et
al. 2016; Aleksa et al. 2018). Fossette et
al. (2014) analyzed available satellite
telemetry data from 1995 to 2010 on
post-nesting females (n = 93) as well as
males (n = 4), females (n = 8), and a
juvenile (n = 1) from foraging grounds
throughout the Atlantic Ocean. They
found widespread use of the North
Atlantic Ocean (Fossette et al. 2014).
High-use areas mainly occurred in the
central (25 to 50° N, 50 to 30° W) and
eastern Atlantic Ocean, in particular in
the waters offshore Western Europe,
around Cape Verde (year-round) and the
Azores (October to March; Fossette et al.
2014). Fossette et al. (2014) found that
seasonal high-use areas also occurred
along the eastern U.S. coast (April to
June and October to December) and off
Canada (July to December). The GOM is
also a high-use foraging area, with a
peak in the northeast GOM during
August and September (Aleksa et al.
2018). Overall, leatherback turtles of the
North Atlantic population appear to
have a diverse array of foraging habitat
available.
Abundance
The total index of nesting female
abundance for the NW Atlantic DPS is
20,659 females. The nesting beaches
with the greatest abundance have been
included in this index, and most
beaches with an unquantified number of
nests likely host few nesting females.
We based this index on 24 nesting
aggregations in 10 nations: Trinidad and
Tobago (n = 11,324), French Guiana (n
= 2,519), Panama (n = 2,251), United
States (n = 1,694), Costa Rica (n =
1,306), Suriname (n = 698), Grenada (n
= 499), Venezuela (n = 215), Guyana (n
= 76), and Nicaragua (n = 10). With the
possible exception of Colombia, our
total index does not include 31
unquantified but likely small nesting
aggregations for which data are not
available. It also does not include
outdated data published by Dow et al.
(2007), which includes binned crawls,
categorized as less than 25, 25 to 100,
100 to 500, 500 to 1000, or unknown
abundance. Crawls or emergences
(measured as females or tracks on
beaches) include both successful egglaying and unsuccessful nesting, so the
number of crawls represents
approximately two to 10 times the
number of nests (Dow et al. 2007).
Because the Dow et al. data, which are
more than 10 years old and do not
provide the number of actual nests, may
not be representative of recent nesting
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trends, we did not include them in our
total index. To calculate the indices of
nesting female abundance, we added the
number of nests over the last 3 years
(representing the most recent
remigration interval; Eckert et al. 2012)
and divided by the clutch frequency
(site-specific values or, when such
values were not available, the average of
the site-specific values, i.e., 5.5 clutches
per season).
Our total index of nesting female
abundance is based on the best available
data for this DPS. It is the most robust
estimate of nesting females at this time
because it only includes available
nesting data from recently and
consistently monitored nesting beaches.
Our total index does not include data
from beaches where we were unable to
quantify the number of nesting females,
either due to the lack of recent or
available nesting data or because only
crawl data were reported (often on
smaller nesting beaches). Scattered
nesting may occur on beaches
throughout the region, but because these
beaches are not monitored, or have not
been recently monitored, recent data are
not available.
Nesting in the NW Atlantic DPS is
characterized by many small nesting
beaches. Large nesting aggregations are
rare; only about 10 leatherback nesting
beaches in the Wider Caribbean Region
(about two percent of the DPS’s total
nesting sites) host more than 1,000
crawls annually (Dow Piniak and Eckert
2011). Only one site, Grande Riviere in
Trinidad, hosts more than 5,000 nesting
females, representing 29 percent of the
total index of nesting female abundance.
Relatively large nesting aggregations are
also found in Matura (Trinidad),
Chiriqui Beach (Panama), and Cayenne
and Remire Montjoly (French Guiana).
In contrast, most known nesting beaches
support a small nesting female
abundance; 71 percent of the total
nesting sites record annual crawls of
less than 100 (Dow Piniak and Eckert
2011). The number of nesting females is
unquantified at 31 beaches (i.e., the
majority of nesting sites for the DPS).
However, for the reasons identified
above, most of those sites have small
abundance levels as inferred from the
numbers of crawls estimated by Dow et
al. (2007). Therefore, our total index of
nesting female abundance represents the
most robust estimate allowed by the best
available data and includes the majority
of nesting females because the largest
nesting aggregations were included. The
data regarding additional nesting
aggregations are not sufficiently recent,
specific, or reliable for inclusion, and
the contribution of these nesting
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aggregations to the total index is
expected to be small.
Our total index of nesting female
abundance is similar in comparison to
other published estimates. TEWG (2007)
estimated the abundance of NW Atlantic
leatherback turtles using nesting data
from 2004 and 2005. At that time, the
number of adult females (equating to
total index of nesting female abundance
in our analysis) was estimated to be
approximately 18,700 (range 10,000 to
31,000). While a wide range was
provided, the point estimate in TEWG
(2007) is similar to, albeit slightly lower
than, our total index of 20,659 nesting
females. The most recent, published
IUCN Red List assessment for the NW
Atlantic Ocean subpopulation estimated
a total of 20,000 mature individuals
(The NW Atlantic Working Group 2019).
Our total index, which only includes
nesting females, exceeds their estimate,
likely due to our use of a 3-year
remigration interval, which has
increased at some locations in recent
years (e.g., 4.5 years at St. Croix; K.R.
Stewart, The Ocean Foundation and C.
Lombard, USFWS, pers. comm., 2019).
We conclude that the total index of
nesting females for the NW Atlantic DPS
is 20,659 females. The nesting beaches
with the greatest abundance have been
included in our total index, and most
beaches with an unquantified number of
nests likely host few nesting females.
Current nesting female abundance is not
at a level where stochastic or
environmental changes would have
catastrophic impacts, but the abundance
at several nesting sites with previously
high density has declined drastically.
However, as we discuss below, a
declining nest trend and several existing
threats will likely continue to reduce
this abundance.
Productivity
The NW Atlantic DPS exhibits
decreasing nest trends at nesting
aggregations with the greatest indices of
nesting female abundance. Though
some nesting aggregations indicate
increasing trends, most of the largest
ones demonstrate declining nest trends.
We evaluated nest trends by using nest
count data consistently collected using
a standardized approach for at least 9
years, with the last year of data in 2014
or more recently and with an average of
more than 50 nests annually. When data
did not meet these criteria, we evaluated
bar graphs provided in the Status
Review Report to consider all available
data. Thus, these data are representative
of the DPS because they include the
largest nesting aggregations. With the
possible exception of Colombia, nesting
aggregations for which data are not
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available are likely small. Significant
declines have been observed at nesting
beaches with the greatest historical or
current nesting female abundance, most
notably in Trinidad and Tobago (Grande
Riviere, Fishing Pond, and Tobago),
Suriname, French Guiana (AwalaYalimapo), Florida, and Costa Rica
(Tortuguero). Therefore, these nest
trends represent the best available data
for this DPS.
In Trinidad and Tobago, trends in
annual nest counts were largely negative
between 2009 and 2017, the years for
which data were available. For
Trinidad, we analyzed trends for three
separately monitored beaches, including
Grande Riviere, Matura, and Fishing
Pond. The long-term trend was negative
for Grande Riviere (median = ¥6.9
percent; sd = 17.4 percent; 95 percent CI
= ¥43.8 to 26.9 percent; f = 0.682; mean
annual nests = 13,272), positive for
Matura (median = 1.8 percent; sd = 15.1
percent; 95 percent CI = ¥29.2 to 33.0
percent; f = 0.561; mean annual nests =
7,359), and negative for Fishing Pond
(median = ¥19.3 percent; sd = 15.1
percent; 95 percent CI = ¥49.8 to 12.0
percent; f = 0.916; mean annual nests =
3,892). For Tobago, the median trend
was ¥0.9 percent annually (sd = 11.3
percent; 95 percent CI = ¥25.0 to 21.5
percent; f = 0.540; mean annual nests =
452).
For French Guiana, we analyzed nest
count data from 2002 to 2017 for AwalaYalimapo beach in the west and data
from 1999 to 2017 for Cayenne and
Remire Montjoly beaches in the east.
There was a steep decline at AwalaYalimapo, with a median trend of
¥19.4 percent annually (sd = 12.2
percent; 95 percent CI = ¥43.2 to 6.0
percent; f = 0.942; mean annual nests =
3,200). In contrast to Awala-Yalimapo,
nest counts at Cayenne and Remire
Montjoly increased by 2.8 percent
annually (sd = 12.9 percent; 95 percent
CI = ¥24.9 to 27.9 percent; f = 0.596;
mean annual nests = 3,498). In addition,
leatherback nesting occurred on remote
beaches in western French Guiana until
2013 (e.g., a high of 4670 nests was
found in 2003, with 1,270 mean annual
nests from 2002 to 2013), but we were
unable to analyze trends because
monitoring on these remote beaches has
been reduced since approximately 2010
due to significant beach erosion and the
disappearance of some previously
monitored beaches.
Suriname, Grenada, and Panama each
had a single time series sufficient for
trend analysis. For Suriname, we
combined datasets from two beaches,
Galibi and Braamspunt, which were
monitored between 2001 and 2017.
Total nests in Suriname declined by
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¥14.6 percent annually (sd = 9.6
percent; 95 percent CI = ¥36.4 to 4.5
percent; f = 0.953; mean annual nests =
4,586). In Grenada, data on the number
of nesting tracks were collected on
Levera beach between 2002 and 2018.
There was a 7.1 percent annual increase
in tracks at Levera during that period
(sd = 8.7 percent; 95 percent CI = ¥10.5
to 25.3 percent; f = 0.827; mean annual
tracks = 895). In Panama, the nest
counts at Chiriqui beach increased by
0.8 percent annually (sd = 7.0 percent;
95 percent CI = ¥14.1 to 14.6 percent;
f = 0.557; mean annual nests = 4,463)
between 2004 and 2017.
In Costa Rica, the four beaches for
which we had sufficient data to analyze
annual nest count trends mostly
exhibited declining trends. Tortuguero
experienced the steepest decrease, with
a median trend of ¥10.9 percent
annually (sd = 4.2 percent; 95 percent
CI = ¥19.5 to 2.2 percent) for data
collected between 1995 and 2017. Nest
counts decreased by ¥3.8 percent
annually at Pacuare beach (sd = 9.3
percent; 95 percent CI = ¥22.6 to 16.9
percent) between 2004 and 2017, but
increased by 1.8 percent annually (sd =
6.0 percent; 95 percent CI = ¥10.8 to
14.2 percent) at the nearby Pacuare
Nature Reserve between 1991 and 2017.
Nest counts at Estacion la Tortuga
deceased slightly, with a median trend
of ¥0.5 percent annually (sd = 7.0
percent; 95 percent CI = ¥15.7 to 13.1
percent) between 2002 and 2017.
For the United States, we analyzed
annual nest count trends for Florida
(statewide data collected between 2008
and 2017), three beaches in Puerto Rico,
including Culebra (1984 to 2017),
Luquillo-Fajardo (1996 to 2017), and
Maunabo (1999 to 2017), and Sandy
Point National Wildlife Refuge in St.
Croix, USVI (1982 to 2017). The median
trend for Florida was a decline of ¥2.1
percent annually (sd = 13.0 percent; 95
percent CI = ¥28.3 to 25.5 percent; f =
0.582; mean annual nests = 1,288).
Culebra nests decreased by ¥3.7
percent annually (sd = 5.3 percent; 95
percent CI = ¥14.9 to 6.8 percent; f =
0.791; mean annual nests = 153), while
nests increased by 15.9 percent annually
at Luquillo-Fajardo (sd = 5.5 percent; 95
percent CI = ¥7.1 to 15.3 percent; f =
0.805; mean annual nests = 283) and by
7.7 percent annually at Maunabo (sd =
4.9 percent; 95 percent CI = ¥2.7 to
17.4 percent; f = 0.945; mean annual
nests = 161). In St. Croix, nests
increased by 1.7 percent annually (sd =
4.6 percent; 95 percent CI = ¥7.8 to
10.7 percent; f = 0.660; mean annual
nests = 399).
These trend data are similar to other
recent findings, adding further
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confidence in declining trends at
multiple large nesting aggregations.
Because of concerns about declining
nest counts throughout the region, the
National Fish and Wildlife Foundation
(NFWF) convened a NW Atlantic
Leatherback Working Group (i.e., the
Working Group) to assess recent nesting
data and complete a region-wide trend
analysis (NW Atlantic Leatherback
Working Group 2018). The trend
analyses conducted by the Working
Group used leatherback nesting data
from 23 sites from 14 different nations
with at least 10 years of data with
consistent within-site methodology,
analyzing data for three time periods:
1990 to 2017, 1998 to 2017, and 2008 to
2017. Our approach to trend analyses
was similar to that used by the Working
Group in that both approaches involved
Bayesian analyses of data meeting set
criteria. However, the Team decided
against aggregating the data over the
DPS due to incongruity of data
collection methods, collection dates and
duration, and reporting. Despite these
differences, the overall conclusion was
the same—an overall declining nest
trend.
The Working Group found that
regional, abundance-weighted trends
were negative for all three time periods
and became more negative in the more
recent time series (NW Atlantic
Leatherback Working Group 2018).
Specifically, overall nesting trends
decreased at ¥4.21 percent annually
from 1990 to 2017 and at ¥5.37 percent
annually from 1998 to 2017, with the
most notable decrease (¥9.32 percent
annually) occurring during the most
recent time frame of 2008 to 2017.
While site-level trends showed variation
within and among sites and across the
time periods, overall the sites also
reflected the same regional pattern:
More negative trends were apparent
during the most recent time frame.
Seven sites had significant positive
nesting trends from 1990 to 2017, but no
sites exhibited significant positive
trends from 2008 to 2017. The
significant decline observed at AwalaYalimapo, French Guiana (¥12.95
percent annually from 1990 to 2017,
¥19.05 percent annually from 1998 to
2017, and ¥31.26 percent annually
from 2008 to 2017), drove the regional
results, but similar significant declines
were found at other nesting beaches for
the longer time period, including: St.
Kitts and Nevis (¥12.43 percent
annually), Tortuguero, Costa Rica
(¥10.42 percent annually), Suriname
(¥5.14 percent annually), and Culebra,
Puerto Rico (¥4.61 percent annually). It
should be noted that the other nesting
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beach in French Guiana (Cayenne)
demonstrated an increasing trend (7.44
percent annually from 1990 to 2017 and
8.19 percent annually from 1998 to
2017). However, it exhibited a
decreasing trend (¥14.21 percent
annually) from 2008 to 2017. While
nesting increased over time at Cayenne,
this increase has apparently not resulted
from females shifting from AwalaYalimapo, as turtles that nest at
Cayenne are genetically distinct
(Molfetti et al. 2013) and females tagged
in Awala-Yalimapo are not seen in
Cayenne or vice versa (NW Atlantic
Leatherback Working Group 2018).
These modeling results demonstrate
that there has been a decline in NW
Atlantic nesting from 1990 to 2017, with
the most significant decreases occurring
from 2008 to 2017. Some nesting
beaches demonstrated positive trends
for the longer time period. However,
none showed significant increases over
the most recent time period. The cause
for the decline is uncertain, but the
Working Group identified
anthropogenic sources (e.g., fisheries
bycatch), habitat losses, and changes in
life history parameters (such as
remigration interval) as potential drivers
of the regional decline. While these
results were taken into consideration by
the Team when evaluating the
extinction risk of the NW Atlantic DPS,
the Team also performed its own trend
analysis of the data provided to the
Team so that the trends were calculated
in a manner consistent with other DPSs.
Regardless, both trend analyses
conclude that the NW Atlantic DPS is
experiencing a significant decline in
nesting.
In-water abundance studies of
leatherback turtles are rare. Archibald
and James (2016) assessed the relative
abundance of turtles at a foraging area
off Nova Scotia, Canada, from 2002 to
2015. This study evaluated
opportunistic sightings per unit effort
and found a mean density of 9.8 turtles
per 100 km2, representing the highest
in-water density of leatherback turtles
reported to date. Archibald and James
(2016) concluded that the relative
abundance of foraging leatherback
turtles off Canada exhibited high interannual variability but, overall, showed a
stable trend from 2002 to 2015. The
authors reported that (at that time) these
results were consistent with the stable
or, in some cases, increasing trends
reported for contributing NW Atlantic
nesting beaches over the last decade
(Dutton et al. 2005; Girondot et al. 2007;
Fossette et al. 2008; McGowan et al.
2008; Stewart et al. 2011; Rivas et al.
2015). While there were no indications
of a decreasing trend, the results should
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be interpreted with caution because of
the small study area, opportunistic data
collection, availability bias variance,
and lack of understanding of the relative
density outside the study area
(Archibald and James 2016).
Despite the declining trend in nesting,
productivity parameters for the DPS are
similar to the species’ averages (though
some may be declining, as we discuss
below). While there is some variation,
most productivity parameters are
relatively consistent throughout the
DPS. The overall survival rate for
nesting females is relatively high at 85
percent (Pfaller et al. 2018), with mean
estimates of 0.70 to 0.99 in French
Guiana (Rivalan et al. 2005, 2008), 0.89
in St. Croix (Dutton et al. 2005), and
0.89 to 0.96 on the Atlantic coast of
Florida (Stewart et al. 2007, 2014).
Remigration intervals range from 1 to 11
years (Schulz 1975; Boulon et al. 1996;
Chevalier and Girondot 1998; Hilterman
and Goverse 2007; Eckert et al. 2012;
Stewart et al. 2014; Rivas et al. 2016;
Garner et al. 2017). In St. Croix and St.
Kitts, the median remigration interval
appears to be increasing (4.5 years; K.R.
Stewart, The Ocean Foundation and C.
Lombard, USFWS, pers. 2019; K.M.
Stewart, Ross University School of
Veterinary Medicine and St. Kitts Sea
Turtle Monitoring Network, pers.
comm., 2019). Averaging all available
data, the mean remigration interval for
the DPS is 2.7 years, rounded to 3 years
for use in our calculation of the index
of nesting female abundance. Average
clutch frequency per nesting season
ranges from 3.6 to 8.3 throughout the
region, with an overall mean of 5.5 nests
per season, interspersed with 9 to 10
day internesting intervals (Eckert et al.
2015; Garner et al. 2017). Recent records
indicate that nesting females deposit 80
to 88 eggs per clutch. However, an early
study by Carr and Ogren (1959) reported
only 67 eggs per clutch. Hatching
success is highly variable for nests that
remain in situ, even for those that are
viable and do not experience significant
inundation or predation, with estimates
as low as 8.9 percent in Costa Rica
(Troe¨ng et al. 2007) and 10.6 percent in
Suriname (Hilterman and Goverse 2007)
and as high as 93.4 percent in Florida
(Perrault et al. 2012). Overall, hatching
success is estimated at approximately 50
percent (Eckert et al. 2012). Hatchling
sex ratios often exhibit a female bias,
but less so than for other sea turtle
species, with estimated production of
anywhere from 30 to 100 percent
females in Suriname, Tobago, Colombia,
and Costa Rica (Mrosovsky et al. 1984;
Dutton et al. 1985; Godfrey et al. 1996;
Leslie et al. 1996; Mickelson and
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Downie 2010; Patin˜o-Martı´nez et al.
2012). However, the proportion of
females documented in foraging
individuals and strandings ranges from
57 to 70 percent (Murphy et al. 2006;
James et al. 2007; TEWG 2007), and the
ratio of females to males during an
individual breeding season is thought to
be closer to 1:1 (Stewart and Dutton
2014).
We conclude that the DPS exhibits a
declining nest trend. In addition, there
are indications of decreased
productivity within the DPS. In St.
Croix, one of the most thoroughly
monitored nesting beaches in this DPS,
the data from 1981 to 2010 indicate that
hatching success and clutch frequency
are declining and remigration intervals
are increasing (Garner et al. 2017).
Overall, we have a high degree of
confidence in the decreasing nest trend
and productivity metrics for this DPS,
due to the large amount of data available
from the largest nesting aggregations.
We acknowledge that data are not
available from all nesting beaches, but
the data that we have relied upon is the
best available and meets established
standards. The declining trends reflect
reduced nesting female abundance. In
addition, longer remigration intervals
and/or reduced clutch frequencies may
play a role in this decline. The decline
reflects a reduction in productivity that
places the DPS at risk given the
magnitude and duration of the
decreasing trend.
Spatial Distribution
The DPS has a broad spatial
distribution for both foraging and
nesting. There is significant genetic
population structure, with
subpopulations connected via various
levels of gene flow and metapopulation
dynamics. Tagging and telemetry
studies indicate considerable mixing of
leatherback turtles among nesting
beaches and at multiple foraging areas
throughout the North Atlantic Ocean.
Nesting is widespread throughout the
NW Atlantic beaches, occurring
primarily as scattered, small
aggregations throughout the Wider
Caribbean, but with larger
concentrations of nesting activity at
certain sites in Trinidad, French Guiana,
Suriname, Trinidad, Colombia, Panama,
Costa Rica, Puerto Rico, St. Croix, and
Florida (Horrocks et al. 2016).
Genetic sampling in the NW Atlantic
DPS has been generally extensive with
good coverage of large populations in
this region. However, sampling from
some smaller Caribbean nesting
aggregations is absent, and there are
gaps in sampling or analysis for nesting
sites along the coasts of South and
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Central America (e.g., Guyana,
Venezuela, Colombia, and Panama). A
comprehensive survey of genetic
population structure in the Atlantic
Ocean included large sample sizes from
five nesting populations representative
of the DPS and analysis of longer
mtDNA sequences in combination with
an array of 17 nuclear microsatellite
DNA loci (Roden and Dutton 2011;
Dutton et al. 2013). The microsatellite
data revealed fine-scale genetic
differentiation among neighboring
subpopulations (Dutton et al. 2013):
Trinidad, French Guiana/Suriname,
Florida, Costa Rica, and St. Croix. The
mtDNA data failed to find significant
differentiation between Florida and
Costa Rica or between Trinidad and
French Guiana/Suriname. However,
Dutton et al. (2013) show that the
mtDNA sequence variation had
relatively low statistical power to detect
fine scale structure compared to the
microsatellite DNA loci. The mtDNA
homogeneity between Costa Rica and
Florida, with differentiation
demonstrated at nuclear DNA loci,
suggests that Costa Rica may be the
source of founders for the Florida
population via one or multiple recent
colonization events, likely indicating
historic connectivity rather than
ongoing demographic connectivity
(Dutton et al. 2013). Likewise the
French Guiana/Suriname and Trinidad
populations were undifferentiated with
mtDNA likely indicating historic
connectivity. However, microsatellite
DNA reveal fine-scale genetic structure
that is consistent with tagging studies
demonstrating a lack of nesting female
movement between the two nesting
aggregations (TEWG 2007). Significant
genetic differentiation has also been
reported for Martinique and Guadeloupe
and the mainland French Guiana
rookery (Molfetti et al. 2013). St. Croix
likely represents a broader Northern
Caribbean subpopulation of the NW
Atlantic population that includes
multiple neighboring island nesting
aggregations in the USVI and Puerto
Rico. However, sampling and analysis
would be required to determine extent
of fine scale structuring (NMFS
unpublished data; Dutton et al. 2013).
The Costa Rica (Tortuguero and
Gandoca) and Guiana (French Guiana
and Suriname) nesting aggregations are
distinct subpopulations based on
microsatellite and mtDNA results
(Dutton et al. 2013), but information on
tag returns indicates movement of
nesting females between adjacent
beaches of Panama, Colombia,
Venezuela and Guyana. Therefore, these
nesting aggregations have ‘‘fuzzy’’
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48341
boundaries, likely a result of flexible
natal homing. Nesting females use
beaches up to 400 km apart between
nesting seasons (Troe¨ng et al. 2004;
Chaco´n-Chaverri and Eckert 2007) and
up to 463 km apart within the same
nesting season (Stewart et al. 2014).
Additional sampling of the remaining
nesting sites will be required to
determine the extent of fine-scale
structuring within the NW Atlantic DPS.
However, the available science indicates
significant substructure within the DPS.
Tagging studies indicate individual
movement and gene flow among nesting
aggregations. This is facilitated by the
species’ flexible natal homing, i.e.,
philopatry to a region, rather than a
specific beach. In adjacent nesting sites
in French Guiana and Suriname, five to
six percent of nesting females were
observed to shift from one site to the
other within a season (TEWG 2007),
while Schulz (1971) reported this
proportion to be slightly higher at 8.5
percent. In contrast, 35 percent of
nesting females in Gandoca, Costa Rica,
were estimated to nest at sites other
than the study site during an individual
season (Chaco´n-Chaverri and Eckert
2007). The predisposition of nesting
females to stray within a nesting season
may be influenced by the proximity of
alternative nesting sites within a range
of approximately 200 km (Horrocks et
al. 2016). However, even within a given
nesting season, females have been
observed to move as far as 369 km
(Grenada), 463.5 km (Florida), and 532
km (Dominica) from their original
location (Horrocks et al. 2016). Among
nesting seasons, interchange between
nesting locations also appears to be
frequent and wide-ranging, with
maximum distance separating two
nesting sites for an individual female
recorded as 1,849 km over an 8-year
span (Horrocks et al. 2016).
Genetic studies have revealed that
turtles from different nesting
aggregations use the same foraging
areas. Analyzing 684 longline bycatch
samples from across the NW Atlantic in
a mixed stock analysis and
microsatellite assignment, Stewart et al.
(2016) found that leatherback turtles
from Costa Rica were caught in a higher
proportion in the GOM (43 percent)
compared to the Northeast Distant
fishing zone, an area in the
northwestern Atlantic Ocean (6
percent), while turtles from Trinidad
and French Guiana comprised 54
percent of bycatch in the GOM and 93
percent in the Northeast Distant fishing
zone. A study of turtles foraging off
Nova Scotia, Canada, similarly assigned
most (82 percent) of the 288 sampled
turtles to Trinidad (n = 164) and French
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Guiana (n = 72), with 15 percent (n =
44) from Costa Rica, and the remainder
from St. Croix (n = 7) and Florida (n =
1; Stewart et al. 2013). These
proportions generally represent the
relative population sizes for these
breeding populations. Microsatellite
DNA assignment of wild captured or
stranded males (n = 122) throughout the
NW Atlantic and Mediterranean found
that all males originated from NW
Atlantic nesting aggregations (Trinidad:
55 percent, French Guiana: 31 percent,
and Costa Rica: 14 percent; Roden et al.
2017). No turtles were identified from
St. Croix or Florida. One turtle that
stranded in Turkey was assigned to
French Guiana, while strandings in
France were assigned to Trinidad or
French Guiana (Roden et al. 2017).
The mixing of nesting aggregations at
foraging areas is also supported by
several tagging and/or satellite telemetry
projects, conducted in U.S. waters
(Murphy et al. 2006; LPRC 2014; Dodge
et al. 2014, 2015; Aleksa et al. 2018),
Canada (James et al. 2005a, 2005b,
2005c, 2006b, 2007; Bond and James
2017), Atlantic Europe and
Mediterranean (Doyle et al. 2008;
Sonmez et al. 2008), and on nesting
beaches of various nations (Hildebrand
1987; Hays et al. 2004; Ferraroli et al.
2004; Eckert 2006; Eckert et al. 2006;
Hays et al. 2006; TEWG 2007; Sonmez
et al. 2008; Evans et al. 2008; Fossette
et al. 2010a, 2010b; Richardson et al.
2012; Bailey et al. 2012; Stewart et al.
2014; Fossette et al. 2014; Horrocks et
al. 2016; Chambault et al. 2017). For
instance, turtles from Nova Scotian
foraging grounds were tracked to nesting
areas off Colombia, Trinidad, Guyana,
and French Guiana (Bond and James
2017). The reverse has also been
demonstrated: some leatherback turtles
from the western Atlantic undertake
annual migrations to Canadian waters to
forage (James et al. 2005c), exemplified
by post-nesting adults tracked to the
waters off Nova Scotia from a variety of
nesting locations, including French
Guiana and Trinidad (Fossette et al.
2014), Costa Rica, Panama (Evans et al.
2008), and Anguilla (Richardson et al.
2012). The eastern and western GOM
also provide foraging areas for this DPS
(Aleksa et al. 2018), as observed from
tracks of post-nesting turtles from
Florida (Hildebrand 1987), Costa Rica
(Tortuguero, Gandoca), and Panama
(Chiriquı´ Beach; Evans et al. 2008;
Evans et al. 2012). Evans et al. (2008)
suggested that the GOM may represent
a significant foraging ground for
leatherback turtles from the Caribbean
coast of Central America.
High use foraging areas may be
identified through available telemetry
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data, but the migration routes to those
areas may vary. Ferraroli et al. (2004)
tracked leatherback turtles from French
Guiana and found turtles dispersed
widely throughout the North Atlantic
but mostly followed two dispersion
patterns: (1) Moving north to the Gulf
Stream area, where they started
following the general ocean circulation;
and (2) traveling east, swimming mostly
against the North Equatorial Current.
Fossette et al. (2014) found a relatively
broad migratory corridor when turtles
departed their nesting sites in French
Guiana/Suriname, and their movements
overlapped with turtles from Grenada
and Trinidad. Fossette et al. (2010a,
2010b) found that turtles tracked from
nesting beaches in French Guiana,
Suriname, and Grenada and turtles
caught in waters off Nova Scotia and
Ireland displayed three distinct
migration strategies: (1) Heading
northwest to fertile foraging areas off the
Gulf of Maine, Canada, and GOM; (2)
crossing the North Atlantic Ocean to
areas off western Europe and Africa; and
(3) residing between northern and
equatorial waters. Essentially, tagging
data coupled with satellite telemetry
data indicate that leatherback turtles of
the NW Atlantic DPS use the entire
North Atlantic Ocean for foraging and
migration (TEWG 2007).
Although adults forage at multiple
areas throughout the North Atlantic
Ocean (Fossette et al. 2014), the range of
juvenile leatherback turtles may be more
restricted. Using an active movement
model, Lalire and Gaspar (2019) found
that most juveniles originating from
nesting beaches in French Guiana and
Suriname cross the Atlantic Ocean at
mid-latitudes with north-south seasonal
migrations; after several years, they
reach the coasts of Europe and North
Africa. Eckert (2002) reviewed the
records of nearly 100 sightings of
juvenile (less than 100 cm curved
carapace length (CCL)) leatherback
turtles and determined they are
generally found in waters warmer than
26 °C, suggesting that the first portion of
their life is spent in tropical and
subtropical waters. After exceeding 100
cm CCL, distribution extends into cooler
waters (as low as 8 °C), which is
considered to be the primary habitat for
the species (Eckert 2002).
The wide distribution of nesting and
foraging areas likely buffers the DPS
against local catastrophes or
environmental changes. The fine-scale
population structure, with movement of
individuals and genes among nesting
aggregations, indicates that the DPS has
the capacity to withstand other
catastrophic events.
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Diversity
The NW Atlantic DPS exhibits spatial
diversity, as demonstrated by insular
and continental nesting, multiple
diverse foraging areas, and moderate
genetic diversity. The DPS nests along
both continental and insular coastlines.
Nesting beach habitat also shows
considerable diversity, ranging from
coarse-grained, sandy beaches to silty,
ephemeral shorelines whose dynamics
are influenced by estuarine input. The
breadth and, in some cases, transiency,
of suitable nesting habitat in the western
North Atlantic may contribute to
consistent, low-level flexibility in natal
homing, both within and among
reproductive seasons (Bra¨utigam and
Eckert 2006), and this flexibility is
thought to surpass that of other sea
turtle species (TEWG 2007).
This DPS exhibits some temporal
variation in nesting. Nesting generally
begins in March or April, peaks in May
or June, and ends in July or August
(Eckert et al. 2012). In French Guiana,
a second small nesting peak was
documented in Awala-Yalimapo during
December and January. However, the
number of nests deposited during that
time frame decreased from 700 in 1986/
1987 to 40 in 1992/1993, and now only
a small number of individuals are
observed to nest during that time
(Girondot et al. 2007). Some evidence
indicates that the timing of nesting may
be modulated by environmental
characteristics distant from the nesting
beach, such as water temperatures at
foraging grounds (Neeman et al. 2015).
The foraging strategies are also
diverse, with turtles using coastal and
pelagic waters throughout the entire
North Atlantic Ocean (Fossette et al.
2014). Foraging habitats include
temperate waters of the GOM, North
Central Atlantic Ocean, northwestern
shelf (United States and Canada),
southeastern U.S. coast, the
Mediterranean Sea, and northeastern
shelf (Europe; TEWG 2007). Some postnesting females also remain in tropical
waters (Fossette et al. 2010). Overall,
leatherback turtles in the North Atlantic
Ocean appear to have a diverse array of
foraging habitat available.
Genetic diversity of the DPS is
moderate, with six mtDNA haplotypes
(Dutton et al. 2013). In St. Croix, a
unique haplotype occurs at high
frequency. The Florida and Costa Rica
nesting aggregations each possess one
unique, low frequency haplotype.
Based upon this information, we
conclude that nesting location and
habitat are diverse, providing some level
of resilience against short-term spatial
and temporal changes in the
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environment. However, high-abundance
nesting occurs only at a few locations
(e.g., Trinidad, French Guiana, and
Panama). The foraging diversity likely
provides resilience against local
reductions in prey availability or
catastrophic events, such as oil spills,
by limiting exposure to a limited
proportion of the total population.
Moderate genetic diversity may provide
the DPS with the raw material necessary
for adapting to long-term environmental
changes, such as cyclic or directional
changes in ocean environments due to
natural and human causes (McElhany et
al. 2000; NMFS 2017). We conclude that
such diversity provides some level of
resilience to threats for this DPS.
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Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
Destruction and modification of
leatherback turtle nesting habitat results
from a variety of activities including
coastal development and construction;
beach erosion and inundation;
placement of erosion control and
nearshore shoreline stabilization
structures and other barriers to nesting;
beachfront lighting; vehicular and
pedestrian traffic; beach sand
placement; sand extraction; removal of
native vegetation; and planting of nonnative vegetation (Lutcavage et al. 1997;
Bouchard et al. 1998; USFWS 1999;
Dow et al. 2007; Eckert et al. 2012;
NMFS and USFWS 2013). As a result,
most nesting beaches are severely
degraded by such activities that
continue to cause adverse impacts
throughout the range of the DPS.
Coastal Development and Construction
In many areas, nesting habitat is
under constant threat from coastal
development and construction (Dow et
al. 2007; Crespo and Diez 2016; Flores
and Diez 2016). Coastal development
impacts include construction of
buildings and pilings on the beach;
increased erosion; artificial lighting;
pollution; recreational beach equipment
and other obstacles on the beach; beach
driving; increased human disturbance;
and mechanized beach cleaning
(Lutcavage et al. 1997; USFWS 1999;
Hernandez et al. 2007; Dow et al. 2007;
Trinidad and Tobago Forestry Division
et al. 2010; Flores and Diez 2016).
Driftwood found on nesting beaches
also has the potential to alter nesting
beach habitat and obstruct nesting
females and hatchlings, as seen in
Gandoca, Costa Rica (Chaco´n-Chaverri
and Eckert 2007). These threats impact
nesting habitat by reducing the amount
and quality of suitable beaches,
preventing or deterring nesting females
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from using optimal locations, destroying
nests, eggs, and hatchlings, and
preventing hatchlings from successfully
reaching the ocean (USFWS 1999;
Chaco´n-Chaverri and Eckert 2007;
Hernandez et al. 2007; Witherington et
al. 2014). Development involving the
construction of tall buildings and
clearing of vegetation can also alter sand
temperatures and skew sex ratios
(Gledhill 2007).
Development occurs to varying
extents throughout the range of the DPS,
but most leatherback nesting occurs in
proximity to some coastal development.
The Florida shoreline is extensively
developed outside wildlife refuges
(Witherington et al. 2011). In Grenada,
nearly 20 percent of all nests surveyed
from 2001 to 2005 occurred in an area
affected by development, resulting in
ongoing run-off onto nesting beaches
(Maison et al. 2010). In Trinidad,
increasing rural and commercial
beachfront development is a concern,
especially on the east coast where the
main nesting beaches are located
(Trinidad and Tobago Forestry Division
et al. 2010), including Grande Riviere,
the largest nesting aggregation of this
DPS. Likewise, several Tobago beaches
are densely developed for commercial
tourism, resulting in reduced turtle
access to potential nesting sites due to
buildings, umbrellas, and other
recreational equipment (Trinidad and
Tobago Forestry Division et al. 2010).
Development in Puerto Rico, in
particular Playa Grande-El Paraiso (i.e.,
Dorado Beach, which is considered to
be the most important nesting beach in
Puerto Rico), is also a notable concern
(Crespo and Diez 2016; Flores and Diez
2016). There, ecosystems continue to be
threatened by coastal development,
even though the coastal zone is
protected by the Maritime-Terrestrial
Zone designation (i.e., Coastal Public
Trust Lands; Flores and Diez 2016).
Coastal development likely influences
leatherback nest placement and
subsequent nest success, which is the
percentage of nesting attempts (i.e.,
emergences onto the beach) that result
in eggs being deposited. On Margarita
Island, Venezuela, Hernandez et al.
(2007) found that leatherback nesting
aggregated towards the portions of the
beach with fewer risk factors, such as
light pollution and concentrations of
beach furniture. This change in nesting
behavior resulted in females nesting in
less optimum areas (e.g., areas with
lower hatching success), thus affecting
the reproductive potential of
leatherback turtles in this region.
The magnitude of development is also
changing in some areas, where nest
placement and success may be affected
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in the future. For instance, the area
around Cayenne, French Guiana, is
undergoing increased urbanization and
recreational use (Fossette et al. 2008). In
recent years, nesting has increased at
Cayenne and eastern beaches compared
to the western Awala-Yalimapo beaches
(Re´serve Naturelle de l’Amana data in
Berzins 2018 and KWATA data in
Berzins 2018). As such, more nesting in
French Guiana is exposed to coastal
development and the associated threats,
and these threats are likely to continue
and increase.
Beach Erosion and Inundation
While erosion is often intensified due
to anthropogenic influences, natural
features in some areas result in high
erosion rates and unstable beaches, thus
affecting leatherback nesting. For
instance, the Maroni River influence in
the Guianas (French Guiana especially)
has resulted in highly dynamic and
unstable beaches, with shifting mudflats
making nesting habitat unsuitable
(Crossland 2003; Goverse and Hilterman
2003; Fossette et al. 2008). Beaches are
often created and lost along the coast of
French Guiana (Kelle et al. 2007). For
example, remote beaches in western
French Guiana experience significant
beach erosion and several disappeared,
reducing or preventing monitoring (and
likely nesting). In Suriname,
Braamspunt Beach at the mouth of the
Suriname River is moving west, out of
the established Wia Wia Nature Reserve
and may disappear in the next several
years (M. Hiwat, WWF, pers. comm.,
2018). This is significant in that
Braamspunt is currently the main
nesting beach in Suriname. The second
highest nesting area in Suriname, Galibi
Beach, is also experiencing significant
erosion and becoming narrower. Similar
beach erosion is occurring in Guyana, as
well as in Trinidad and Tobago
(Reichart et al. 2003; Trinidad and
Tobago Forestry Division et al. 2010). At
some Trinidad and Tobago nesting sites
(e.g., Fishing Pond, Matura, Grande
Riviere, and Great Courland Bay), rivers
emerge onto nesting beaches and create
additional erosion during the nesting
season (Godley et al. 1993; Lee Lum
2005), intensifying nest loss (up to 35
percent of nests; Trinidad and Tobago
Forestry Division et al. 2010).
Seasonal erosion also occurs at most
Caribbean nesting beaches. A survey of
Wider Caribbean Regions found that
erosion/accretion was the highest threat
to nesting habitat (Dow et al. 2007). For
example, at Playa Gandoca, Costa Rica,
erosion from strong coastal drift
currents is thought to be one of the
largest obstacles to hatching success,
destroying greater than 10 percent of all
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nests laid in some years (Chaco´nChaverri and Eckert 2007). In 2006 and
2007, coastal erosion and inundation
accounted for 33 to 42 percent of nest
loss in southern Panama and 29 to 48
percent on Caribbean Colombia beaches
(Patin˜o-Martı´nez et al. 2008).
Inundation of nests is also a concern.
Leatherback turtles generally nest closer
to the water than other sea turtles (Caut
et al. 2010). If nests are laid too close
to the high tide line, they are subjected
to erosion and inundation, which can
result in egg mortality from suffocation
or curtailed embryonic development
(Chaco´n-Chaverri and Eckert 2007; Caut
et al. 2010). This inundation
phenomenon occurs on multiple nesting
beaches and is particularly of concern in
areas with high tidal influence and
dynamic coastlines. On Krofajapasi
beach in Suriname, 31.6 percent of nests
laid by females were below the spring
high tide level and determined to be
‘‘doomed’’ clutches (Dutton and
Whitmore 1983). Similarly, in Gandoca,
Costa Rica, 37 percent of nests from
1990 to 2004 were laid in the low tide
zone and would have been inundated if
not relocated (Chaco´n-Chaverri and
Eckert 2007). In St. Croix, 43 percent of
the nests (with a range of 25 to 68
percent) were considered to be
‘‘doomed’’ each season (McDonaldDutton et al. 2001), but beginning in
1983, all doomed clutches were
relocated to improve hatching success
(Dutton et al. 2005). Without
intervention, these nests would likely
have been lost. On Awala-Yalimapo,
French Guiana, 27 of 89 nests were
overlapped by tide at least once during
the incubation period, and the hatching
success was on average significantly
lower in overwashed nests (Caut et al.
2010). Observed mortality was 100
percent in the intertidal zone at sites
along the coasts of Panama and
Colombia, with an overall nest loss by
erosion and inundation ranging from 16
to 48 percent among three major nesting
sites (Patin˜o-Martı´nez et al. 2008).
While levels of inundation and resulting
declines in hatching success have been
noted at multiple sites throughout the
range of the NW Atlantic DPS, the
specific impacts of inundation may be
variable. Hilterman and Goverse (2007)
noted that leatherback nests can tolerate
relatively high levels of inundation, so
hatching may still be successful despite
proximity to the tide line. Because of
this, and because it may affect natural
sex ratios (Mrosovsky and Yntema
1980), the relocation of nests susceptible
to inundation was abandoned in 2002 in
Suriname (Hilterman and Goverse
2007); only nests directly threatened by
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beach erosion are relocated, under
certain circumstances. Other nations
still relocate nests to reduce the impacts
of erosion. However, as mentioned, such
practices may result in cooler nests and
affect sex ratios (Spanier 2008). While
eggs relocated to hatcheries could have
been lost under natural circumstances,
due to coastal erosion and inundation in
some areas (Dutton and Whitmore 1983,
Chaco´n-Chaverri and Eckert 2007),
hatching success in relocated nests is
often lower than in situ nests (Revuelta
et al. 2014; Valentin-Gamazo et al. 2018;
Florida Department of Environmental
Protection unpublished data 2018).
Such naturally dynamic areas make it
difficult to protect nesting beach habitat
and accurately assess leatherback
nesting trends. This is particularly
noteworthy given that nesting females
use high energy, erosion-prone beaches,
which often result in high nest loss
(Chaco´n-Chaverri and Eckert 2007;
TEWG 2007; Spanier 2008; Trinidad
and Tobago Forestry Division et al.
2010). However, leatherback turtles in
the Guianas seem to have adapted to
this constant geomorphological change
of beaches. When new beaches develop,
they may be colonized within months
by nesting females, who take advantage
of the fresh, clean sand (or seashells, in
Guyana) and absence of entangling or
deep-rooted beach vegetation (TEWG
2007).
Nest site selection by leatherback
turtles is still poorly understood
(Maison et al. 2010), but nesting females
may be changing their nesting patterns
due to erosion. Spanier (2008) found
that nesting females at Playa Gandoca,
Costa Rica, appear to actively select nest
sites that are not undergoing extensive
erosion, with slope considered to be the
cue for site selection. A similar result
was found on Grande Riviere, Trinidad,
with a nesting shift from east to west
throughout the season as an apparent
response to erosion on the eastern end
of the nesting beach (Lee Lum 2005).
Further, Maison et al. (2010) studied
nest placement in Grenada and
discovered that leatherback turtles
seemed to respond to the accretion of
the north facing beach and erosion of
the east facing beach in 2005 by nesting
more often on the north facing beach. If
erosion is increasing in existing nesting
locations, nesting may occur in areas
with lower success rates, thus affecting
productivity. In addition, leatherback
nests are deeper than those of other sea
turtles; water content and salinity
typically increase with depth, leading to
a decrease in sea turtle hatching success
(Foley et al. 2006).
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Erosion Control, Nearshore Shoreline
Stabilization Structures, and Other
Barriers
A widespread strategy to reduce
coastal erosion is to construct erosion
control structures. However, these
structures reduce the amount of
available nesting habitat. Also, when
beachfront development occurs, the site
is often engineered to protect the
property from erosion. This type of
shoreline engineering, collectively
referred to as beach armoring, includes
sea walls, rock revetments, riprap,
sandbag installations, groins and jetties.
Beach armoring can result in permanent
loss of a nesting beach through
accelerated erosion and prevention of
natural beach/dune accretion. These
impacts can prevent or hamper nesting
females from accessing suitable nesting
sites (USFWS 1999). Clutches deposited
seaward of these structures may be
inundated at high tide or washed out
entirely by increased wave action near
the base of the erosion control
structures. As these structures fail and
break apart, they spread debris on the
beach, thus creating additional impacts
to hatchlings and nesting females.
In the southeastern United States,
numerous erosion control structures
that create barriers to nesting have been
constructed. In Florida, the total amount
of existing and potential future armoring
along the coastline is approximately 24
percent (164 miles; FDEP, pers. comm.,
2018). This assessment of armoring does
not include other structures that are a
barrier to sea turtle nesting, such as
dune crossovers, cabanas, sand fences,
and recreational equipment.
Additionally, jetties have been placed at
many ocean inlets in the United States
to keep transported sand from closing
the inlet channel. The installation of
jetties resulted in lower loggerhead and
green turtle nesting density updrift and
downdrift of the inlets, leading
researchers to propose that beach
instability from both erosion and
accretion may discourage turtle nesting
(Witherington et al. 2005). Leatherback
nesting near jetties and inlets is low,
possibly reflecting their avoidance of
such areas. There are some efforts, such
as the Coastal Construction Control Line
Program, that provide protection for
Florida’s beaches and dunes while
allowing for continued use of private
property. However, armoring structures
on and adjacent to the nesting beach
continue to be permitted and
constructed on the nesting beaches of
Florida, as in other nations where the
DPS nests.
Due to erosion, beach nourishment is
a frequent activity in some developed
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areas, and many beaches are on a
periodic nourishment schedule. Beach
nourishment may result in direct burial
and disturbance to nesting females, if
conducted during the nesting season. It
may also result in changes in sand
density, beach hardness, beach moisture
content, beach slope, sand color, sand
grain size, sand grain shape, and sand
grain mineral content, if the placed sand
is dissimilar from the original beach
sand (Nelson and Dickerson 1988;
USFWS 1999). These changes can affect
nest site selection, digging behavior,
incubation temperature (and hence sex
ratios), gas exchange parameters within
incubating nests, hydric environment of
the nest, hatching success and hatchling
emerging success (Lutcavage et al. 1997;
Steinitz et al. 1998; Ernest and Martin
1999; USFWS 1999; Rumbold et al.
2001; Brock et al. 2009). On severely
eroded sections of beach, where little or
no suitable nesting habitat previously
existed, beach nourishment has been
found to result in increased nesting
(Ernest and Martin 1999). However, on
most beaches in the southeastern United
States, nesting success typically
declines for the first year or two
following nourishment, even though
more nesting habitat is available for
turtles (Trindell et al. 1998; Ernest and
Martin 1999; Herren 1999; Brock et al.
2009). Further, nourishment projects
result in heavy machinery, pipelines,
increased human activity and artificial
lighting on the project beach, further
affecting nesting females and beach
habitat. Overall, the impacts of beach
nourishment to this DPS are not as
widespread as other threats to nesting
habitat, as Dow et al. (2007) found that
only four nations (Anguilla, Cuba,
Mexico, and United States) reported
frequent or occasional beach
nourishment.
Artificial Lighting
Coastal development also contributes
to habitat degradation by increasing
light pollution, which can result in
hatchling and nesting female
disorientation, altering behavior and
leading to mortality. In Florida, from
2013 to 2017, a total of 341 leatherback
nests (representing the whole or
majority of hatchlings in the nest) and
five nesting females were disoriented
(FWC unpublished data 2018). Artificial
lighting ranked as the third highest
threat to nesting/hatching turtles in the
Wider Caribbean Region (Dow et al.
2007). For example, urban development
is significant in Puerto Rico, with light
pollution (as well as coastal erosion and
deforestation) occurring near
leatherback nesting beaches (Crespo and
Diez 2016). Fortunately, some of the
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major nesting beaches in this DPS are
located in comparatively remote areas,
and large-scale development is currently
less of an issue there (Trinidad and
Tobago Forestry Division et al. 2010;
NMFS and USFWS 2013). That said,
even within the same country, light
pollution is variable. Fossette et al.
(2008) reported that in French Guiana,
light pollution from residential areas is
a problem at Cayenne Beach, but it is
not an issue at Awala-Yalimapo.
Similarly, lighting is not a significant
problem on nesting beaches in Trinidad,
but is a concern in Tobago (Trinidad
and Tobago Forestry Division et al.
2010). With the risk of increased
development in some of these relatively
remote areas, additional light pollution
is anticipated, and disorientation of
hatchlings and adults from such lighting
may become a bigger problem. In Costa
Rica, beachfront lighting is increasing
and may become problematic at
Gandoca Beach (Chaco´n-Chaverri and
Eckert 2007) and Tortuguero (de Haro
and Troe¨ng 2006).
Light pollution has been managed to
some extent (Witherington et al. 2014).
Lighting in Florida is regulated by
multiple rules and regulations including
Florida statutes, the Florida Building
Code, and local lighting ordinances
(Witherington et al. 2014). In addition,
the Florida Department of
Transportation and local governments
have adopted lighting-design standards.
A total of 82 municipalities in Florida
have adopted lighting ordinances to
minimize the impact of lighting on
adjacent sea turtle nesting beaches
(Witherington et al. 2014). However,
compliance and enforcement is lacking
in some areas. Further, lighting away
from areas covered by beachfront
ordinances is unregulated, resulting in
urban glow. Although outreach and
conservation programs control the
impacts of lighting in some other
locations, such as Costa Rica, Mexico,
and Puerto Rico (Lutcavage et al. 1997;
Crespo and Diez 2016), a majority of
nations do not have regulations in place.
Sand Extraction
Extracting sand from nesting beaches
for construction projects has a
detrimental effect on the amount of
available nesting beach habitat and also
accelerates erosion (resulting in the
aforementioned associated impacts).
Sand mining occurs in most Wider
Caribbean nations to varying extent and
frequency (Dow et al. 2007). In
particular, beach sand mining has been
extensive at Matura Bay and
Blanchisseuse in Trinidad (Trinidad
and Tobago Forestry Division et al.
2010). Some nations regulate sand
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48345
mining: In St. Lucia, the Conservation
and Management Act of 2014 requires a
certificate of environmental approval for
projects removing sand from nesting
beaches.
Removal of Native Vegetation
In some nations, upland deforestation
and the resultant deposition of debris
and garbage can destroy or modify
nesting beaches. The debris can block
access of gravid (pregnant) females and
fatally trap emergent hatchlings
(Chaco´n-Chaverri and Eckert 2007). The
accumulation of logs reduces the
amount of available nesting habitat,
possibly forcing leatherback females to
nest in suboptimal locations (TEWG
2007). Deforestation due to coastal
development is a notable concern in
Puerto Rico (Crespo and Diez 2016).
Vehicular Traffic
Beach driving also occurs in most
nations throughout the range of this DPS
(Chaco´n-Chaverri and Eckert 2007; Dow
et al. 2007; Trinidad and Tobago
Forestry Division et al. 2010). In the
United States, vehicular driving is
allowed on certain beaches in Florida
(e.g., Duval, St. Johns, and Volusia
Counties). Beach driving reduces the
quality of nesting habitat in several
ways. Vehicle ruts on the beach can
prevent or impede hatchlings from
reaching the ocean following emergence
from the nest (Mann 1977; Hosier et al.
1981; Cox et al. 1994; Hughes and Caine
1994). Sand compaction by vehicles
hinders nest construction and hatchling
emergence from nests (Mann 1977;
Gledhill 2007). Vehicle lights and
vehicle movement on the beach after
dark can deter females from nesting and
disorient hatchlings. Additionally,
vehicle traffic contributes to erosion,
especially during high tides or on
narrow beaches where driving is
concentrated on the high beach and
foredune.
Vegetation
Beach vegetation (native and nonnative) can affect turtle nesting
productivity by obstructing nest
construction and potentially drying the
sand (resulting in egg chamber
collapse). Vegetation can form
impenetrable root mats that can invade
and desiccate eggs and affect developing
embryos, impede hatchling emergence,
and trap hatchlings (Conrad et al. 2011).
Non-native vegetation has invaded
many coastal areas and often
outcompetes native plant species
(USFWS 1999). The occurrence of exotic
vegetation (or loss of native vegetation)
was recognized as a medium-ranked
threat in many Wider Caribbean nations
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(Dow et al. 2007). The Australian pine
(Casuarina equisetifolia) is particularly
harmful to sea turtles (USFWS 1999).
Australian pines cause excessive
shading of the beach that would not
otherwise occur. Studies of loggerhead
turtles in Florida suggest that nests laid
in shaded areas are subjected to lower
incubation temperatures, which may
alter the natural hatchling sex ratio
(Marcus and Maley 1987; Schmelz and
Mezich 1988). Fallen Australian pines
limit access to suitable nest sites and
can entrap nesting females (Reardon and
Mansfield 1997). The shallow root
network of these pines can interfere
with nest construction (Schmelz and
Mezich 1988). Dense stands of
Australian pine have overtaken many
coastal areas throughout central and
south Florida.
While non-native vegetation can affect
nesting habitat throughout the range of
the DPS, native vegetation can also
affect productivity. For instance, at
Sandy Point, St. Croix, changing
erosion-accretion cycles led to native
Ipomoea pes-caprae, a creeping vine,
extending into the nesting area in some
years. Nesting females at Sandy Point
typically avoided nesting in vegetation,
resulting in more nests laid near the
high-tide line (Conrad et al. 2011). As a
result, Ipomoea pes-caprae decreased
nest productivity by reducing
leatherback hatching and emergence
(percentage of hatchlings that emerge
from the nest) success rates (Conrad et
al. 2011).
Mitigations to Habitat Modification
Nesting habitat disruptions are
minimized in some areas. Several areas
in the NW Atlantic DPS range are under
U.S. Federal ownership as National
Wildlife Refuges in Florida (Archie Carr
and Hobe Sound), Puerto Rico (Culebra
and Vieques) and St. Croix (Sandy
Point). Beaches in some Wider
Caribbean countries are also protected.
In Trinidad, Matura and Fishing Pond
beaches were declared Prohibited Areas
in 1990, and the nesting beach at
Grande Riviere in 1997. In 1998, the
Amana Nature Reserve, which includes
Awala-Yalimapo beach and a 30 m wide
marine fringe, was established in French
Guiana. In Suriname, the Wia Wia
Nature Reserve was implemented in
1961 (amended and enlarged in 1966 to
protect sea turtles), and in 1969, the
Marowijne beaches were declared a
sanctuary (the Galibi Nature Reserve;
Schulz 1971). In addition, Tortuguero
National Park, Costa Rica, was
established in 1976 to protect nesting
habitat (Bjorndal et al., 1999).
Terrestrial habitat in these areas is
therefore protected from the above
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threats to some extent. USFWS and
NMFS also designated as critical habitat
for leatherback turtles the nesting
beaches at Sandy Point, St. Croix (43 FR
43688; September 26, 1978) and
surrounding marine waters (44 FR
17710; March 23, 1979), which benefits
the turtles in this DPS. However, if ESA
protections did not continue (i.e., if this
species were no longer listed), these
protections would be lost.
Marine Habitat Modifications
In the marine environment, habitat
threats include anthropogenic noise and
offshore lighting. We discuss other
threats to marine habitat and prey (e.g.,
marine pollution, oil exploration, and
climate change) in later sections.
Anthropogenic noise impacts the
marine habitat of the DPS. Dow Piniak
et al. (2012) measured hearing
sensitivity of leatherback hatchlings.
They found that hatchlings are able to
detect sounds underwater and in air,
responding to stimuli between 50 and
1200 Hz in water and 50 and 1600 Hz
in air, with maximum sensitivity
between 100 and 400 Hz in water and
50 and 400 Hz in air. This sensitivity
range overlaps with the frequencies and
levels produced by many anthropogenic
sources used in the North Atlantic,
including seismic airgun arrays,
drilling, low frequency sonar, shipping,
pile driving, and operating wind
turbines. These noise sources may affect
leatherback turtles’ marine habitat and
subsequently impact distribution and
behavior. Offshore artificial lighting
occurs in some marine waters of this
DPS (Dow et al. 2007) but is less of a
threat than beachfront lighting
throughout the range of the DPS.
Summary
We conclude that nesting females,
hatchlings, and eggs are exposed to the
loss and modification of nesting habitat,
especially as a result of coastal
development and armoring, erosion, and
artificial lighting. These threats impact
the DPS by reducing nesting and
hatching success, thus, lowering the
productivity of the DPS. Based on the
information presented above, we
conclude that habitat reduction and
modification pose a threat to the NW
Atlantic DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization is a threat to the NW
Atlantic DPS, mostly due to poaching of
turtles and eggs in certain nations. Legal
harvest of turtles and eggs also occurs in
some nations.
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While the vast majority of nations
within the range of the NW Atlantic
DPS protect leatherback turtles from
harvest, it is legal in some Caribbean
and Central American nations
(Brautigam and Eckert 2006; Dow et al.
2007; Richardson et al. 2013; Horrocks
et al. 2016). For example, the harvest of
leatherback turtles over 20 pounds is
allowed in Montserrat and Dominica
from October 1 to May 31; Saint Lucia
allows leatherback turtles over 65
pounds to be taken from October 2 to
February 27; and St. Kitts and Nevis
allows take of leatherback turtles over
350 pounds from October 2 to February
27 (Montserrat Turtles Act 2002;
Bra¨utigam and Eckert 2006). In some
nations, commercial use is prohibited,
but traditional use is allowed, which
can still diminish protection. In
Colombia, subsistence fishing of sea
turtles is permitted, and indigenous use
is allowed in Honduras. Traditional or
cultural use is permitted in Belize with
prior approval (Bra¨utigam and Eckert
2006). However, regular leatherback
nesting does not occur in Belize, and its
occurrence in surrounding waters is
infrequent, reducing the impact of such
mortality. Legal harvest throughout the
range of this DPS is not monitored, and
the precise magnitude of this threat is
not clear. However, we conclude that
legal harvest of turtles is significant
because, when it occurs, nesting turtles
are targeted, removing the most
important individuals from the
population. More often, leatherback
eggs, rather than turtle meat, are
harvested (TEWG 2007; Patin˜o-Martı´nez
et al. 2008), reducing productivity in the
DPS.
Poaching of turtles and eggs occurs
throughout the NW Atlantic DPS, and
Dow et al. (2007) ranked it as a threat
for all turtle species on the beaches in
the Wider Caribbean Region. In Panama,
interviews with locals revealed that the
development of a new way for cooking
leatherback turtle meat has resulted in
a recent increase of its consumption in
Changuinola, Bocas del Toro Province
(CITES Secretariat 2019). Adult turtles
are killed in Panama and on remote
beaches in Trinidad and Tobago (Troe¨ng
et al. 2002; Ordon˜ez et al. 2007;
Trinidad and Tobago Forestry Division
et al. 2010). Most poaching, however,
targets eggs, and the level often is
determined by how much monitoring
and activity to deter poachers occur on
the nesting beaches. Some of the highest
levels of egg poaching occur throughout
Costa Rica (Troe¨ng et al. 2004). Troe¨ng
et al. (2007) found that, at a minimum,
between 13 to 21.5 percent of nests
between 2000 and 2005 were illegally
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collected at Tortuguero. Poaching of
leatherback nests was higher outside
Tortuguero National Park (minimum 33
percent) than within the National Park
(minimum 9 percent) in 2005 (de Haro
and Troe¨ng 2006). At Pacuare Playa,
Costa Rica, 55 percent of nests were
poached in 2012 (Fonseca and Chaco´n
2012) and 42 percent were poached in
2017, which was the lowest level since
Latin American Sea Turtles (LAST)
started to monitor in 2012 (LAST 2017).
Poaching at Gandoca Beach has
decreased over time (previously 100
percent of nests were poached), but
rates still averaged 15.5 percent
annually from 1990 to 2004 (Chaco´nChaverri and Eckert 2007). In the
Dominican Republic, poaching is also
high. Revuelta et al. (2012) determined
the poaching of clutches in Jaragua
National Park and Saona Island ranged
from 0 to 100 percent from 2006 to
2010, with averages of 19 percent on
western Jaragua National Park beaches,
71 percent on eastern Jaragua National
Park beaches, and 74 percent on Saona.
Poaching also occurs at relatively high
levels in Colombia (e.g., 22 to 31 percent
of clutches at Playona in 2006 and 2007;
Patin˜o-Martı´nez et al. 2008) and, to
some extent, in most other Caribbean
nations (e.g., Guyana and Grenada).
Poaching is likely more prevalent, and
occurs at higher levels, on unmonitored
or unprotected beaches (Dow et al.
2007; TEWG 2007; Troe¨ng et al. 2007;
Trinidad and Tobago Forestry Division
et al. 2010; K. Charles, Oceans Spirits
Inc., pers. comm., 2018).
Poaching has been significantly
reduced at some nesting beaches. In
Suriname, high levels of egg poaching
(at least 26 percent of nests) occurred in
the late 1990s, but due to better
monitoring and enforcement, that level
has been significantly reduced
(Hilterman and Goverse 2007; M. Hiwat,
WWF, pers. comm., 2018). Poaching
was also a major problem in Trinidad,
but levels have been reduced with more
people monitoring the beach (Trinidad
and Tobago Forestry Division et al.
2010). The Marine Turtle Conservation
Act of 2004 (MTCA) funds activities in
Panama in an attempt to reduce
poaching. At Chiriqui Beach, Panama,
intense monitoring efforts have
attempted to reduce poaching. However,
of the monitored nests, 29 leatherback
nests (0.7 percent) were still poached in
2017 (Sea Turtle Conservancy 2017).
Further, poaching in Panama outside
the monitored areas still occurs, with
the clandestine sale of eggs widespread
(Brautigam and Eckert 2006). In St.
Croix, almost 100 percent of nests were
lost to poaching prior to 1981 (Garner et
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al. 2017). However, the establishment of
the USFWS Sandy Point National
Wildlife Refuge has reduced egg
poaching to 0 to 1.8 percent annually as
a result of nightly patrols (Garner et al.
2017).
Poaching of eggs is widespread
throughout the Caribbean, especially on
beaches of Costa Rica, Dominican
Republic, and Colombia. The total
number of individuals affected by
poaching cannot be quantified at this
time. However, we conclude that many
eggs and some adults are affected by
illegal poaching at nesting beaches.
Adults and eggs are also exposed to
legal harvest in some nations. The legal
and illegal harvest of nesting females
reduces both abundance (through loss of
nesting females) and productivity
(through loss of reproductive potential),
resulting in a high impact to the DPS.
Legal and illegal egg harvest reduces
productivity only. Thus, we conclude
that overutilization poses a threat to the
DPS.
Disease or Predation
For the NW Atlantic DPS, information
on diseases is limited, but predation is
a well-documented threat.
Much of the available information on
disease in leatherback turtles was
obtained by necropsy of stranded large
juvenile and adult turtles; the health
implications of various conditions
reported in this species are
incompletely understood. Solitary large
intestinal diverticulitis of unknown
etiology was found in 31 subadult and
adult leatherback turtles stranded in
U.S. waters (Stacy et al. 2015). All
lesions were chronic and unrelated to
the cause of death in all cases, although
risk of perforation and other
complications are possible. Adrenal
gland protozoal parasites were found in
17 leatherback turtles in North
American waters examined from 2001 to
2014; it is not currently known whether
parasitism affects adrenal function
(Ferguson et al. 2016). In addition,
leatherback turtles are hosts for several
trematode parasites (flatworms), known
species of which also occur in hardshelled sea turtles (Manfredi et al. 1996,
Greiner et al. 2013). In general,
trematodes are frequently encountered
without any apparent clinical effect on
the turtle host but can affect some
heavily parasitized individuals. With
regard to other types of potential
disease-causing organisms, there are a
small number of reports of bacterial
infections in stranded individuals
(Poppi et al. 2012; Donnelly et al. 2016).
A variety of other bacteria have been
documented in nesting females on
beaches in Costa Rica (Santoro et al.
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2008) and St. Kitts (Dutton et al. 2013);
the majority of identified bacterial
species may be considered as potential
or opportunistic pathogens for sea
turtles. A putative case of
fibropapilloma, a virus-associated
tumor-causing disease in sea turtles, has
been reported in a leatherback; this
disease is considered very rare in the
species (Huerta et al. 2002).
An in-water health assessment was
performed on 12 turtles directly caught
at-sea and seven turtles bycaught in
fishing gear in the NW Atlantic Ocean
(Innis et al. 2010). Most were
determined to be in good health, but
several exhibited evidence of past
injuries. The blood chemistry of
entangled turtles indicated stress,
seawater intake, and reduced food
consumption associated with
entanglement. In addition, Perrault et al.
(2012) examined baseline blood
chemistry metrics (i.e., plasma protein
electrophoresis, hematology, and
plasma biochemistry) as indicators of
health for nesting females in Florida.
They found that multiple measures of
maternal health significantly correlated
with leatherback hatching and
emergence success (the percentage of
hatchlings that emerge from the nest).
From these data, we estimate that the
exposure of eggs, juveniles, and adults
to disease is low. The impact of disease
cannot be quantified at this time as we
have no documentation of any deaths or
reductions in productivity directly
related to disease. However, disease
may compound the effects of or have
synergistic effects with other threats to
the species and related physiologic
derangements. We conclude that
disease, alone or in combination with
other threats, is likely a threat to the
DPS.
Throughout the range of the DPS,
predation is a threat to leatherback eggs,
hatchlings, and adults. Eckert et al.
(2012) provides an exhaustive list of the
documented predators for each life stage
and area. For eggs in the NW Atlantic
DPS, predators include ants (Dorylus
spininodis), fly larvae (Diptera spp.),
locust larvae (Acrididae spp.), mole
crickets (Scapteriscus didactylus), ghost
crabs (Ocypode quadratus), vultures
(Cathartidae), dogs (Canis familiaris),
cattle (Bos taurus; due to trampling),
armadillo (Dasypodidae), opossum
(Didelphis marsupialis), coati (Nasua
spp.), and raccoons (Procyon lotor); see
Eckert et al. 2012).
In particular, dog predation of eggs
occurs in many areas (e.g., Colombia,
French Guiana, Guyana, Panama, Puerto
Rico, and Trinidad and Tobago). In
Trinidad, where the largest nesting
aggregation occurs, feral dogs are
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considered to be the primary threat to
eggs, even above poaching and coastal
erosion (Trinidad and Tobago Forestry
Division et al. 2010). On Chiriqui Beach,
Panama, 54 percent of the monitored
leatherback nests were depredated by
dogs in 2003 and approximately eight
percent in 2004 (Ordon˜ez et al. 2007).
Such predation may been reduced as a
result of protection efforts funded by the
MTCA. In Playa California, Maunabo,
Puerto Rico, more than 30 percent of the
leatherback nests were depredated by
stray dogs in 2012 (Crespo and Diez
2016). A public outreach project in
Puerto Rico was established in 2013 to
reduce this impact. Puerto Rico is a U.S.
territory; if ESA protections were
removed, it is likely that predation rates
would be higher.
Egg predation by other species is also
a notable concern in some areas. On
Gandoca Beach, Costa Rica, dipteran
larvae infestation exceeded 75 percent
of nests in 2005 and 2006 (Gautreau et
al. 2008). In French Guiana, on average,
mole crickets preyed on 18 percent of
all eggs (Maros et al. 2003). These
threats are likely to continue, as no
predator screening typically occurs in
Wider Caribbean nations due to the
potential for increased poaching as well
as logistical difficulties in these areas of
high density nesting. Nest loss to
predators was found to be the seventh
ranked threat to turtles (all species, not
specific to leatherback turtles) on
nesting beaches in the Wider Caribbean
Region, and have been noted to
frequently occur in Honduras, Mexico,
Panama, Puerto Rico, and Venezuela
(Dow et al. 2007).
Hatchlings are preyed upon by a wide
variety of species, including mole
crickets, ghost crabs, horse-eye jack fish
(Caranx latus), gray snapper (Lutjanus
griseus), tarpon (Megalops atlanticus),
vultures, hawks (Accipitridae), gulls
(Larus spp.), night heron (Nyctanassa
violacea), frigate birds (Fregatidae),
dogs, mongoose (Atilax paludinosus),
coati, and raccoons (Eckert et al. 2012).
Again, dogs are a serious threat to
leatherback hatchlings in some areas,
and especially in Puerto Rico (Crespo
and Diez 2016).
There are few documented predators
to subadults and adult leatherback
turtles, presumably because of their
large size and pelagic behavior.
Predation by sharks (Elasmobranchii)
and killer whales (Orcinus orca) has
been reported in Barbados and St.
Vincent, respectively (Caldwell and
Caldwell 1969; Horrocks 1989). Sharks
have also been reported to prey on
nesting females off St. Croix, USVI
(DeLand 2017; Scarfo et al. 2019). Over
the past 6 years, researchers at Sandy
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Point have observed an apparent
increase in injuries to leatherback
turtles (K. Stewart, NMFS, pers. comm.,
2019). These injuries, many of them
consistent with shark predation, affect
up to 70 percent of all nesting females
at the beach (Scarfo et al. 2019). While
some turtles probably survive these
encounters, it is unknown how many
encounters result in mortality or
reduced nesting effort. Jaguars (Panthera
onca) prey on nesting females in some
areas, including Suriname, French
Guiana, Guyana, and Costa Rica (see
Eckert et al. 2012). While three nesting
females were killed by jaguars at
Tortuguero, Costa Rica, from 1998 to
2005, this mortality is only considered
to be a minor threat and is therefore
unlikely to cause a population decline
on its own (Troe¨ng et al. 2007).
Archibald and James (2018) examined
228 leatherback turtles for injuries off
Atlantic Canada and on Matura,
Trinidad, and found 15.7 percent of
turtles exhibited injuries of suspected
predatory origin.
Predation on early life stages is
natural; however, at high rates, it
reduces the viability of the DPS (see the
Status Review). Predation primarily
reduces productivity via reduced egg
and hatching success and the loss of
hatchlings. Predation on nesting females
reduces abundance and productivity.
We conclude that predation is a threat
to the NW Atlantic DPS.
Inadequacy of Existing Regulatory
Mechanisms
Many regulatory mechanisms
(including state, Federal and
international) have been promulgated to
protect leatherback turtles, eggs, and
nesting habitat throughout the range of
the NW Atlantic DPS. We reviewed the
objectives of each regulation and to
what extent they adequately address the
targeted threat (i.e., the threat that the
regulation was intended to address).
The effectiveness of many international
regulations was evaluated by Hykle
(2002), who found that international
instruments often do not realize their
full potential, either because they do not
include all key countries, do not
specifically address sea turtle
conservation, are handicapped by the
lack of a sovereign authority that
promotes enforcement, or are not legally
binding.
National regulatory mechanisms are
described in full in the Status Review
Report. Although these regulatory
mechanisms provide some protection to
the species, most inadequately reduce
the threat they were designed to
address, generally as a result of poor
implementation or incomplete
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enforcement. Specifically, existing
regulatory mechanisms continue to be
inadequate to control impacts to nesting
beach habitat and overutilization
(harvest of turtles and eggs) for this DPS.
In addition, regulatory mechanisms are
inadequate to reduce several other
threats including bycatch in fishing
gear, vessel strikes, and marine debris.
Despite existing regulatory mechanisms,
bycatch from fisheries (discussed in
detail along with existing regulatory
mechanisms in the Fisheries Bycatch
section), incomplete nesting habitat
protection, and poaching remain major
threats to the DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat
to the NW Atlantic DPS. Bycatch occurs
throughout the range of the DPS,
affecting juveniles, subadults, and
adults.
Finkbeiner et al. (2011) analyzed sea
turtle bycatch across all commercial
U.S. fisheries from 1990 to 2007. They
examined sea turtle bycatch reduction
based on the year a particular fishery
implemented bycatch reduction
measures. Prior to implementing
bycatch reduction measures,
approximately 3,800 leatherback
interactions, of which 2,300 were lethal,
occurred in U.S. Atlantic Ocean and
GOM commercial fisheries annually.
After bycatch reduction measures were
implemented, 1,400 leatherback turtles,
40 of those dead, were estimated to be
taken annually in the Atlantic Ocean.
The Atlantic/GOM pelagic longline
fishery was responsible for the most
annual interactions (n = 900) and
mortality events (n = 17) in the Atlantic
Ocean, followed by the southeast
Atlantic/GOM shrimp trawl fishery
(Finkbeiner et al. 2011). These estimates
represent minimum numbers of actual
bycatch and mortality. Because the
observer coverage for these fisheries is
low (so some bycatch may not be
observed and observed effort may not be
a true representation of actual fleet
effort), not all fisheries are observed and
thus some are not included in these
estimates. Interactions are difficult to
observe if gear modifications are in
place, and so the methods used are
conservative (Finkbeiner et al. 2011).
In the Wider Caribbean Region,
reports of leatherback bycatch in
fisheries are common. In a survey of
Caribbean nations, Dow et al. (2007)
ranked fisheries bycatch among the
highest in-water threat to sea turtles.
Many fisheries in less industrialized
nations are coastal and small-scale, but
these fisheries are reported to have
significant ecological impacts due to
their high bycatch discards and impacts
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to the marine environment (Shester and
Micheli 2011). Of particular concern are
leatherback bycatch in artisanal
nearshore and offshore gillnet, longline
and trawl fisheries (Barrios-Garrido and
Montiel-Villalobos 2016). Information
on fisheries bycatch is collected mostly
from stranding records but also from
fisher surveys (Moncada et al. 2003;
Delamare 2005; Madarie 2006, 2010,
2012) and observations of nesting
females. Hilterman and Goverse (2007)
recorded fisheries related injuries on
nesting females in Suriname. In 2002,
16.9 percent of the nesting females had
fisheries- related injuries; in 2003, at
least 18.3 percent had such injuries; and
in 2005, 9 percent (Hilterman and
Goverse 2007). From 2000 to 2003, an
average of 28 leatherback turtles
stranded on the Suriname survey
beaches. Although no cause of death
was immediately apparent, Hilterman
and Goverse (2007) indicated that the
mortalities were fisheries-related, based
upon the fisheries that occur offshore
with high bycatch and documented
fisheries-related injuries on nesting
leatherback turtles at the same time. On
the western oceanic nesting beaches of
French Guiana, injuries consistent with
fisheries interactions (e.g., scars,
wounds) were recorded on 8.4 percent
(n = 1,259) of nesting females in 2003
(Morisson et al. 2003). In Venezuela, 55
percent of strandings from 2001 to 2007
(n = 57) exhibited evidence of fisheries
interactions (Barrios-Garrido and
Montiel-Villalobos 2016). Most recently,
an injury assessment of 228 leatherback
turtles from two foraging areas off the
Atlantic coast of Canada and Trinidad
nesting beaches found 19 percent of
turtles exhibited injuries indicative of
entanglement in lines or nets, and 17
percent showed evidence of hooks; 62
percent of turtles assessed exhibited a
minimum of one external injury
(Archibald and James 2018).
Fisheries bycatch also occur in the
Mediterranean and eastern North
Atlantic Ocean. Casale et al. (2003)
analyzed 411 records of leatherback
turtles in the Mediterranean, of which
152 were collected from Italy. Most of
these records were from fishery captures
(n = 170) or found in unknown
circumstances (n = 127). Of those
reported by fishermen, set or drift nets
had the highest number of interactions
(29.4 percent), followed by unknown
fishing equipment (22.9 percent),
longlines (20.6 percent), unspecified
nets (12.9 percent), other fishing
equipment (9.4 percent), and trawls (4.7
percent). The main fisheries affecting
turtles in the Mediterranean (all turtle
species, not just leatherback turtles) are
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Spanish and Italian surface longlines,
North Adriatic Italian trawls, Tunisian
trawls, Turkish trawls, Moroccan
driftnets, and Italian driftnets (Camin˜as
2004). The same types of fishing gear
from other nations also affect turtles, but
the bycatch numbers are lower (Camin˜as
2004). Stranding records from Portugal
from 1978 to 2013 found that 49 of 275
leatherback turtles exhibited evidence of
fishery interactions (the cause of
stranding could not be determined in
most cases due to decomposition state;
Nicolau et al. 2016). Multifilament nets
accounted for approximately 41 percent
of the strandings, followed by
monofilament nets, traps/pots, and
longlines. Coastal artisanal fisheries
were recognized as a particular threat in
Portugal.
Based upon these summary reports
and stranding assessments, it is clear
that fisheries have a large impact on the
NW Atlantic DPS. In the following
paragraphs, we review information on
specific gear interactions, including the
following fisheries: Gillnet, longline,
trawl, pot/trap, and other.
Gillnet Fisheries
Gillnet fisheries are common
throughout the range of this DPS. Due
to the nature of the gear and fishing
practices (e.g., relatively long soak
times), bycatch in gillnets is among the
highest source of direct sea turtle
mortality (Upite et al. 2013; Wallace et
al. 2013; Upite et al. 2018). Upite et al.
(2018) evaluated observed fishery
interactions and post-interaction
mortality and determined a 79 percent
sea turtle mortality rate for Northeast
and Mid-Atlantic gillnet gear from 2011
to 2015. Wallace et al. (2013) calculated
leatherback bycatch in gillnets
throughout the NW Atlantic Ocean of
0.015 turtles/set, with a 21 percent
median mortality rate (not considering
post-interaction mortality). This gear
was classified as having a relatively
high bycatch impact on the NW Atlantic
leatherback population. Small scale
fisheries are of particular concern, given
the magnitude of bycatch, nearshore
distribution, and limited monitoring
(Lewison et al. 2015). When nets are
used in waters off nesting beaches,
where leatherback turtles mate, nesting
females and mature males are often
captured and killed.
The largest documented bycatch of
leatherback turtles in gillnet gear occurs
off the coast of Trinidad. Lee Lum
(2006) estimated that more than 3,000
leatherback turtles were captured by
coastal surface gillnets off Trinidad
annually, with an approximate 30
percent mortality rate. These captures
involved adult turtles, occurring off the
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north and east coasts of Trinidad during
January to August, i.e., the breeding and
nesting season, when nesting females
and adult males occur in the waters off
nesting beaches (Lee Lum 2006). Gilman
et al. (2010) extrapolated leatherback
bycatch estimates (Lee Lum 2006;
Gearhart and Eckert 2007) to the entire
Trinidad Spanish mackerel and king
mackerel surface gillnet fishery, and
estimated that almost 7,000 turtles were
captured in 2000. Additionally, Eckert
et al. (2013) worked with drift gillnet
fishermen to identify leatherback
bycatch hot spots off the north and east
coasts of Trinidad (where the nesting
beaches are), with capture probability
increasing from March to July and a
secondary peak in October.
Whereas most of the documented
leatherback bycatch off Trinidad occurs
in surface drift gillnet fisheries, bottom
set gillnet fishing also captures
leatherback turtles (Gass 2006; S. Eckert,
WIDECAST, pers. comm., 2018). The
magnitude of effort and turtle bycatch in
this fishery are lower than for surface
nets, but mortality rates are higher
(approximately 70 percent; Gass 2006).
As such, the bottom set gillnet fishery
is thought to have a comparable level of
mortality to the drift gillnet fishery
(approximately 500 to 1,000 leatherback
turtles annually; Gass 2006; S. Eckert,
WIDECAST, pers. comm., 2018). The
Sea Turtle Recovery Action Plan for the
Republic of Trinidad and Tobago noted
that drowning in gillnets is that nation’s
most significant cause of sea turtle
mortality (Trinidad and Tobago Forestry
Division et al. 2010). Bond and James
(2017) tracked a female from Canadian
waters to a nesting beach off Trinidad,
but the turtle was confirmed dead,
entangled in coastal fishing gear, just
prior to the date of her first predicted
nesting event. Venezuelan fishers have
also been seen hauling leatherback
turtles from Trinidad waters into their
boats (Brautigam and Eckert 2006).
Together, drift and bottom-set gillnets
off the Trinidad beaches, which host the
largest nesting aggregation in the DPS,
are estimated to kill well over 1,000
leatherback turtles annually, and they
thus pose a large threat to the DPS.
High levels of gillnet bycatch occur in
other Caribbean and South American
nations, also off major nesting beaches.
In French Guiana, bycatch was
confirmed to be high in the Maroni
estuary (Chevalier 2001; Girondot 2015).
In 2003, 26 leatherback turtles were
caught in coastal gillnets and released
off the Cayenne and Montjoly nesting
sites (Gratiot et al. 2003 in TEWG 2007).
Delamare (2005) conducted fishermen
interviews and estimated an average of
1,149 leatherback captures in 2004 and
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2005 by bottom-set or drifting gillnets in
French Guiana. No estimate of mortality
was provided, but it is likely similar to
Trinidad fisheries, i.e., 70 and 30
percent, respectively. In Suriname, a
World Wildlife Fund survey of
fishermen estimated leatherback
bycatch in drifting gillnets at 584 in
2006, 174 in 2010, and 424 in 2012
(Madarie 2006; Madarie 2010; Madarie
2012). Most of the turtles were captured
alive. In Colombia, 10 to 40 leatherback
turtles are killed annually by gillnets
(Patin˜o-Martı´nez et al. 2008). Longline
and driftnet gillnet fisheries in
Moroccan waters off the northwestern
Africa coast capture approximately 100
leatherback turtles annually
(Benhardouze et al. 2012).
Although not at as high a rate as in
the Caribbean (based upon observed
interactions), gillnet bycatch occurs in
U.S. and Canadian waters. Although
South Carolina, Georgia, Florida,
Louisiana, and Texas have prohibited
gillnets in their State waters, active
gillnet fisheries remain in other states
and U.S. Federal waters. No cumulative
estimates of leatherback bycatch in
gillnet fisheries in U.S. waters are
available due to the limited observed
interactions. However, from 2003 to
2017, fishery observers recorded lethal
and non-lethal bycatch in fixed sink,
drift sink, and drift floating gillnets
throughout the U.S. Atlantic Exclusive
Economic Zone (EEZ) and GOM (NMFS
unpublished data). From 2012 to 2016,
27 leatherback turtles (coefficient of
variation = 0.71, 95 percent CI over all
years: 0–68) were bycaught with 21
mortalities in sink gillnet gear in the
Georges Bank and Mid-Atlantic regions
(Murray 2018). From 1989 to 1998, U.S.
drift pelagic gillnets captured 54
leatherback turtles, but that gear is no
longer used. Hamelin et al. (2017)
reviewed leatherback entanglement
records reported by Canada in Atlantic
Canadian waters between 1998 and
2014. Gillnets, mainly targeting
groundfish, were involved in 24 of 205
entanglements (11.7 percent),
particularly in Newfoundland and
Labrador (n = 15). Often, gillnet
entanglements involve the vertical lines
associated with gear (M. James, DFO,
pers. comm., 2019).
Gillnet bycatch occurs in the eastern
North Atlantic Ocean and in the
Mediterranean Sea. As in other areas,
sea turtles have the potential to interact
with set gillnets and drift gillnets. The
United Nations (UN) established a
worldwide moratorium on drift gillnet
fishing effective in 1992; the General
Fisheries Commission for the
Mediterranean prohibited driftnet
fishing in 1997; a total ban on driftnet
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fishing by the European Union fleet in
the Mediterranean went into effect in
2002; and the International Commission
for the Conservation of Atlantic Tunas
(ICCAT) banned driftnets in 2003.
Nevertheless, unregulated driftnetting
continued to occur in some areas (e.g.,
the Mediterranean Sea and off Europe;
Pierpoint 2000; Camin˜as 2004). In the
Atlantic Ocean, leatherback bycatch has
been reported from NE Atlantic tuna
driftnet fisheries by English, French and
Irish vessels (Pierpoint 2000). Of 20
leatherback turtles found in nets in
British and Irish waters (1980 to 2000),
eight were caught in the NE Atlantic
tuna driftnet fishery (with 25 percent
mortality) and one was caught in a hake
gillnet (Pierpoint 2000).
Historically, driftnet fishing in the
Mediterranean Sea caught large
numbers of sea turtles. And today an
estimated 600 illegal driftnet vessels
operate in the Mediterranean, including
fleets based in Algeria, France, Italy,
Morocco, and Turkey (Environmental
Justice Foundation 2007). Out of 411
records of leatherback turtles (stranded,
captured, sighted, or found in unknown
circumstances) in the Mediterranean
Sea, 170 turtles were captured by
fishermen, of which 29.4 percent were
caught by set or drift nets (Casale et al.
2003). Driftnets and gillnets in Greece,
Israel, Italy, Tunisia and Turkey have
reported documented leatherback
interactions, and occasional leatherback
bycatch occurs in Croatian artisanal
gillnet fisheries (Camin˜as 2004; Ergene
and Ukar 2017). In particular, Karaa et
al. (2013) reviewed 36 leatherback
bycatch records from Tunisia fisheries
in the Gulf of Gabes, and found that
gillnets are the dominant threat to
leatherback turtles in the region. A
similar result (e.g., gillnets being a high
threat to leatherback turtles in the area)
was found in the Adriatic Sea (Lazar et
al. 2012). The first leatherback recorded
on the Aegean coast of Turkey was
caught in a gillnet (Taskavak et al.
1998). Further, a review by Casale
(2008) found that leatherback turtles are
taken in the drift gillnet fishery in Spain
at a rate of 0.065 turtles/day-boat.
Throughout the range of the NW
Atlantic DPS, effective gillnet bycatch
reduction measures have not been
required, but measures to reduce
leatherback bycatch have been
discussed in some areas (e.g., Trinidad;
Eckert 2013). If nations have a closed
season for fishing, at least in the nesting
season (e.g., Suriname; Madarie 2006),
nesting females are afforded some level
of protection from gillnet bycatch. Some
nations have prohibited gillnet gear; St.
Barthelemy does not allow trammel nets
in its territorial waters and St. Lucia
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prohibits fishing within 100 meters of
shore to protect nesting turtles. There
are gillnet and trammel net restrictions
in Curacao (Ministry of Health,
Environment, and Nature 2014, UN
Environment Programme 2017). In the
United States, gillnets with stretched
mesh seven inches and larger are
prohibited at certain times off North
Carolina and Virginia to protect sea
turtles (50 CFR 223.206(d)(8); 71 FR
24776, April 26, 2006). While no gear
modifications are currently required for
U.S. gillnet fisheries, Federal U.S.
fisheries are subject to section 7 of the
ESA, 16 U.S.C. 1536(a)(2), and through
formal consultations on specific
fisheries, measures may be required to
minimize the impact of incidental take
in gillnets (NMFS 2013). Regardless of
some of these protective measures,
gillnet bycatch (especially off nesting
beaches) results in the loss of thousands
of mature individuals annually.
Longline Fisheries
Leatherback turtles are known to
interact with longline fishing gear, most
commonly pelagic longlines (Lewison et
al. 2004; Zollett 2009; Wallace et al.
2010; Wallace et al. 2013). There is
significant concern over the effects of
pelagic longline fishing, which extends
globally throughout temperate and
tropical waters, including several high
pressure fishing areas in the North
Atlantic Ocean (Fossette et al. 2014;
Gray and Diaz 2017). In international
waters, numerous flag states have high
seas longline fisheries that frequently
catch leatherback turtles (Lewison et al.
2004). Individuals are found entangled
and hooked in this gear, mostly by the
flippers (Witzell and Cramer 1995;
Coelho et al. 2015; Huang 2015).
Leatherback bycatch in longlines
throughout the NW Atlantic Ocean was
calculated at 0.062 turtles per set,
classifying the gear as a relatively low
bycatch impact relative to other sea
turtle populations (Wallace et al. 2013;
Lewison et al. 2015). However, because
longline fisheries are widespread across
leatherbacks’ distribution and use
millions of hooks each year, they pose
a large threat to the NW Atlantic DPS
and are estimated to kill thousands of
individuals (mature and immature)
annually.
Pelagic longline fishing is widespread
throughout the range of the DPS and
involves a number of nations, so an
accurate estimate of total bycatch is
difficult to obtain. In the Atlantic Ocean
from 2002 to 2013, the largest longline
fishing fleets belonged to Taiwan, Japan,
Spain, Belize, and China, with the
Taiwanese fleet comprising the largest
distant-water longline effort throughout
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the region (Angel 2014; Huang 2015). In
an assessment of the impact of ICCAT
fisheries on sea turtles, Gray and Diaz
(2017) estimated leatherback
interactions with pelagic longlines in
the ICCAT area from 2012 to 2014 (15
to 16 fleets). Using a combination of
published and assigned sea turtle
bycatch rates as a function of estimated
fishing effort submitted to ICCAT by its
members, Gray and Diaz (2017) found a
high degree of overlap in the central
North Atlantic Ocean and equatorial
waters (some of which are outside this
DPS). Within the NW Atlantic region, an
estimated 7,138 leatherback interactions
occurred in 2012, 6,036 in 2013 and
4,991 in 2014 (Gray and Diaz 2017).
Applying a reasonable estimated
mortality rate of 21.4 percent, as seen in
other high seas pelagic longline gear
(Huang 2015), results in an average
annual estimated mortality of 1,296
leatherback turtles from 2012 to 2014.
However, this is likely an underestimate
of total mortality, as the high seas
mortality rate in Huang (2015) was
based upon the disposition of the turtle
when boarded and therefore did not
account for post-interaction mortality;
240 of 459 leatherback turtles caught
from 2002 to 2013 were alive and 121
were of unknown status (Huang 2015).
Angel et al. (2014) conducted a risk
assessment of turtles from the impacts
of tuna fishing in the ICCAT region and
found the NW Atlantic RMU (which is
comparable to the NW Atlantic DPS;
Wallace et al. 2010) has high-moderate
vulnerability to longline gear, with as
many as 270 million longline hooks
annually from 2000 to 2009. In
particular, Fossette et al. (2014)
analyzed leatherback satellite tracks
(converted to densities) overlaid with
longline fishing effort from 1995 to 2009
in the Atlantic Ocean. In the North
Atlantic Ocean, a total of four seasonal
high-susceptibility areas were
identified: one in the central northern
Atlantic in international waters, one
along the east coast of the United States,
and one each in the Canary and Cape
Verdean basins (Fossette et al. 2014).
These areas partly occurred in the EEZs
of eight nations (Cape Verde, Gambia,
Guinea Bissau, Mauritania, Senegal,
Spain/Canaries, United States, and
Western Sahara). Given the species’
flexible diving behavior, it is reasonable
to expect that turtles are likely to
encounter pelagic longlines throughout
the Atlantic Ocean, regardless of
whether they are engaged in foraging or
migratory behavior (Fossette et al.
2014).
Bycatch in U.S. Atlantic and GOM
pelagic longlines has been extensively
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studied in the last decade. Current
estimates of leatherback interactions
with the U.S. Atlantic pelagic longline
fishery are lower than previous years. In
the late 1990s and early 2000s, estimates
of Atlantic U.S. pelagic longline bycatch
were around 1000 leatherback turtles
annually (NMFS 2001; Yeung 2001;
NMFS 2018), with bycatch rates of
about 0.15 to 0.5 turtles per 1000 hooks
(Watson et al. 2005). In 2005, after the
United States required pelagic longline
gear modifications (50 CFR 635.21), the
fleet was estimated to have interacted
with 351 leatherback turtles outside
experimental fishing operations (Walsh
and Garrison 2006). NMFS (2018)
estimated 239 leatherback interactions
in the U.S. Atlantic pelagic longline
fishery in 2011, 596 in 2012, 363 in
2013, 268 in 2014, 299 in 2015, and 339
in 2016. The majority of interactions
occurred in the GOM, Mid-Atlantic
Bight, Northeast Coastal, and Northeast
Distant areas (NMFS 2018). The postinteraction mortality estimate for the
most recently available 3-year period
(2013 to 2015) for leatherback turtles is
30.13 percent (L. Desfosse, NMFS, pers.
comm., 2018). Based on the average
leatherback interaction estimate for the
entire U.S. pelagic longline fleet from
2011 to 2016 (351), the estimated
average annual mortality for the U.S.
pelagic longline fishery is 106
leatherback turtles.
Leatherback interactions also occur in
Canadian pelagic longline fisheries.
From summer to fall, primarily on the
Scotian Shelf, encounters with
leatherback turtles have been
documented in the large pelagic
longline fishery since 2001 (DFO 2012).
With observer coverage ranging from 5
to 30 percent since 2001, there were 102
reported interactions with pelagic
longlines from 2001 to 2005, and 36
from 2006 to 2010 (DFO 2012).
Mortality rates are estimated to be in the
range of 21 to 49 percent, resulting in
an estimated mortality of 13 to 44
leatherback turtles annually. Based on
an analysis of Canadian observer data
from 2002 to 2010, the bycatch rate in
this fishery is estimated to have
declined from 120–190 leatherback
turtles annually from 2002 to 2006 to
60–90 leatherback turtles annually from
2006 to 2010, largely as a result of gear
modifications (Hanke et al. 2012).
In the Mediterranean Sea, longlining
is prevalent. Drifting longlines targeting
swordfish (Xiphias gladius), albacore
(Thunnus alalunga), and bluefin tuna
(T. thynnus) are considered to be the
most dangerous fishing gear for turtles
in the Mediterranean Sea (Lucchetti and
Sala 2010). Drifting longlines (mainly
for albacore tuna) in Spain, Italy,
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Greece, and Albania have documented
leatherback interactions (Camin˜as 2004).
In the western Mediterranean, swordfish
longlines appeared to be responsible for
most of the leatherback bycatch and
entanglements (Camin˜as 1998; Camin˜as
2004). Casale et al. (2003) reviewed
bycatch rates for longline fisheries
targeting swordfish and estimated the
average Mediterranean longline bycatch
rates at 0.0025 leatherback turtles/1000
hooks, with a maximum rate of 0.0510
leatherback turtles/1000 hooks in the
Tyrrhenian Sea of Italy (Casale et al.
2003; Casale and Margaritoulis 2010). Of
170 leatherback fishery captures in
fisheries from the Mediterranean Sea,
approximately 35 involved longlines
(Casale et al. 2003). While leatherback
turtles are encountered in
Mediterranean longlines, loggerheads
are the most common species caught;
only 0.1 percent of turtles captured
during an observer program in Spain,
Italy and Greece were leatherback
turtles (3 out of 2,370 observed turtles;
Laurent et al. 2001). However, given the
extensive longline effort in the
Mediterranean Sea (Casale 2008),
leatherback bycatch in the
Mediterranean is still a concern.
Lewison et al. (2004) estimated a range
of 250 to 10,000 leatherback turtles
bycaught in the Mediterranean in 2000,
with 6 percent observer coverage.
Longline bycatch of leatherback
turtles in the range of the NW Atlantic
DPS also occurs in waters off Cape
Verde (Melo and Melo 2013; Coelho et
al. 2015), Morocco (Benhardouze et al.
2012), and Brazil (Pacheco et al. 2011).
Given the wide distribution of both
pelagic longline gear and leatherback
turtles, bycatch of individuals in
longline gear can occur wherever and
whenever the gear and sea turtle
distribution overlap.
Large circle hooks (non-offset) have
been found to reduce leatherback
bycatch by as much as 55 percent
compared to traditional J-style hooks
(Andraka et al. 2013; Coelho et al.
2015). While the vessels of certain
nations may employ large circle hooks,
there are no obligations for international
longline fleets to adopt such bycatch
mitigation measures (Richardson et al.
2013). In 2005, an ICCAT resolution
encouraged circle hook research (ICCAT
2005), but no legally binding measure to
require circle hooks exists (Gilman
2011). Without the widespread use of
non-offset circle hooks, it is likely that
the high bycatch rates of leatherback
turtles in pelagic longline gear will
continue throughout the North Atlantic
high seas fisheries.
Since 2004, the United States has
issued regulations that require
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modifications to pelagic longline gear in
the U.S. Atlantic and GOM to reduce the
bycatch and post-interaction mortality
of sea turtles; these regulations (50 CFR
635.21(c)(2)) specify hook type and size
(18/0 or 16/0 circle hooks depending on
the area), bait type, use of turtle
disentangling equipment and handling
guidelines. Swimmer et al. (2017)
recently analyzed pelagic longline
interactions before (1992 to 2001) and
after (mid-2004 to 2015) these
regulations were promulgated.
Throughout the study period, 844
leatherback turtles were captured.
Overall, turtle bycatch was highest in
the Northeast Distant statistical
reporting area (0.3 turtles/1000 hooks),
followed by the Northeast Coastal,
GOM, and Caribbean areas. Bycatch
rates were higher for years prior to 2004;
after the regulations, Atlantic
leatherback bycatch rates declined by 40
percent (0.13 to 0.078 turtles/1000
hooks). Within the Northeast Distant
area alone, where additional restrictions
include a large circle hook (18/0) and
limited use of squid bait, rates declined
by 64 percent (0.44 to 0.16 turtles/1000
hooks; Swimmer et al. 2017). Gilman
and Huang (2017) found similar results:
Fish versus squid bait lowered catch
rates of leatherback turtles, and wider
circle hooks reduced leatherback catch
rates relative to narrower J and tuna
hooks. Capture probabilities are lowest
when using a combination of circle
hook and fish bait.
Efforts have been made to reduce
interactions in Canadian waters as well.
Circle hook use has been recommended
in the swordfish-directed Canadian
longline fleet since 2003, whereas
corrodible circle hooks have been
required in the pelagic longline fishery
since 2012 (DFO 2013; C. MacDonald,
DFO, pers. comm., 2019). There is no
mandatory hook size restriction for the
Canadian longline fleet, but license
holders almost exclusively use 16/0
circle hooks (C. MacDonald, DFO, pers.
comm., 2019). De-hooking and linecutting kits are required on swordfish
longline fishery vessels (C. MacDonald,
DFO, pers. comm. 2019).
Some fishing fleets in the Atlantic
Ocean (e.g., U.S., Canadian, ICCAT
vessels) use large circle hooks and
modified bait, but these measures are
not required in all areas (Watson et al.
2005; Gilman et al. 2007; Gilman 2011).
Some nations in the Wider Caribbean
Region have implemented circle hook
provisions; in Belize, the high seas
fishing fleet adopted the use of circle
hooks on 10 percent of the fleet and are
required to report capture of sea turtles
by longlines (Belize Fisheries
Department 2017). Because the
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measures are not widely required, the
number of vessels that do not employ
bycatch reduction measures is likely
higher than the number of vessels that
do, and so we conclude on the basis of
the best available information that
leatherback bycatch in pelagic longline
fisheries is still a significant threat
(Lewison et al. 2015).
Leatherback interactions with bottom
longlines also occur. Directed shark
fisheries using bottom longlines in the
Atlantic Ocean and GOM may capture
or entangle leatherback turtles (NMFS
2012), and the GOM reef fishery is also
anticipated to take leatherback turtles
(NMFS 2011). On February 7, 2007,
NMFS published a rule that required
commercial shark bottom longline
vessels to carry the same dehooking
equipment as the pelagic longline
vessels; this rule was promulgated to
reduce post-interaction mortality (72 FR
5633).
The Canadian east coast groundfish
longline fishery targets a wide variety of
groundfish species, including cod,
haddock, pollock and white hake.
Observer coverage has ranged from 2 to
30 percent depending on area, and there
have been no reported interactions of
leatherback turtles in the observer
database since 2001 (DFO 2012).
However, there have been three reports
from Quebec logbooks and 10 reports of
interactions with groundfish longline
gear to non-governmental groups (DFO
2012). This indicates that the risk of
interactions in this gear may be higher
than documented through the observer
program.
Bottom longlines are also used in the
Mediterranean Sea (Casale 2008). While
there have not been any documented
leatherback captures from this gear type,
loggerheads have been caught at high
rates in Tunisia, Libya, Greece, Turkey,
Egypt, Morocco, and Italy (Casale 2008),
and interactions with leatherback turtles
are possible.
Commercial pelagic longline fisheries
do not operate in some Caribbean
nations, such as in Panama where effort
is limited to vessels under six tons
(Executive Decree 486, December 28,
2010). However, other Caribbean
nations allow commercial pelagic
longline fishing, and many find
leatherback turtles with longline hooks
(Re´serve Naturelle de l’Amana data in
Berzins, Office National de la Chasse et
de la Faune Sauvage, pers. comm., 2018
and KWATA data in Berzins 2018).
While no longlines exist in the
Caribbean Dutch nations of Bonaire, St.
Eustatius and Saba, there are efforts to
introduce circle hooks into the trolling
fishery (Ministry of Economic Affairs
2014). We consider longline bycatch to
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be a widespread threat to this DPS,
likely resulting in the loss of thousands
of individuals annually.
Trawl Fisheries
Leatherback turtles may interact with
bottom and midwater trawl gear
throughout the North Atlantic Ocean.
The highest reported trawl bycatch of
leatherback turtles of the NW Atlantic
DPS is likely from the southeastern U.S.
shrimp trawl fishery. Epperly et al.
(2003) anticipated an average of 80
leatherback mortalities a year in shrimp
trawl interactions, dropping to an
estimate of 26 leatherback mortalities in
2009 due to reduction in fishing effort
(Memo from Dr. B. Ponwith, SEFSC, to
Dr. R. Crabtree, SERO, January 5, 2011).
The 2014 NMFS Southeast U.S. Shrimp
Fishery Biological Opinion estimated
167 annual leatherback captures (144
mortalities) in the Atlantic Ocean and
GOM shrimp otter trawl fishery, with an
additional 34 captures in try nets (single
nets testing for shrimp concentrations;
NMFS 2014). The majority of these
interactions were in the GOM. However,
a more recent study of the GOM and
southeastern U.S. Atlantic coast shrimp
otter trawl fishery found fewer
leatherback captures: From 2007 to
2017, only 3 leatherback turtles were
reported in the observer data (with
coverage levels around 2 percent of
nominal days at sea; Babcock et al.
2018).
In the mid-Atlantic and northeastern
U.S. waters, observers reported 9
leatherback captures in bottom otter
trawl gear and 5 captures in midwater
trawls from 1993 to 2017 (NMFS
unpublished data 2018). In the Wider
Caribbean Region, leatherback turtles
are reported captured in trawls in
French Guiana (Ferraroli et al. 2004;
TEWG 2007), Guyana (Reichart et al.
2003), Suriname (Madarie 2010),
Trinidad (Forestry Division et al. 2010),
and Venezuela (Alio et al. 2010).
Since 1980, there were eight reports of
leatherback turtles incidentally captured
by trawl gear in British and Irish waters
(Pierpoint 2000). In the Mediterranean
Sea, leatherback bycatch in bottom
trawls off Tunisia (Caminas 2004) and
Egypt (Casale 2008) has also been
reported.
Trawl bycatch reduction measures
(e.g., turtle excluder devices (TEDs) are
in place in some nations. The
southeastern U.S. shrimp fishery has
required TEDs since the early 1990s.
However, TEDs that were initially
required for use in the U.S. Atlantic
Ocean and GOM shrimp fisheries were
less effective for leatherback turtles as
compared to smaller, hard-shelled turtle
species, because the TED openings were
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too small to allow leatherback turtles to
escape. To address this problem, NMFS
issued a final rule on February 21, 2003,
to amend the TED regulations (68 FR
8456) to require modified TEDs in the
southeastern United States (Atlantic
Area and GOM Area) that exclude
leatherback turtles, as well as large
benthic immature and sexually mature
loggerhead and green sea turtles. TEDs
are also required in summer flounder
trawls operating off Virginia (south of
Cape Charles) and North Carolina (64
FR 55860, October 15, 1999; 67 FR
19933, April 17, 2002).
TEDs are also used outside the United
States. Shrimp harvested with
commercial fishing technology that may
adversely affect sea turtles generally
cannot be imported into the United
States per Public Law 101–162, Section
609(b), enacted on November 21, 1989
(16 U.S.C. 1537 note). The import ban
does not apply to nations that have
adopted sea turtle protection programs
comparable to that of the United States
(i.e., require and enforce TED use) or
whose fishing activity does not present
a threat to sea turtles (e.g., nations
fishing in areas where sea turtles do not
occur). Although most certifications are
done on a national basis, the U.S. State
Department guidelines allow some
individual shipments of TED-harvested
shrimp from uncertified countries with
appropriate documentation.
Approximately 40 nations are currently
certified to import shrimp into the
United States, and five fisheries have
been determined as having their
products eligible for importation with
proper documentation (83 FR 22739,
May 16, 2018). Specifically, on May 8,
2018, the U.S. State Department
certified 13 nations on the basis that
their sea turtle protection programs (e.g.,
use of TEDs) are comparable to that of
the United States: Colombia, Costa Rica,
Ecuador, El Salvador, Gabon,
Guatemala, Guyana, Honduras, Mexico,
Nicaragua, Nigeria, Panama, and
Suriname. It also certified 26 shrimpharvesting nations and one economy as
having fishing environments that do not
pose a danger to sea turtles. In addition,
one fishery from a non-certified nation
within the range of the NW Atlantic
DPS (the French Guiana domestic trawl
fishery) has been authorized to import
shrimp products, provided certain
documentation accompanies the
imports. Sixteen nations have shrimping
grounds only in cold waters where the
risk of taking sea turtles is negligible:
Argentina, Belgium, Canada, Chile,
Denmark, Finland, Germany, Iceland,
Ireland, the Netherlands, New Zealand,
Norway, Russia, Sweden, the United
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Kingdom, and Uruguay. Ten nations
(Bahamas, Belize, China, the Dominican
Republic, Fiji, Jamaica, Oman, Peru, Sri
Lanka, and Venezuela) and Hong Kong
only harvest shrimp using small boats
with crews of less than five that use
manual rather than mechanical means
to retrieve nets or catch shrimp using
other methods that do not threaten sea
turtles. Use of such small scale
technology is not believed to adversely
affect sea turtles. For those nations
within the geographical range of the NW
Atlantic DPS, the threat of shrimp
trawling is minimized with TED use.
TEDs are also required in trawl fleets
in Trinidad, Belize, Brazil, and
Venezuela, but those gear modifications
do not currently meet the U.S.
certification protocol. On June 20, 2019,
the European Union passed a regulation
(PE–CONS 59/1/19 Rev 1) that requires
technical measures concerning: The
taking and landing of marine biological
resources; the operation of fishing gear;
and the interaction of fishing activities
with marine ecosystems. Specific to sea
turtles, the regulation requires shrimp
trawl fisheries to use a TED in European
Union waters of the Indian and West
Atlantic Oceans, consisting of waters
around Guadeloupe, French Guiana,
Martinique, Mayotte, Re´union and Saint
Martin.
TEDs are not required in
Mediterranean trawls. Some nations,
like Belize, St. Barthelemy, Venezuela
(industrial fishing only), and the
Caribbean Netherlands (Bonaire, St.
Eustatius, Saba), have banned trawling
(Bolivarian Republic of Venezuela
Official Gazette N° 5.877, March 14,
2008; Ministry of Economic Affairs
2016; Belize Fisheries Department
2017), and Costa Rica does not allow the
issuance of any new permits for shrimp
trawling (Costa Rica Ministry of
Environment and Energy 2017). Curacao
prohibits fishing in its territorial waters
and inland bays with dragnets (and
certain fish traps). These initiatives
reduce the impact of trawling on
leatherback turtles.
Pot/Trap Fisheries
Leatherback turtles are commonly
entangled in the vertical lines of pot and
trap gear. Entanglements have been
mostly reported from U.S. and Canadian
waters, but line entanglements have
occurred in other areas where similar
gear is used (e.g., Britain; Godley et al.
1998).
Due to high numbers of entanglement
reports, a Sea Turtle Disentanglement
Network (STDN) was established by
NMFS in the northeastern United States
(Maine to Virginia) in 2002. This
program relies primarily on reports from
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the public and subsequent
documentation and disentanglement by
trained responders. From 2008 to 2017,
267 leatherback entanglements were
reported in vertical fishing line (STDN
unpublished data). Of those fisheries
that could be identified, 79 were lobster,
21 were fish traps or fish lines, 18 were
conch (or a combination of conch and
lobster), and 5 were crab gear; 144
entanglements were from unidentifiable
fishing gear. While most unknown
vertical line entanglements likely
involve pot/trap gear, this cannot
always be conclusively determined. The
majority of the leatherback turtle reports
(67 percent) were from Massachusetts
waters. Of the 267 leatherback
entanglements, 221 were released alive
and 46 were found dead.
Given the nature of their injuries, it is
probable that not all animals released
alive from entanglements survived.
Currently there are limited empirical
data on leatherback survival from pot/
trap entanglements. Innis et al. (2010)
found that at least some of the
disentangled individuals were able to
resume normal behavior and migratory
patterns, but two leatherback turtles
were entangled at least twice, and a
third disentangled turtle had significant
forelimb skin and muscle injuries. The
effects of entanglement may be sublethal initially, but could result in
subsequent mortality. By assessing the
injuries experienced by each turtle that
was documented to have been entangled
and using NMFS’ post-interaction
mortality guidance (NMFS 2017), the
resulting mortality rate for northeastern
U.S. vertical fishing line interactions for
all sea turtle species combined was
calculated at 55 percent from 2013 to
2017 (NMFS unpublished data). When
the mortality estimate includes those
turtles that were not disentangled and
assumed to have died, the rate increases
to 61 percent. As a result (and applying
the latest 5 year mortality rate to the last
10 years of entanglement data), 147 to
163 leatherback turtles died from
vertical fishing line gear (most of which
were likely pot/trap gear) in the
northeastern U.S. waters from 2008 to
2017, based on opportunistically
reported data. An additional 36
leatherback turtles were reported
entangled in trap/buoy lines from North
Carolina to Texas from 2008 to 2017
(STSSN unpublished data). Of those 36
entanglements, 32 turtles were found
alive and 4 dead, but these southeastern
U.S. numbers do not incorporate
potential post-interaction mortality so
the total lethal interactions were likely
higher. Further, this information is
likely an underestimate of actual
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entanglements and mortality given the
opportunistic reporting nature of the
program; therefore, it is clear that
leatherback interactions with vertical
fishing lines are a threat to this DPS.
Entanglements in Canadian waters are
also frequently reported under
circumstances similar to the U.S. STDN
program, i.e., opportunistically by
fishermen or the public. Between 1998
and 2014, 205 leatherback
entanglements were reported in Canada
along the Atlantic coast, with most from
Nova Scotia (136) and Newfoundland
(40; Hamelin et al. 2017). Entanglements
mostly involved pot fisheries (44
percent; n = 91), including snow crab (n
= 37), inshore lobster (n = 31), rock crab
(n = 10), whelk (n = 8), and hagfish (n
= 3) fisheries. Trap net fisheries were
involved in 26 percent of the
entanglements (n = 53). Of the overall
205 reports, the majority of turtles were
reported alive and successfully released
(n = 174), and the other 15 percent (n
= 31) were reported dead in gear.
However, the number of dead turtles is
likely an underestimate of actual
entanglement-associated mortality
(Hamelin et al. 2017).
Leatherback turtles are also found
entangled in vertical fishing lines in
European waters. Since 1980, 83
leatherback turtles were bycaught in
British and Irish waters, with the
method of capture identified in 58 cases
(Piedpoint 2000). The majority of
captures (n = 36) were rope
entanglements, usually buoy lines used
in pot fisheries for crustaceans or whelk,
with a 61 percent recorded mortality
(Pierpoint 2000).
Some types of aquaculture use
vertical lines similar to pot/traps and
may pose an entanglement risk (Price et
al. 2017). Four leatherback turtles (two
alive, two dead) in Canadian and U.S.
waters have been opportunistically
reported in aquaculture gear to date
(Price et al. 2017). However, as this
industry is anticipated to grow in the
near future, leatherback interactions
with aquaculture lines, and subsequent
injury or mortality, may increase.
These data comprise the best available
information on pot/trap fishery
interactions with the NW Atlantic DPS.
However, due to the high probability of
underreporting leatherback turtle
entanglements by fishers, the ad hoc
nature of public reporting, and the
uncertainty about post-release
survivorship, the leatherback mortality
rate due to entanglements in vertical
lines is likely underestimated (Hamelin
et al. 2017). Estimates indicate that
approximately 622,000 vertical lines are
deployed from fishing gear in U.S.
waters from Georgia to the Gulf of
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Maine (Hayes et al. 2018). There are
currently no existing mitigation
measures to reduce leatherback bycatch
in vertical fishing lines, but efforts to
reduce the amount of vertical lines in
the water to assist with large whale
conservation in the United States may
help reduce the impact to the DPS
(https://
www.greateratlantic.fisheries.noaa.gov/
protected/whaletrp/).
Other Gear Types
Leatherback turtles are also
susceptible to bycatch in pound nets,
weirs, and purse seine fisheries. In the
United States, pound nets set in Virginia
waters have entangled leatherback
turtles. On June 23, 2006, NMFS issued
a regulation (71 FR 36024) requiring
offshore pound nets set in a portion of
the lower Chesapeake Bay from May 6
through July 15 of each year to use
modified pound net leaders, a gear
modification consisting of vertical hard
lay lines spaced at least two feet apart
on the top portion of the leader, and
eight inch or smaller stretched mesh on
the bottom portion of the leader. From
2013 to 2017, 16 leatherback turtles
have been found entangled in the hard
lay lines of the leaders, of which two
were dead (NMFS 2018). While
individuals may continue to be
entangled in modified pound net
leaders, the impact of the pound net
fishery on the NW Atlantic DPS is likely
minor given the few nets set in the
lower Chesapeake Bay using this gear
(approximately four to six) and the
frequency of live interactions. From
2008 to 2017, the STDN also
documented leatherback captures in
weirs set off Massachusetts; these turtles
were found alive, either entangled in the
netting (n = 2) or free swimming in the
weir (n = 4).
Purse seines are used to catch a
variety of fish species and are
commonly used in the ICCAT area to
catch tuna (Angel et al. 2014).
Leatherback captures have occurred in
Atlantic purse seine fisheries, and this
bycatch may have a minor impact on the
DPS. In British and Irish waters, two
leatherback turtles were reported to be
captured in purse seine gear between
1980 and 2000 (Pierpont 2000).
Clermont et al. (2012) reported a total
capture of 67 leatherback turtles in more
than 9000 observed Atlantic purse seine
sets between 1995 and 2011, with only
four found dead (representing 10
percent observer coverage). Most of the
interactions were adults (75 percent).
However, not all of the purse seine
effort reported by Clermont occurs in
the NW Atlantic DPS range. Thus, purse
seine interactions with this DPS may be
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a fraction of the total captures reported.
For those purse seines in the ICCAT
region using fish aggregating devices
and for those setting over freeswimming tuna schools, the effort
(through 2011) was concentrated in the
tropics, off West Africa between
Namibia and Mauritania and off
Venezuela (Clermont et al. 2012; Angel
et al. 2014). While leatherback and
purse seine interactions may occur
where distribution and effort overlap,
the magnitude of the purse seine
impacts on the NW Atlantic DPS is
lower than the bycatch values presented
in Clermont et al. (2012). Further, Angel
(2014) found that the direct impacts on
turtles from purse seine fishing
operations appears to be minor in
comparison to the impacts from longline
fishing, especially as most purse seine
captures are released alive.
Summary of Fisheries Bycatch
We conclude that most immature and
adult leatherback turtles of this DPS are
exposed to bycatch in multiple fisheries
throughout their range. Bycatch in
gillnet fisheries, in particular, is a major
threat with high mortality rates (Lee
Lum 2006; Gilman et al. 2010; Girondot
2015), annually killing thousands of NW
Atlantic leatherback turtles. When set
off nesting beaches, gillnets result in
high mortality of nesting females and
mature males (Lee Lum 2006; Eckert
2013). Longline bycatch is considered to
be a widespread threat throughout the
DPS and a primary source of leatherback
mortality (Lewison et al. 2004),
resulting in the death of thousands of
leatherback turtles annually. In general,
bycatch mortality reduces abundance by
removing individuals from the
population. When nesting females are
killed, it also reduces productivity. We
conclude that fisheries bycatch is the
primary threat to the NW Atlantic DPS.
Vessel Strikes
Vessel strikes are a threat to the NW
Atlantic DPS. Injuries from vessel
strikes may include blunt force trauma
and propeller parallel slicing wounds
affecting the carapace, flippers, head,
and/or underlying organs (Work et al.
2010). Most of what is known about
vessel strikes comes from stranding
records; the most extensive stranding
network is found in the United States:
The Sea Turtle Stranding and Salvage
Network (STSSN). In the United States
(Maine through Texas), 957 leatherback
turtles were reported stranded,
captured, or entangled from 2008 to
2017, and of those, 204 had probable
vessel-related injuries (STSSN
unpublished data). For example, at least
72 leatherback turtles stranded in
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Massachusetts with vessel strike
wounds between 2006 and 2018,
including at least three adult females
that had previously been documented
nesting in the Caribbean (Dourdeville et
al. 2018; Mass Audubon Wellfleet Bay
Wildlife Sanctuary, unpublished data,
2019). It is sometimes difficult to
determine whether the vessel related
wounds occurred before or after the
turtle died (Stacy et al. 2015). However,
a recent study estimated that
approximately 93 percent of Florida
stranded turtles with vessel strike
wounds were killed by those injuries
(Foley et al. 2019). Based on the best
available information, it is reasonable to
conclude that approximately 190
leatherback turtles were killed as a
result of vessel strikes in U.S. Atlantic
and GOM waters from 2008 to 2017.
This number is likely an underestimate
as strandings represent a small
percentage of turtles that are injured or
die at sea, and many vessel strikes are
not reported, detected, or recovered.
Vessel strikes have been documented
in other nations as well, including in
Portugal (Nicolau et al. 2016), Britain
(Godley et al. 1998), and off the coast of
Tunisia in the Strait of Sicily (Karaa et
al. 2013; Caracappa et al. 2017). While
there is very limited observational
information on vessel collisions in
Atlantic waters of Canada, there has
been at least one recorded vessel strike
(DFO 2012). More recently, an injury
assessment of leatherback turtles (n =
228) on Atlantic Canada foraging
grounds and on a Trinidad nesting
beach found only 1.3 percent of turtles
exhibited injuries consistent with vessel
strikes (Archibald and James 2018).
However, this low injury rate may
indicate that there is low survivorship
of vessel strikes. Females with carapace
damage from propellers have been also
observed on Costa Rican nesting
beaches (de Haro et al. 2006).
Leatherback behavior data can help
predict the potential for vessel strikes.
Based on telemetry data for leatherback
turtles (n = 15) on the northeastern U.S.
shelf, leatherback turtles spent over 60
percent of their time in the top 10 m of
the water column and over 70 percent
of their time in the top 15 m (Dodge et
al. 2014). Additional turtle-borne
camera and autonomous underwater
vehicle research in the waters off
Massachusetts suggests that turtles
surface frequently and engage in
subsurface swimming (within the top 2
m) when occupying shallow, wellmixed, coastal environments, increasing
the probability of a vessel strike (Dodge
et al. 2018). Based on 24 free swimming
leatherback turtles tagged in Canadian
waters from 2008 to 2013, Wallace et al.
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(2015) found these leatherback turtles
primarily occupied the upper 30 m of
the water column and had shallow 4 to
6 minute dives. Given most leatherback
activity occurs in the top 15 to 30
meters of the water column in temperate
shelf waters of the NW Atlantic Ocean
and vessel traffic is high along the U.S.
East coast, the risk of vessel strikes is
likely higher than the documented
interactions would suggest (DFO 2012;
Hamelin et al. 2014).
While observational data are limited,
it is reasonable to conclude that, based
upon the best available information,
mortality due to vessel strikes may
occur wherever vessel traffic and
leatherback distribution (juvenile and
adult) overlap. The impact is likely
minimized in areas with less frequent
vessel traffic (e.g., less developed areas)
and decreased leatherback turtle
presence. Nesting females and mature
males may be especially vulnerable to
vessel strikes because they occur in the
waters off nesting beaches, which are
coastal areas where vessel traffic is more
prevalent. Vessel strikes affect the NW
Atlantic DPS by lowering abundance (if
the interaction results in mortality) and
affecting future reproductive potential
(productivity) when nesting females are
killed. We conclude that vessel strikes
pose a threat to the NW Atlantic DPS.
Pollution
Pollution includes contaminants,
marine debris, and ghost fishing gear.
The detection of pollution impacts on
leatherback turtles is opportunistic and
thus likely underestimated. While
plastic ingestion is not always fatal, it
can reduce ability to feed, affect
swimming behavior and buoyancy
control, potentially lead to chemical
contamination and chronic effects, and
weaken physical condition, which
could impair the ability to avoid
predators and survive threats (Nelms et
al. 2016). Entanglement in marine
debris results in injuries that can reduce
fitness, cause eventual death, reduce
ability to avoid predators, reduce ability
to forage and/or swim efficiently due to
drag, and lead to starvation or drowning
(Nelms et al. 2016). Pollution on the
beach and in the water occurs
throughout the range of the NW Atlantic
DPS.
Dow et al. (2007) defined marine
pollution as agriculture, petroleum,
sewage, industrial runoff, vessel
discharges, declining water quality, and
marine debris. They found pollution in
the marine environment to be among the
greatest threats to all sea turtle species
in the Wider Caribbean Region. Dow et
al. (2007) defined beach pollution as
agriculture, petroleum/tar, sewage,
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industrial runoff, and beach litter/
debris; they found pollution on the
beach to be a threat. Pollution on the
beach and in the water occurs
throughout the range of the NW Atlantic
DPS.
Leatherback turtles are susceptible to
adverse effects from pollution. Marine
pollution, including direct
contamination and structural habitat
degradation, can also affect leatherback
habitat. In particular, the Mediterranean
is an enclosed sea, so organic and
inorganic wastes, toxic effluents, and
other pollutants rapidly affect the
ecosystem (Camin˜as 2004).
Of particular concern, due to their
immune, reproductive, and endocrine
disrupting nature, are persistent organic
pollutants (POPs), such as
polychlorinated biphenyls (PCBs),
polybrominated diphenyl ethers
(PBDEs), and pesticides (Bergeron et al.
1994; Bishop et al. 1991, 1998; Keller et
al. 2004). These chemicals have been
identified in both adults and eggs in
several areas occupied by this DPS.
Guirlet et al. (2010) measured maternal
transfer of organochlorine contaminants
(OCs) from 38 nesting females in French
Guiana. PCBs were found to be the
dominant OC, followed by pesticides,
but OC concentrations were lower than
concentrations measured in other
marine turtles (potentially due to the
lower trophic level diet and offshore
foraging areas). All OCs detected in
nesting adults were detected in eggs,
suggesting a maternal transfer of OCs. In
French Guiana, hatching success has
been shown to be low when OCs are
present in the sand (most likely
originating from pesticide use in
plantations and malaria prophylaxis
(Guirlet 2005). However, a link between
OCs and embryonic mortality could not
be determined (Guirlet et al. 2010).
Stewart et al. (2011) also recorded PCB,
OC, and PBDE concentrations for
nesting and stranded leatherback turtles
in the southeastern United States. Their
results also suggested maternal transfer
of POPs in leatherback turtles, but
Stewart et al. (2011) found higher levels
of PCBs and pesticides than those found
in French Guiana (Guirlet et al. 2010).
While finding that leatherback
contaminant concentrations were
substantially lower than concentrations
in other reptile studies that
demonstrated toxic effects, Stewart et al.
(2011) suggested that sub-lethal effects
(especially on hatchling body condition
and health) may nevertheless be
occurring in this species. De Andres et
al. (2016) similarly monitored PCB and
PBDE concentrations in eggs laid in
Costa Rica (18 nests). POP levels were
similar to those reported in French
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Guiana nesting females (Guirlet et al.
2010) and slightly lower than those in
Florida (Stewart et al. 2011). Further, De
Andres et al. (2016) found a significant
negative relationship between PBDE
levels and hatching success, suggesting
potential harmful effects of these
contaminants on leatherback
reproduction. OCs (and mercury) have
also been documented in turtles that
stranded in the United Kingdom
(Godley et al. 1998). A leatherback that
stranded off the coast of Wales, U.K.
was found with PCB levels one-to-three
orders of magnitude higher than the
lowest levels reported for fish taken in
the North Atlantic, but similar to the
lowest concentrations reported from
oceanic cetaceans (Davenport et al.
1990). Even with the recent restriction
of the use of POPs, due to the
widespread persistent nature of these
chemicals and continuing atmospheric
deposition (Ross et al. 2009) it is
probable that similar chemical
concentrations occur in other areas of
this DPS.
Other contaminants have also been
documented in leatherback turtles and
their eggs. Heavy metals (e.g., arsenic,
cadmium, chromium, mercury, lead,
etc.) enter the environment from a
variety of sources (Guirlet et al. 2008;
Perrault 2012). In particular, mercury
can affect a variety of functional
processes in wildlife, including the
nervous, excretory and reproductive
systems (Wolfe et al. 1998). Mercury,
cadmium, and lead were recorded in
nesting females (n = 46) and eggs in
French Guiana (Guirlet et al. 2008).
Maternal transfer of all three elements
was documented, and female lead levels
increased throughout the nesting season
(Guirlet et al. 2008). This could be
explained, in part, by external
contamination via ingestion of
contaminated prey or polluted water
during nesting, as the French Guiana
coast environment is exposed to
significant environmental pollution via
anthropogenic and natural sources.
While mercury concentrations were
lower than values reported for other sea
turtle species, cadmium levels
documented in French Guiana were at
the same level shown to impact gonadal
development in other turtle species and
may impact reproductive processes and
lower fertility (Guirlet et al. 2008). In
Massachusetts, entangled turtles had
significantly higher blood lead
concentrations than directly captured
turtles (Innis et al. 2010). While similar
to those reported in French Guiana
(Guirlet et al. 2008), blood
concentrations of mercury and cadmium
were at levels high enough to induce
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carcinogenic, teratogenic, and toxic
effects in a variety of species (Innis et
al. 2010).
Mercury and selenium have also been
recorded in nesting females and eggs in
Florida and St. Croix. Animals
persistently exposed to mercury can
experience selenium deficiency, which
is of concern because selenium is
important to hatching and emergence
success (Perrault et al. 2011). However,
high levels of selenium can be toxic and
negatively impact hatching success
(Perrault et al. 2013). Mercury
concentrations in nesting females from
Florida were found to be higher than in
St. Croix, which could be a result of
different migratory and foraging areas,
whereas hatchling blood mercury values
were higher in St. Croix (Perrault et al.
2011; Perrault et al. 2013). It is
interesting to note that in St. Croix, no
correlations were found between
mercury or selenium concentrations and
hatching or emergence success, which is
different from results in Florida
(Perrault et al. 2011; Perrault et al.
2013). Hazard quotient results by
Perrault et al. (2013, 2014) imply that
mercury and selenium levels could pose
a threat to leatherback turtle
reproductive success and/or hatchling
health and survival. Leatherback
hatching and emergence success rates
are already low compared to other
species of sea turtles (Bell et al. 2004;
Perrault et al. 2011), so the impacts of
pollution and contamination on
hatching success is a notable concern. In
addition, mercury was found to be
higher in adults than juveniles/subadults stranded along the U.S. Atlantic
coast, suggesting potential physiological
concerns due to accumulation and
ongoing inputs into the environment
(Perrault et al. 2012). It is clear that
additional long-term research is needed
to better understand the relationship of
non-essential elements in turtle
development and reproduction.
Marine debris (most notably plastic
pollution) is a threat throughout the
range of the NW Atlantic DPS (Girondot
2015). Several global reviews have
outlined the persistent and widespread
nature of the issue, both as an ingestion
and an entanglement threat (Mrosovsky
et al. 2009; Schuyler et al. 2014; Nelms
et al. 2016; Lynch 2018). Law et al.
(2010) assessed plastic content at the
surface of the western North Atlantic
Ocean and Caribbean Sea from 1986 to
2008, and found the highest
concentration of plastic debris was
observed in subtropical latitudes and
associated with large-scale convergence
zones, which include foraging areas
targeted by leatherback turtles.
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Ingestion of marine debris is a
concern for leatherback turtles,
especially given the similarity of their
preferred prey (e.g., gelatinous
zooplankton) to some plastics. In
particular, plastic bags appear similar to
jellyfish in the marine environment,
leading to mistaken and inappropriate
triggering of the sensory cue to feed
(Schuyler et al. 2014; Nelms et al. 2016).
While plastic ingestion is not always
fatal, it can reduce ability to feed, affect
swimming behavior and buoyancy
control, potentially lead to chemical
contamination and chronic effects, and
weaken physical condition, which
could impair the ability to avoid
predators and survive threats (Nelms et
al. 2016).
Marine debris ingestion can occur in
any location, but given the enclosed
nature of the sea and intense human
pressure, the Mediterranean Sea in
particular is a hot spot for plastic
marine debris and other pollutants
(Camin˜as 2004; Cozar et al. 2015).
Marine debris ingestion has been
documented from leatherback turtles
stranded in Tunisia (Karaa et al. 2013),
Israel (Levy et al. 2005), the northern
Adriatic Sea (Poppi et al. 2012), and the
Strait of Sicily (Caracappa et al. 2017).
Of particular note, 30 to 73 percent of
turtles stranded in the Bay of Biscay
(France) were found to have ingested
plastic annually from 1979 to 1999 (out
of 87 leatherback turtles necropsied;
Duguy et al. 2000). The seasonal rate of
ingestion was inversely related to the
abundance of jellyfish, leading the
authors to propose that the depletion of
jellyfish led to debris ingestion as
potential prey. Cozar et al. (2015)
conclude that the effects of plastic
pollution on marine life are anticipated
to be frequent in the high plasticaccumulation region of the
Mediterranean Sea.
In U.S. waters, marine debris
ingestion has also been documented in
stranded leatherback turtles. However,
ingestion does not always cause
mortality and is typically an incidental
finding. Of 41 leatherback turtles
necropsied from North Carolina to
Texas between 2008 and 2017, 17 had
ingested plastics or marine debris
(STSSN unpublished data 2018). From
Maine to Virginia during that same time
period, 10 necropsies detected
ingestion, but the total number of
necropsied turtles, out of the 677
strandings in the region, is currently
unknown. It is likely that many more
stranded turtles ingested some level of
marine debris (STSSN unpublished data
2018). Out of 33 leatherback turtles
examined in New York Bight (an area
with dense population), 30 percent had
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synthetic material ingestion, mostly
consisting of thin, clear plastic (Sadove
et al. 1989). Of two leatherback turtles
stranded in North Carolina during 2017
whose gastrointestinal tracts were
analyzed, microplastics were present in
both (Duncan et al. 2018).
Marine debris ingestion is not limited
to microplastics or plastic bags. Off the
northeastern U.S. coast, necropsies of
disentangled leatherback turtles that
have died post-release have documented
considerably large pieces of plastic (e.g.,
83 by 35 cm) in their stomachs (Innis et
al. 2010). These numbers likely
underestimate the true marine debris
ingestion rate because many turtles
likely ingest marine debris and do not
strand.
Leatherback turtles can also become
entangled in marine debris. From 2008
to 2017, the Northeast U.S. STDN
documented 24 entanglements from
miscellaneous sources not attributed to
obvious fisheries entanglements, as
described above (STDN unpublished
data). These unknown entanglements
could involve a myriad of sources but
are considered as entangling marine
debris. The Sea Turtle Recovery Action
Plan for the Republic of Trinidad and
Tobago noted that entanglement in lost
or abandoned fishing gear (primarily
nets) poses a threat to leatherback
turtles in the marine and terrestrial
environment (Forestry Division et al.
2010).
Marine debris is also a problem on
nesting beaches and can reduce nesting
success. Pollution and debris often are
deposited on high energy beaches,
which are also the preferred nesting
habitat of leatherback turtles (TEWG
2007). Coastal and inland littering
(which can ultimately reach the sea) is
a problem throughout Trinidad and
Tobago, and ocean borne debris is
particularly prevalent on the east and
north coasts, which host the main
leatherback nesting beaches (Trinidad
and Tobago Forestry Division et al.
2010). Extensive debris on nesting
beaches is not uncommon throughout
the Caribbean, often carried by rivers to
the sea and later washed ashore (e.g., in
Costa Rica; Chaco´n-Chaverri and Eckert
2007). Debris on nesting beaches may
impede females during the nest-site
selection stage, limit and degrade the
amount of habitat available, and/or
result in aborted nesting attempts
(Chaco´n-Chaverri and Eckert 2007). If
line or netting is encountered on nesting
beaches, entanglement of nesting
females and hatchlings is also a risk.
The majority of the NW Atlantic DPS
is exposed to pollution throughout all
life stages. These threats are a result of
the developed nature of many of the
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nations within the range of the DPS. The
degree of impact is difficult to quantify,
especially given the widespread nature
of pollution and the diverse types of
impacts. Contaminants may affect this
DPS by reducing productivity, if
hatching success is lowered, and by
lowering abundance, if contamination
results in mortality. Marine debris
affects the DPS by lowering abundance,
when it causes death through ingestion
or entanglement, and reducing
productivity, when hatchlings and
nesting females are affected. While, we
do not have quantitative estimates of the
number of individuals that are killed or
injured as a result of pollution, we
conclude that it is prevalent throughout
the range of the DPS and constitutes a
threat to the NW Atlantic DPS.
Oil and Gas Exploration
Oil and gas activities have the
potential to impact the NW Atlantic
DPS directly (e.g., exposure to oil
following oil spills) and indirectly (e.g.,
increased probability of vessel strikes
and habitat degradation/destruction). In
addition to lethal effects, sublethal
effects may occur and include
displacement from primary foraging
areas with accompanying energy costs
(TEWG 2007).
Several areas within the range of the
NW Atlantic DPS have intense oil and
gas development and exploration close
to major nesting beaches. The potential
for oil spills is of particular concern in
the Wider Caribbean Region due to its
effect on all life stages in the marine
environment. The biggest oil producing
nations in South America are Brazil,
Mexico, Venezuela, and Colombia.
Although only three Caribbean nations
currently have exportable oil and
natural gas reserves (Barbados, Cuba,
and Trinidad and Tobago, with Trinidad
and Tobago the only significant
exporter), in 2017, a major oil field was
discovered off Guyana, which will
likely lead to extensive new
development and extraction. As a result,
marine traffic is likely to increase in the
area as well as the possibility for oil
spills. In Panama, contamination from
oil spills, primarily in area of the TransIsthmus oil pipeline and the Panama
Canal, is of particular concern
(Bra¨utigam and Eckert 2006; Ruiz et al.
2006). Some Caribbean nations (e.g.,
Belize, French Guiana) have permanent
moratoria on oil and gas exploration in
offshore waters.
In the United States, oil and gas
extraction primarily occurs in the GOM
(BOEM 2016; BOEM 2017), an area with
leatherback foraging and migratory
habitat (Aleksa et al. 2018). Increased
shipping traffic and marine noise due to
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oil and gas explorations in the GOM
pose a direct threat for leatherback
turtles in foraging grounds and
migratory routes, due to the potential for
vessel strikes and harassment (Wallace
et al. 2017; Ward 2017). Oil spills
regularly occur in the GOM, from small
amounts of varying types of oil product
to large catastrophic spills. In 2010, a
major oil spill occurred in the northcentral GOM, affecting important
foraging habitat used by leatherback
turtles (Deepwater Horizon NRDA
Trustees 2016). Evans et al. (2012)
tracked a post-nesting leatherback from
Chiriqui Beach, Panama, into the GOM
during the Deepwater Horizon oil spill.
The track followed similar tracks from
turtles in previous years and did not
seem to change once entering areas with
visible oil slicks (on two occasions).
Injuries to leatherback turtles caused by
the GOM Deepwater Horizon oil spill
could not be quantified (Deepwater
Horizon NRDA Trustees 2016).
However, given that the GOM is
important habitat for leatherback turtles
(Aleksa et al. 2018) and leatherback
turtles were documented in the
Deepwater Horizon oil spill zone during
the oil spill period, the Deepwater
Horizon NRDA Trustees (2016)
concluded that leatherback turtles were
exposed to Deepwater Horizon oil, and
some portion of those exposed likely
died.
In Atlantic Canada, impacts from oil
and gas may also occur. Several
petroleum production projects occur
offshore of Nova Scotia (https://
www.cnsopb.ns.ca/offshore-activity/
offshore-projects). Howard (2012)
determined that oil pollution from
coastal refineries, ships, small engine
vessels, and oil and gas exploration and
production is a risk to leatherback
survival in Canada. There are also
offshore oil and gas platforms in the
North (United Kingdom, Denmark) and
Mediterranean Seas, where similar
impacts to leatherback turtles may also
occur (EU Offshore Authorities Group
2018; https://euoag.jrc.ec.europa.eu/
node/63). In particular, the
Mediterranean Sea has been declared a
‘‘special area’’ by the International
Convention for the Prevention of
Pollution from Ships (MARPOL), in
which deliberate petroleum discharges
from vessels are banned, but numerous
repeated offenses are still thought to
occur (Pavlakis et al. 1996). Some
estimates of the amount of oil released
into the region is as high as 1,200,000
metric tons (Alpers 1993). Direct oil
spill events also occur, as in Lebanon in
2006 when 10,000 to 15,000 tons of
heavy fuel oil spilled into the eastern
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Mediterranean (UN Environment
Programme 2007).
In summary, oil and gas activities are
prevalent in foraging, migratory, and
offshore nesting habitats of the NW
Atlantic DPS, potentially exposing all
life stages to oil associated threats, such
as direct miring in oil, oil ingestion,
vessel strikes, and nesting beach
contamination. Oil and gas activities
have the potential to affect this DPS by
reducing productivity (e.g., if hatching
success is reduced by oil spills) and
potentially lowering abundance (e.g., if
oil exposure results in mortality). As
such, oil and gas activities are a threat
to the NW Atlantic DPS.
Natural Disasters
Natural disasters, such as hurricanes
and other storms, and natural
phenomena, such as Sargassum events
on or near nesting beaches, pose a threat
to the NW Atlantic DPS.
Hurricanes are common in the
Caribbean and southeastern United
States. Hurricanes and tropical storms
impact nesting beaches by increasing
erosion and sand loss and depositing
large amounts of debris. In 2017,
Hurricane Maria devastated the islands
of Dominica, St. Croix, and Puerto Rico,
and even though the nesting season was
nearly over, many beaches were
impacted, including Maunabo, Puerto
Rico (one of the most abundant nesting
beaches on the island; R. Espinoza,
Conservacio´n ConCiencia, pers. comm.,
2017). Dewald and Pike (2014) found
that a lower level of leatherback nesting
attempts occurred on sites that were
more likely to be impacted by
hurricanes. These types of storm events
may ultimately affect the amount of
suitable nesting beach habitat,
potentially resulting in reduced
productivity, especially as leatherback
turtles typically nest on high energy
beaches (TEWG 2007).
Hurricanes may also result in egg loss
by destroying and inundating nests.
However, hurricanes are usually
aperiodic so the impacts are expected to
be infrequent. Hurricanes also typically
occur after the peak of the leatherback
hatching season and would not be
expected to affect the majority of
incubating nests (USFWS 1999). That
said, according to the Intergovernmental
Panel on Climate Change (IPCC), climate
change may be increasing the frequency
and patterns of hurricanes (IPCC 2014)
potentially causing such impacts to
nests to become more common in the
future.
Sargassum is a genus of macroalgae
found in temperate and tropical waters.
When large amounts of Sargassum wash
ashore, they form thick mats that have
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the potential to disrupt females’ nesting
activities and impede hatchlings’ access
to the ocean (Maurer et al. 2015). In
2011 and 2015, large amounts of
Sargassum were present in the
Caribbean (mainly Trinidad and Tobago
and Grenada) and frequently washed
ashore, covering large expanses of sandy
shoreline on nesting beaches. While
females still nested in these areas,
hatchlings needed intervention to reach
the ocean (Wang and Hu 2016;
Audroing, TVT, pers. comm., 2018; K.
Charles, Ocean Spirits Inc., pers. comm.,
2018). Most recently, large amounts of
Sargassum were found in 2018 on
Caribbean beaches, causing Barbados to
declare a national emergency in June
2018. Such widespread blanketing of
Sargassum on leatherback nesting
beaches throughout the Caribbean has
the potential to impact future hatching
success and survival.
In summary, natural disasters and
phenomena have the potential to impact
the NW Atlantic DPS. However, given
the infrequent and temporary nature of
the occurrences, only a small proportion
of eggs, hatchlings, and nesting females
are exposed to these threats. Impacts
include egg and hatchling mortality that
affect productivity of the DPS. Seasonal
losses at individual beaches may be
large, but we do not expect such
impacts to be spatially or temporally
widespread. However, we conclude that
natural disasters pose a threat to the
DPS.
Climate Change
Climate change is a threat to the NW
Atlantic DPS. The impacts of climate
change include increases in
temperatures (air, sand, and sea
surface); sea level rise; increased coastal
erosion; more frequent and intense
storm events; and changes in ocean
currents. These impacts may affect
leatherbacks through alterations of the
incubation environment, reduction of
nesting habitat, and changes in prey as
described in the following subsections.
Modeling results show that global
warming (rise in average surface
temperature) poses a ‘‘slight risk’’ to
females nesting in French Guiana and
Suriname relative to those nesting in
Gabon, Congo, and West Papua (Dudley
et al. 2016). As global temperatures
continue to increase, some beaches will
experience changes in sand
temperatures, which in turn will alter
the thermal regime of incubating nests.
Changing sand temperatures at nesting
beaches may result in changing sex
ratios of hatchling cohorts and reduced
hatching output (Hawkes et al. 2009).
Leatherback turtles exhibit temperaturedependent sex determination (Binckley
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and Spotila 2015) and warmer
temperatures produce more female
embryos (Mrosovsky et al. 1984;
Hawkes et al. 2007). In the NW Atlantic
DPS, the pivotal temperature (the
temperature at which a sex ratio of 1:1
is produced) is estimated to be between
29.25 °C and 30.5 °C (Eckert et al. 2012)
but there are variations in
measurements (Girondot et al. 2018),
over time, and among locations. An
increase over that temperature would
result in more female hatchlings. Such
increases in female hatchling output
have already been documented (Patin˜oMartı´nez et al. 2012), and with an
increase in temperatures from climate
change, these trends are likely to
continue if other nesting factors remain
constant. For example, Patin˜o-Martı´nez
et al. (2012) developed a model to relate
measured incubation temperature to sex
ratio and estimated that females nesting
at Caribbean Colombian beaches
currently produce approximately 92
percent female hatchlings. Under all
future climate change scenarios,
complete feminization could occur as
soon as 2021 (Patin˜o-Martı´nez et al.
2012). In St. Eustatius, leatherback
hatchling production was female biased
from 2002 to 2012, with less than
approximately 24 percent of males
produced every year (Laloe¨ et al. 2016).
Future warming air temperatures will
exacerbate this female bias, and female
leatherback sex ratios are projected to
consistently reach 95 percent after 2028
on that island, which has dark and light
sand beaches (Laloe¨ et al. 2016).
Warming trends in Costa Rica are
expected to be higher than the global
average and resulting female-biased sex
ratios are also expected (Gledhill 2007).
While the assumption is that most
nesting beaches will become femalebiased due to increased sand
temperatures, this may not be the case
in all areas. In Grenada, increased
rainfall (another effect of climate
change) was found to have a cooling
influence on nests, so that more male
producing temperatures (less than 29.75
°C) were found within the clutches
(Houghton et al. 2007). Further, due to
the tendency of nesting females to
deposit some clutches in the cooler
intertidal zone of beaches, the effects of
long-term climate on sex ratios may be
mitigated (Kamel and Mrosovsky 2004;
Patin˜o-Martı´nez et al. 2012).
Hatching success is affected by
warming temperatures. Extremely high
sand/nest temperatures are anticipated
to result in embryonic mortality
(Gledhill 2007, Santidria´n Tomillo et al.
2012, Valentin-Gamazo et al. 2018). In
Costa Rica, warmer conditions can
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exacerbate the effects of biotic
contamination and mold infestations of
developing embryos (Gledhill 2007),
resulting in reduced hatching success.
Temperature increases are likely to be
associated with more extreme
precipitation and faster evaporation of
water, leading to greater frequency of
both very wet and very dry conditions
that reduce productivity (Patin˜oMartı´nez et al. 2014; Santidria´n Tomillo
et al. 2015). These impacts may affect
nests in different ways, but the result
(e.g., reduced hatching output) is
similar. Very wet conditions may
inundate nests or increase fungal and
mold growth, reducing hatching success
(Patin˜o-Martı´nez et al. 2014). Very dry
conditions may affect embryonic
development and decrease hatchling
output. Under climate change scenarios,
very dry conditions are expected for St.
Croix, an area already showing
decreased productivity and reduced first
time nesting female abundance
(Santidria´n Tomillo et al. 2015; Garner
et al. 2017). Santidria´n Tomillo et al.
(2015) assessed climatic conditions on
hatchling output at four nesting sites
(Sandy Point, St. Croix; Pacuare,
Caribbean Costa Rica; Playa Grande,
Pacific Costa Rica; Maputaland, South
Africa), and found that St. Croix had the
highest projected warming rate (+ 5.4
°C), highest absolute temperature and
lowest precipitation levels. With these
further increases in dryness and air
temperatures, hatchling productivity is
expected to be compromised by the end
of the 21st century in this area
(Santidria´n Tomillo et al. 2015).
Santidria´n Tomillo et al. (2015)
suggested that the lack of rain is what
reduces developmental success and
hatchling emergence. However, Rafferty
et al. (2017) evaluated long-term climate
data for St. Croix, using climate data
collected from a nearby weather station,
and found no significant trend in
incubation temperatures or precipitation
that could be associated with observed
decreases in productivity at this
location.
Finally, incubation temperatures can
also influence hatchling morphology
and locomotion (Mickelson and Downie
2010). Leatherback hatchlings
originating from nests incubated at
lower temperatures exhibited carapace
and front flipper length-width ratios
that significantly improved their
crawling speeds relative to those
hatchlings incubated at high
temperatures (Mickelson and Downie
2010).
Sea level rise is another threat to
leatherback turtles. Thornalley et al.
(2018) found that the Labrador Sea deep
convection and the Atlantic Meridional
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Overturning Circulation, a system of
ocean currents in the North Atlantic,
have been unusually weak over the past
150 years or so, and this weakened state
may have modified northward ocean
heat transport, as well as atmospheric
warming by altering ocean-atmosphere
heat transfer. Further, the documented
weakening of this system is related to
above-average sea level rise along the
U.S. East Coast (Caesar et al. 2018). Sea
level rise may result in intensified
erosion and loss of nesting beach habitat
(Fish et al. 2005; Fuentes et al. 2010;
Fonseca et al. 2013). In Bonaire, up to
32 percent of the current beach area
could be lost with a 0.5 m rise in sea
level, with lower, narrower beaches
being the most vulnerable (Fish et al.
2005). Ussa (2013) predicted a 20 to 25
percent loss in beach areas due to sea
level rise by the year 2100 within the
Archie Carr National Wildlife Refuge,
Florida, as well as areas adjacent to the
Refuge. With the threat of increasing sea
level rise, protection of developed
coastlines often involves shoreline
armoring that reduces the amount of
beach available, thus creating a smaller
amount of space for turtles to nest
(Hawkes et al. 2009). Along such
developed coastlines, rising sea levels
may cause severe effects on eggs,
because nesting females are forced to
deposit eggs seaward of shoreline
armoring, potentially subjecting them to
repeated tidal inundation and/or egg
exposure from exacerbated wave action
near the base of these structures.
Sea level rise is expected to result in
more nests being inundated, reducing
hatching success. On Playona Beach,
Colombia, Patin˜o-Martı´nez et al. (2014)
found that nests in wet sand suffered
higher mortality (emergence success of
zero percent for wettest nests to 64
percent for the driest nests), suggesting
that nesting success should be expected
to decrease under future climate change
sea level rise scenarios. Inundation is
likely to reduce hatching success
(Patin˜o-Martı´nez et al. 2008; Caut et al.
2010) and will continue to occur (or
worsen) with sea level rise.
However, leatherback turtles may be
less susceptible than other species of sea
turtles to loss of nesting habitat, because
they exhibit lower nest-site fidelity
(Dutton et al. 1999). Nesting beaches in
the Guianas are already highly dynamic
and interseasonally variable, and
leatherback nesting females have been
successful in those areas despite the fact
that some beaches disappear between
nesting years (Plaziat and Augustinus
2004; Kelle et al. 2007; Caut et al. 2010).
If global temperatures increase and there
is a range shift northwards, beaches not
currently used for nesting could in the
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48359
future become used by leatherback
turtles, potentially offsetting some loss
of accessibility to beaches in southern
portions of the range. Leatherbacks’
behavioral flexibility may allow for
opportunities to colonize new beaches,
but whether turtles can colonize nesting
areas that become available, either
thermally or geographically, by climate
change, and whether these colonized
areas provide incubation regimes that
will lead to successful nesting,
emergence success, and hatchling
fitness cannot be known at this time
(Hawkes et al. 2009).
Observed changes in marine systems
are associated with other aspects of
climate change, including rising water
temperatures, as well as related changes
in ice cover, salinity, oxygen levels, and
circulation. Ocean temperatures of the
U.S. northeastern continental shelf and
surrounding NW Atlantic waters have
warmed faster than the global average
over the last decade (Pershing et al.
2015). New projections for the U.S.
northeastern shelf and NW Atlantic
Ocean suggest that this region will
warm two to three times faster than the
global average and existing projections
from the IPCC may be too conservative
(Saba et al. 2015). This increase in
northeastern shelf waters is relevant for
NW Atlantic leatherback turtles, as they
rely on U.S. and Canadian waters to
forage during the warmer months (James
2005a, 2006b, 2007; Dodge 2014, 2015).
Global warming is expected to expand
leatherback foraging habitats into, and
increase residency time in, higher
latitude waters (James et al. 2006a;
McMahon and Hays 2006; Robinson et
al. 2009). For example, leatherback
turtles have extended their range in the
Atlantic north by around 200 km per
decade over the last two decades as
warming has caused the northerly
migration of the 15 °C sea surface
temperature (SST) isotherm, the lower
limit of thermal tolerance for
leatherback turtles (McMahon and Hays
2006). Documented weakening of the
Meridional Overturning Circulation is
related to above-average warming in the
Gulf Stream region and an associated
northward shift of the Gulf Stream
(Caesar et al. 2018). This weakening of
the deep, cold-water circulation in the
North Atlantic is likely to continue to
occur with global warming. Migratory
routes may be altered by climate change
as increasing ocean temperatures shift
range-limiting isotherms north
(Robinson et al. 2009). Post-nesting
females from French Guiana were found
to migrate northward toward the Gulf
Stream north wall, targeting similar
habitats in terms of physical
characteristics, i.e., strong gradients of
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SST, sea surface height, and a deep
mixed layer (Chambault et al. 2017).
Hatchling dispersal may also be affected
by changes in surface current and
thermohaline circulation patterns
(Hawkes et al. 2009; Pike 2013).
The effects of global warming are
difficult to predict, but changes in
reproductive behavior (e.g., remigration
intervals, timing and length of nesting
season) could occur (Hawkes et al. 2009;
Hamann et al. 2013). Robinson et al.
(2014) found that the median nesting
date at Sandy Point (St. Croix) occurred
on average 0.17 days earlier per year,
between 1982 and 2010. However,
Neeman et al. (2015) found that
increased temperatures at the foraging
grounds tend to delay leatherback
nesting. Temperatures at the nesting
beaches (Playa Grande, Costa Rica;
Tortuguero, Costa Rica; and St. Croix)
did not affect the timing of leatherback
nesting (Neeman et al. 2015). Because
the relation between temperatures (local
sea surface and the foraging grounds)
and timing of nesting is complex,
Neeman et al. (2015) indicated that
further study is needed at the nesting
beaches to determine how
environmental conditions change
within the season and how these
changes affect nesting success. Robinson
et al. (2014) suggests that shifts in the
nesting phenology may make the
Atlantic populations more resilient to
climate change.
Extreme precipitation events over
most of the mid-latitude and tropical
regions will very likely become more
intense and more frequent (IPCC 2014).
Changes in the frequency and timing of
storms or changes in prevailing currents
could lead to increased beach loss via
erosion (Van Houtan and Bass 2007;
Fuentes and Abbs 2010). More frequent
and intense storm events will have the
same effect on leatherback nesting
success as previously described for
natural disasters.
In summary, climate change is likely
to affect multiple life stages of turtles in
the NW Atlantic DPS. Likely impacts
include altering sex ratios and reducing
nest success, reducing nesting beach
habitat and nests due to sea level rise
and storms, and potentially changing
distribution. Climate change therefore
has the potential to alter productivity
and diversity. These impacts could be
more severe in certain areas with more
dynamic beach environments, or could
be widespread throughout the DPS.
Impacts are likely to range from small,
temporal changes in nesting season to
large losses of productivity. That said,
leatherback turtles are considered to be
the best able to cope with climate
change of all sea turtle species due to
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their wide geographic distribution and
relatively weak nesting site fidelity.
Overall, we conclude that climate
change is a threat to the NW Atlantic
DPS.
Conservation Efforts
Next we consider ‘‘conservation
efforts’’ under Section 4(b)(1)(A) (16
U.S.C. 1533(b)(1)(A)).1 There are
numerous efforts to conserve the
leatherback turtle. The following
conservation efforts apply to the NW
Atlantic DPS (for a description of each
effort, please see the section on
conservation efforts for the taxonomic
species): African Convention on the
Conservation of Nature and Natural
Resources (Algiers Convention); Central
American Regional Network;
Convention on the Conservation of
Migratory Species of Wild Animals;
Convention on Biological Diversity;
Convention on International Trade in
Endangered Species of Wild Fauna and
Flora; Convention Concerning the
Protection of the World Cultural and
Natural Heritage (World Heritage
Convention); Convention for the
Protection and Development of the
Marine Environment of the Wider
Caribbean Region, Specially Protected
Areas and Wildlife (SPAW); Convention
on the Conservation of European
Wildlife and Natural Habitats;
Convention for the Co-operation in the
Protection and Development of the
Marine and Coastal Environment of the
West and Central African Region
(Abidjan Convention); Memorandum of
Understanding Concerning
Conservation Measures for Marine
Turtles of the Atlantic Coast of Africa
(Abidjan Memorandum); Convention for
the Protection and Development of the
Marine Environment of the North East
Atlantic; Convention on Nature
Protection and Wildlife Preservation in
the Western Hemisphere (Washington or
Western Hemisphere Convention);
Convention for the Protection and
Development of the Marine
Environment of the Wider Caribbean
Region (Cartagena Convention);
Cooperative Agreement for the
Conservation of Sea Turtles of the
Caribbean Coast of Costa Rica,
Nicaragua, and Panama (Tri-Partite
Agreement); Council Regulation (EC)
No. 1239/98 of 8 June 1998 Amending
Regulation (EC) No. 894/97 Laying
Down Certain Technical Measures for
the Conservation of Fishery Measures
(Council of the European Union);
1 For a related discussion of existing regulatory
mechanisms to protect turtles, which are
considered separately under Section 4(a)(1)(D), see
the discussion above at ‘‘Inadequacy of Existing
Regulatory Mechanisms.’’
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Council Directive 92/43/EEC on the
Conservation of Natural Habitats and of
Wild Fauna and Flora (EC Habitats
Directive); Food and Agricultural
Organization (FAO) Technical
Consultation on Sea Turtle-Fishery
Interactions; Inter-American Convention
for the Protection and Conservation of
Sea Turtles (IAC); MARPOL; InterAmerican Tropical Tuna Convention
(IATTC); IUCN; North American
Agreement for Environmental
Cooperation; Protocol Concerning
Specially Protected Areas and Biological
Diversity in the Mediterranean; Ramsar
Convention on Wetlands; Regional
Fishery Management Organizations
(RFMOs); UN Convention on the Law of
the Sea (UNCLOS); and UN Resolution
44/225 on Large-Scale Pelagic Driftnet
Fishing. Although numerous
conservation efforts apply to the turtles
of this DPS, they do not adequately
reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the NW Atlantic DPS is at high risk of
extinction. The total index of nesting
female abundance is 20,659 females at
consistently monitored beaches, and the
most recent annual rate of decline is
estimated to be approximately nine
percent (NW Atlantic Leatherback
Working Group 2018). The best
available nest data reflect a steady
decline for more than a decade,
becoming more pronounced since 2008
(Eckert and Mitchell 2018; NW Atlantic
Leatherback Working Group 2018). This
decreasing trend is observed when all
available nest data are combined and at
most nesting beaches (NW Atlantic
Leatherback Working Group 2018),
including the largest nesting aggregation
in Trinidad (i.e., Grande Riviere, which
is declining at 6.9 percent annually). In
terms of productivity, the DPS exhibits
low hatching success, while other key
parameters such as clutch size,
remigration interval, and clutch
frequency are similar to species’
averages. There are also indications of
decreased productivity within the DPS
at one of the most intensively monitored
nesting beaches (i.e., Sandy Point, St.
Croix; Garner et al. 2017). The declining
region-wide nest trend and potential
changes in productivity make the DPS
highly vulnerable to threats.
However, the DPS exhibits broad
spatial distribution and some diversity.
Based upon genetic data, as well as
flipper tagging and satellite telemetry
data, this DPS shows significant spatial
structure with some connectivity among
nesting and foraging areas. Further,
nesting occurs in a variety of habitats,
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including islands and mainland, as well
as muddy, sandy, and shelly beaches.
The DPS uses multiple, distant, and
diverse foraging areas, including
oceanic and coastal waters throughout
the North Atlantic Ocean,
Mediterranean Sea, and GOM, providing
some resilience against reduced prey
availability. While the numerous and
diverse nesting and foraging locations,
along with moderate levels of genetic
diversity, provide some level of buffer to
the DPS, the highest concentrations of
nesting occur in Trinidad, French
Guiana, and Panama, where a
catastrophic event could have a
disproportionate impact on the DPS.
The primary threat to the NW Atlantic
DPS is bycatch in commercial and
artisanal, pelagic and coastal fisheries.
Gillnet fisheries, in particular those off
nesting beaches, are the greatest concern
given the high mortality rate. In
particular, the coastal surface drift
gillnet fishery off Trinidad kills an
estimated 1,000 adult leatherback turtles
annually (Lee Lum 2006; Eckert et al.
2008; Eckert 2013). Bycatch, and
subsequent mortality, in Trinidad
bottom set gillnets and surface gillnets
in Suriname and French Guiana are
major threats to the NW Atlantic DPS.
Trinidad and French Guiana host the
highest number of nesting females in
this DPS, so the continued mortality of
adults in that area is of significant
concern. Further, no adequate
regulatory mechanism is currently in
place (e.g., no gear modifications or
closures) to address this incidental
bycatch. These fisheries and the related
mortality rates have been occurring for
years (Lee Lum 2006; Eckert 2013).
Longline fisheries are the most
widespread threat, occurring throughout
the Atlantic Ocean by fisheries from
multiple nations, incidentally capturing
thousands of leatherback turtles
annually based on the best available
data. Longline gear modifications (e.g.,
circle hooks) are sometimes, but not
consistently, used. Fishery bycatch in
pot/trap gear, especially off the
northeastern U.S. coast and in Canadian
waters, and trawls are also significant
threats. Fisheries bycatch reduces
abundance by removing individuals
from the population; when those
individuals are nesting females, it
reduces productivity as well. Given the
lack of observer coverage and reporting,
cumulative mortality due to fisheries
bycatch is likely higher than available
estimates.
Additional threats to the DPS include
habitat loss, the legal and illegal harvest
of turtles and eggs, predation, vessel
strikes, pollution, climate change, oil
and gas activities, and natural disasters.
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Coastal development and armoring,
erosion (natural and anthropogenic),
and artificial lighting are some of the
most significant stressors on nesting
beach habitat, reducing nesting and
hatching success (i.e., productivity).
Habitat loss and modification is also
anticipated to increase over time with
additional development and climate
change. Legal and illegal harvest of
turtles and eggs reduces abundance and
productivity. Illegal egg poaching occurs
in several nations, particularly Costa
Rica, Dominican Republic, and
Colombia. While reduced in some
nations, illegal poaching still occurs on
unmonitored beaches throughout most
of the Caribbean, including Suriname
and Trinidad. While leatherback eggs
and hatchlings are preyed upon by
many species, the biggest threat is from
feral dogs. Egg predation by dogs occurs
in many nations, but it is a particular
concern in Colombia, French Guiana,
Guyana, Panama, Puerto Rico, and
Trinidad and Tobago. Intervention (e.g.,
screening) to reduce predation is not
used in most places, partially due to the
concern of attracting poachers as well as
the infeasibility of implementing
effective measures at high-density or
remote beaches. Egg predation reduces
productivity.
Vessel strikes are also a threat, killing
numerous leatherback turtles each year.
While exposure to vessel strikes may be
most severe in developed areas, the total
impacts are high, affecting both
abundance and productivity. Pollution,
ingestion of plastics, and entanglement
in marine debris are threats to all
leatherback turtles, most likely resulting
in injury and compromised health, and
sometimes mortality. Exposure to
pollution is widespread in the NW
Atlantic Ocean, but effect data are
limited. Oil and gas activities are threats
with the potential to grow in some
Caribbean areas. Natural disasters
(hurricanes) and phenomenon (large
Sargassum events) have an intermittent
impact to the NW Atlantic DPS. Climate
change is likely to result in reduced
productivity due to greater rates of
coastal erosion and sea level rise and
subsequent nest inundation and habitat
loss, reduced hatching success,
changing sex ratios, and distributional
changes. Although many international,
national, and local regulatory
mechanisms are in place, they do not
adequately reduce the impact of these
threats.
The cumulative impact of these
multiple threats is potentially large
(Andersen et al. 2017). Innis et al.
(2010) reported that many individuals
are simultaneously exposed to multiple
threats, including: entanglement, injury,
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plastic ingestion, adrenal gland
parasitism, diverticulitis, and burdens
of environmental toxins (Innis et al.
2010). Such cumulative pressures affect
individual survival and productivity. In
some cases, it is possible to directly link
individual threats to demographic
reductions (e.g., high mortality in
gillnets off nesting beaches reduces
nesting female abundance). More often,
however, several threats contribute
synergistically to demographic
reductions. For example, reductions in
hatching success may be caused by one
or more of the following threats alone or
in combination: erosion, poaching,
predation, climate change, and
pollution.
We find that the NW Atlantic DPS is
affected by numerous severe threats.
These present, ongoing threats injure or
kill turtles and contribute to the
declining nest trend. The Team
evaluated whether the DPS is at risk of
extinction currently or would become so
within the foreseeable future. To answer
this question, they asked how long it
would take for the total index of nesting
female abundance to be reduced by 50
percent, a drastic decline that would
reduce abundance to a level where
demographic risks would present an
independent threat to the DPS’s
continued existence, and whether this
time period places the DPS at risk
currently or within the foreseeable
future. Using estimates of the mean time
to maturation for the population
(approximately 19 years; Avens et al. in
review) and mean nesting longevity
(approximately 11 years; Avens et al. in
review) of the species, they estimated a
generation time of approximately 30
years. They then considered three
different scenarios. First, they
calculated the time until 50 percent
reduction in the total index of nesting
female abundance using data on a
significant and influential, welldocumented, threat: Gillnet bycatch
mortality of 1,000 adult turtles annually
off the largest nesting aggregation, i.e.,
Trinidad. Assuming that half of the
turtles at Trinidad killed are female,
total index of nesting female abundance
would decrease by 50 percent in 28
years, which is approximately one
generation.
Second, the Team used regional nest
trend data from the NW Atlantic
Leatherback Working Group (2018).
Using the most recent trends as is
typical for population projections (i.e.,
¥9.32 percent per year from 2008 to
2017), they found that the total index of
nesting female abundance would fall by
50 percent within 8 years (95 percent CI:
6 to 13 years). Using trends from the
longer data set (¥4.21 percent per year
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from 1990 to 2017), the total index of
nesting female abundance would fall by
50 percent within 17 years (95 percent
CI: 11 to 31 years). Finally, using their
calculation of nest trend for the highest
abundance nesting area in the DPS,
Trinidad (¥7.3 percent per year, 95
percent CI: ¥34 to 18 percent), the
Team found that the total index of
nesting female abundance would
decrease by 50 percent within 10 years
(95 percent CI: 3 years to ‘‘never;’’
however, ‘‘never’’ is highly unlikely,
given that there is a 75 percent
likelihood that the true value of the nest
trend in Trinidad is negative (f =
0.754)). There are several caveats with
using nest trend data: Adult females
typically account for, at most, a small
percentage of the population; trends in
nesting female abundance may not be an
index of the remainder of population;
stable age distribution is assumed; and
time series of available data do not
always span one generation (let alone
multiple generations required to reach
stable age distribution). Despite these
caveats, all scenarios resulted in a 50
percent reduction in the total index of
nesting female abundance in less than
one generation. While the first scenario
did not involve the use of nest trend
data, it did result in a 50 percent
reduction within one generation when
considering only one threat (albeit the
most severe), and we know that the DPS
faces many large-impact threats,
(suggesting that the first scenario
understates the DPS’s degree of risk).
For the purpose of the extinction risk
analysis, the Team discussed whether
the resulting range of time periods (8 to
28 years) suggests a present risk of
extinction or a risk of extinction within
the foreseeable future. The Team did not
have a unanimous view. All but one
Team member were present to vote on
the level of extinction risk. Eight Team
members concluded with moderate
confidence that the DPS is at high
extinction risk due to threats and the
declining trend that has accelerated in
recent years. Their confidence was
moderate rather than high due to some
resilience provided by the abundance,
spatial distribution, and diversity for
this DPS. Two Team members
concluded with low confidence that the
DPS is at moderate extinction risk. Their
confidence in this conclusion was low
due to the declining trend that has
accelerated in recent years. The Terms
of Reference called for a simple
majority, and after voting, the Team
concluded that the DPS met the
definition for high risk of extinction. We
agree with the Team’s overall
conclusion that a 50 percent decline in
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less than one generation equates to a
current, high risk of extinction. We find
support for this conclusion in well
documented examples of other
leatherback populations that have
quickly declined despite larger
abundances (e.g., the Mexico nesting
aggregation declined from 70,000
nesting females in 1982 to under 1,000
nesting females by 1994; Spotila et al.
2000).
We conclude that the NW Atlantic
DPS is presently in danger of extinction
due to the number and magnitude of
threats, of which fisheries bycatch is the
greatest concern. These present and
ongoing threats have resulted in
imminent and substantial demographic
risks (i.e., declining trends and reduced
abundance). Although numerous
conservation efforts apply to the turtles
of this DPS, they do not adequately
reduce the risk of extinction. We
conclude that the NW Atlantic DPS is in
danger of extinction throughout its
range and therefore meets the definition
of an endangered species. The
threatened species definition does not
apply because the DPS is currently at
risk of extinction (i.e., at present), rather
than on a trajectory to become so within
the foreseeable future.
SW Atlantic DPS
The Team defined the SW Atlantic
DPS as leatherback turtles originating
from the SW Atlantic Ocean, north of
47° S, east of South America, and west
of 20° W; the northern boundary is a
diagonal line between 5.377° S, 35.321°
W and 12.084620° N, 20° W. The
southern boundary is based on the
Antarctic circumpolar current which
prevents sea turtles from nesting further
south. The western end of the northern
boundary is based at the ‘‘elbow’’ of the
Brazilian coast, where the Brazilian
Current begins and likely restricts the
northern nesting range of this DPS. We
placed the eastern boundary at the 20°
W meridian as an approximate midpoint
between SW Atlantic and SE Atlantic
(i.e., turtles that nest in western Africa)
nesting beaches and to reflect both
DPS’s wide foraging range throughout
the South Atlantic Ocean. However, due
to its low abundance, the SW Atlantic
DPS is less likely to be encountered
compared to the more abundant SE
Atlantic DPS.
The SW Atlantic DPS only nests on
the southeastern coast of Brazil,
primarily in the state of Espı´rito Santo,
on a continuous stretch of beach, less
than 100 km in length, with
concentrated nesting in Povoac¸a˜o and
Comboios. While there is occasional,
limited nesting south of these primary
nesting beaches, the sand becomes
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coarser further south, and the
excavation of nests becomes more
difficult because the sand falls back into
the holes (Thome´ et al. 2007).
While nesting is limited
geographically, the overall range of this
DPS (i.e., all areas of occurrence) is
extensive, as demonstrated by
individuals tracked to numerous
foraging areas. Leatherback turtles of
this DPS use coastal waters off South
America from the ‘‘elbow’’ of Brazil
southwards to Uruguay and Argentina,
where quality foraging areas allow for
coastal foraging in addition to openocean foraging (Almeida et al. 2011).
Individuals of this DPS are also known
to migrate to the waters off western
Africa and forage in the oceanic habitat
in between South America and Africa
(Almeida et al. 2011). Likewise,
Prosdocimi et al. (2014) found 84 to 86
percent of leatherback turtles sampled
from the foraging grounds off Argentina
and Elevac¸a˜o do Rio Grande (an
elevated offshore area across from
Brazil) to originate from western African
beaches.
Abundance
The total index of nesting female
abundance for the SW Atlantic DPS is
27 females. We based this index on nest
monitoring data from Projeto TAMAR,
the Brazilian Sea Turtle Conservation
Program, which has established an
index nesting survey area along 47 km
of beach (10 km along Povoac
¸a˜o and 37
km along Comboios; IAC Brazil Annual
Report 2018), where complete daily
surveys have been conducted during the
primary nesting season from September
through March, since the 1986/1987
nesting season. Some nesting occurs
along the non-index stretches of
Povoac
¸a˜o and the beaches to the
northern part of the area, but it is minor
relative to nesting on the index survey
area (Thome´ et al. 2007). To calculate
the index of nesting female abundance
(i.e., 27 nesting females) for the Espı´rito
Santo index area, we divided the total
number of nests between the 2014/2015
and 2016/2017 nesting seasons (i.e., a 3year remigration interval; Thome´ et al.
2007) by the clutch frequency (5
clutches/season; Thome´ et al. 2007;
Tiwari et al. 2013).
Minimal, scattered nesting has been
reported on beaches outside Espı´rito
Santo (Barata and Fabiano 2002; Thome´
et al. 2007; Bezerra et al. 2014), but
these beaches exhibit suboptimal sand
characteristics for nesting, limiting the
possibility of substantial nesting
expansion into those areas (Thome´ et al.
2007). Therefore, while the nest counts
from the index beach surveys do not
provide a full estimate of all nesting for
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the DPS, they provide a high-quality
dataset, account for the majority of the
nests (approximately 80 percent;
Colman et al. 2019), and are used for
determining our index of nesting female
abundance and the nest trend in the
next section.
Our total index of nesting female
abundance is similar to the IUCN Red
List assessment’s estimate of 35 mature
individuals (female and male, assuming
a 3:1 sex ratio) based on nesting data
through 2010 (Tiwari et al. 2013).
The total index of nesting female
abundance (i.e., 27 nesting females at
the index beach) places the DPS at risk
for environmental variation, genetic
complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). These
processes, working alone or in concert,
place small populations at a greater
extinction risk than large populations,
which are better able to absorb losses in
individuals. Due to its small size, the
DPS has limited capacity to buffer such
losses. Given the intrinsic problems of
small population size, we conclude that
the nesting female abundance is a major
factor in the extinction risk of the SW
Atlantic DPS.
Productivity
The SW Atlantic DPS exhibits an
increasing, although variable nest trend.
Long-term monitoring data for this small
DPS are limited to the index nesting
survey area in southeastern Brazil,
where data has been collected between
the 1986/1987 and 2016/2017 nesting
seasons. Over the 31-year data
collection period, the mean annual
number of nests for these beaches was
35. While this is below the criterion of
50 annual nests for conducting a trend
analysis, we determined that this site
should nevertheless be included due to
the high quality and consistency of the
data, and the fact that these data
accurately represent the low level of
nesting of this DPS. The median
increase in nest counts was 4.8 percent
annually (sd = 5.8 percent; 95 percent
CI = ¥8.4 to 15.5 percent; f = 0.832;
mean annual nests = 35). As the index
area hosts the majority of known nesting
activity, these data are representative of
the entire DPS. We conclude that
nesting has increased from 1986 to
2017. Our trend estimate is similar to
that of the IUCN Red List assessment,
which characterizes the population as
increasing (Tiwari et al. 2013). It is also
in agreement with the recent study by
Colman et al. (2019), which describes
the trend as increasing but variable,
with the mean annual number of nests
increasing from 25.6 nests in the first 5
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years to 89.8 nests in the last 5 years of
monitoring (between 1988 and 2017).
While the long term trend indicates
an increase in nesting, the most recent
3 years of data (i.e., 30, 64, and 38 nests
from 2014 to 2016) show a marked
reduction in nests compared to the
previous 3 years (i.e., 78, 124, and 102
nests from 2011 to 2013). The reason for
this reduction is unknown. It could
reflect declining nesting female
abundance or changes in productivity
metrics (i.e., a longer remigration
interval or reduced clutch frequency)
related to environmental shifts or prey
availability. Therefore, there is
uncertainty regarding whether the
increasing trend will continue.
The productivity parameters for this
DPS are fairly typical for the species. In
Brazil, the average clutch size appears to
be on the lower end of the range for
Atlantic populations; conversely,
Brazilian nests tend to have a higher
average number and percentage of eggs
per clutch (Thome´ et al. 2007).
Therefore, the egg production of this
DPS appears to be weighed more
towards production of viable, hatchlingproducing eggs compared to other
Atlantic populations (Thome´ et al.
2007). Nesting females produced an
average of 3,496 hatchlings annually
over the past 10 years of nesting, which
was calculated by multiplying 60.4
nests annually, 87.7 eggs per nest, and
66.0 percent hatching success (Colman
et al. 2019). This estimate does not
include the limited nesting outside the
index area. The mean size of nesting
females (CCL) has changed from 159.8
cm, with a range of 139 to 182 cm
(Thome´ et al. 2007) to 152.9 cm ± 10.0
SD, with a range of 124.7 to 182.0 cm;
the decrease was statistically significant
and may indicate recruitment (Colman
et al. 2019). Hatching success has
increased from a mean of 65.1 percent
(with a range of 53.3 to 78 percent;
Thome´ et al. 2007) to a mean of 66
percent (with a range of 38.8 to 82.4
percent; Colman et al. 2019).
While the overall nest trend for this
DPS is increasing, there is uncertainty
regarding the continuation of this trend,
given the data for the past 3 years. The
population remains extremely small,
and thus overall productivity is limited.
Additionally, the potential for
population growth is not clear, given the
limited suitable nesting habitat
available. We conclude that limited
productivity places the DPS at risk of
extinction.
Spatial Distribution
The SW Atlantic DPS comprises a
single, small nesting aggregation
concentrated on the beaches of one state
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48363
in Brazil (Espı´rito Santo). A tagging
study has shown internesting
movements along 300 km of the coast,
including over 100 km on either side of
known nesting beaches (Almeida et al.
2011), indicating connectivity
throughout this area. The nesting spatial
distribution is extremely restricted, with
nesting constrained to a small area, with
little suitable nesting habitat into which
it can expand. Conversely, the DPS
exhibits a broad foraging range,
extending south to waters off Uruguay
and Argentina, throughout the pelagic
waters of the South Atlantic, and across
to western Africa (Almeida et al. 2011).
The wide distribution of foraging
areas likely provides some level of
buffer for the DPS against local
catastrophes or environmental changes
that could limit prey availability.
However, the limited nesting range, and
apparent lack of suitable nesting
beaches into which to expand, renders
the DPS highly susceptible to
detrimental environmental impacts,
both acute (e.g., storms and singular
events) and chronic (e.g., sea level rise
and temperature changes). Any such
change would impact the entire extent
of the DPS’s nesting habitat. With no
metapopulation structure, the DPS has
reduced capacity to withstand other
catastrophic events. Thus, despite
widely distributed foraging areas, the
extremely narrow nesting distribution
and lack of population structure
increases the extinction risk of the SW
Atlantic DPS.
Diversity
Despite its extremely low nesting
female abundance, the Brazilian nesting
aggregation has the second-highest
haplotype diversity among all Atlantic
populations (h = 0.498¥0.532; Dutton
et al. 2013; Vargas et al. 2017).
According to Thome´ et al. (2007), while
most nesting occurs from September
through March, sporadic nesting has
been recorded throughout the year,
which may provide temporal resilience
if environmental conditions limit
nesting during the primary nesting
season. The use of estuarine waters (of
the Rio de la Plata) as a year-round
foraging ground is an unusual
characteristic shared with the SE
Atlantic DPS (Lopez-Mendilaharsu et al.
2009; Prosdocimi et al. 2014). Despite
genetic and foraging diversity, the
limited size and range of the nesting
aggregation reduces the resilience of this
DPS.
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Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
Within the limited nesting range of
the SW Atlantic DPS, habitat
modification is a threat. The 2015
collapse of a tailings dam at an ore mine
upstream of the index nesting survey
area had an undetermined, but
potentially long-term, impact on the
nesting beach of the DPS. Tens of
millions of cubic meters of heavy metalladen mining waste entered the Doce
River and ultimately passed through the
mouth of the river, in the middle of the
index nesting area. Nests laid near the
river mouth were relocated to prevent
hatchlings from entering polluted
waters. Hatching success was not
significantly different between years in
the period of 2012 to 2017, which
include three seasons before (2012–
2014) and three seasons after (2015–
2017) the mining event (Colman et al.
2019). While no difference was noted in
the distribution of nests following the
dam breach, non-lethal impacts to
individuals encountering the polluted
waters, especially hatchlings, could not
be measured. Such impacts may have
occurred but may not be evident for
decades following the spill. Projeto
TAMAR is monitoring for heavy metals
in eggs and nesting females and is
closely watching for changes in fitness
and reproductive parameters (Thome´ et
al. 2017). As a result of the dam’s
collapse, the Brazilian Federal
government is implementing a marine
protected area (APA-Area de Protecao
Ambiental da Foz do Rio Doce),
including about 100 kilometers of
coastline, which should encompass the
entire extension of the index nesting
beaches, with both coastline and
surrounding marine areas. Such a
measure is an environmental
compensation for the dam’s collapse,
and should be implemented with
specific resources in the coming years
(ICMBio, MMA, Brazil; J. Thome´,
Projeto TAMAR, pers. comm., 2019).
Beach erosion and tidal flooding are
also threats to this DPS. According to
Thome´ et al. (2007), occasional
relocation of nests and nest protection
occur when inundation or predation
risk is considered high. The majority of
nests are relocated when in danger of
beach erosion or tidal flooding (J.
Thome´, Projeto TAMAR, pers. comm.,
2019).
Although coastal light pollution has
been documented to be increasing in
Brazil, nesting has not been notably
impacted thus far (Colman et al. 2018).
The lack of impact may be attributable
to conservation strategies including the
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creation of protected areas and
minimization of direct lighting on the
nesting beaches. Nests are relocated
from heavily lit areas. All light sources
with a light intensity greater than 0 lux
(lux = lumen per m2) on these beaches
are prohibited by a Federal ordinance
(Portaria IBAMA 11/1995).
Construction, lighting, and poaching
were not considered a significant
problem at the leatherback nesting
beaches by Thome´ et al. (2007).
However, such problems persist in
several other turtle nesting beaches in
Brazil (Mascarenhas et al. 2004; Lara et
al. 2016). More recently, coastal
development and artificial lighting have
been identified as potential threats for
leatherback turtles on the beaches of
Espı´rito Santo (TAMAR/Unpublished
data) and further research is needed to
better understand these threats. Nests
are relocated from heavily lit areas.
Colman et al. (2018) found a negative
relationship between nest density and
light levels. Additionally, as oil industry
and other economic developments are
explored, the potential threat to the
nesting habitat may increase (Thome´ et
al. 2007).
A significant portion of the nesting
beach is protected as a Federal reserve
under Brazilian Decree no. 90222
(September, 25 1984), which covers 15
km of Comboios Beach, south of the
mouth of the Doce River. An additional
22 km, south of the reserve, falls within
indigenous land that has restricted
access under Federal law. No Federally
protected areas exist north of the Doce
River mouth, where Povoac¸a˜o Beach
occurs. However, local, state, and
Federal regulations provide some
coastal zone protections in that area.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization poses a threat to the
SW Atlantic DPS. Though specific
information on leatherback turtle
harvests is not available, there was
historically traditional harvest of sea
turtles and eggs in Espı´rito Santo (Hartt
1941; Medeiros 1983). This harvest,
however, has been largely curtailed
through the work of Projeto TAMAR,
which promoted other economic
activities and hired ex-turtle hunters to
protect nests (Marcovaldi et al. 2005;
Almeida and Mendes 2007). The
capture of leatherback turtles was
banned in Brazil in 1968, and full
protection for all sea turtles was enacted
in 1986 (Marcovaldi and Marcovaldi
1999). At present, egg poaching has
been reduced, and there is no known
subsistence hunting for sea turtles of
any species (Thome´ et al. 2007). As
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previously noted, there is protection for
or limited access to much of the nesting
habitat south of the Doce River.
However, this protection does not
extend north of the river, where
additional nesting occurs. Because of
the very small size of the population,
even very low levels of egg poaching
have the potential to impact the
population. Therefore, we conclude that
overutilization poses a threat to the SW
Atlantic DPS.
Disease or Predation
While we could not find any
information on disease for this DPS,
predation is a threat to the SW Atlantic
DPS. Invertebrates, reptiles, and
mammals prey on eggs, while hatchlings
fall prey to land, air, and marine
predators. According to Thome´ et al.
(2007), relocation and protection of
nests may be undertaken when
inundation (primarily) or predation
(secondarily) risk is considered high (J.
Thome´, Projeto TAMAR, pers. comm.,
2019). Predators include foxes
(Cerdocyon thous), raccoons (Procyon
cancrivorus), and domestic dogs,
although there are no quantitative
estimates of predation for this DPS (J.
Thome´, Projeto TAMAR, pers. comm.,
2019). Some predation of large juveniles
and adults occurs in the marine
environment, especially by sharks
(Bornatowski et al. 2012), but the
frequency and impact on those
populations is not well understood. For
this DPS, predation primarily impacts
productivity (i.e., reduced egg and
hatching success). We conclude that
predation is a threat to the SW Atlantic
DPS, but that there is insufficient
information to classify disease as a
threat.
Inadequacy of Existing Regulatory
Mechanisms
The SW Atlantic DPS is protected by
several regulatory mechanisms. For
each, the Team reviewed the objectives
of the regulation and to what extent it
adequately addresses the targeted threat.
Beach habitat is protected throughout
much of the nesting range of this DPS.
The vast majority of nesting occurs in
Espı´rito Santo, where beaches have been
protected since 1982. All light sources
with a light intensity greater than 0 lux
(lux = lumen per m2) on these beaches
are prohibited by a Federal ordinance
(Portaria IBAMA 11/1995).
The take of leatherback turtles is
illegal throughout the SW Atlantic
Ocean. Regional regulations include:
Brazil Portaria, Manter proibida a
captura de tartarugas marinhas das
espe´cies Caretta, Dermochelys coriacea,
Eretmochelys imbricata e Lepidochelys
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olivacea 2 No.27/1982; Uruguay
Presidential Decree 144 and additional
legislation to reduce bycatch and
prevent habitat alteration, and to
prevent the removal of individuals from
their natural environment; Argentina
National Decree 666 from 1997; and
various laws prohibiting hunting and
selling sea turtles. Harvest and
consumption of sea turtles are illegal
under Brazilian law (Law on
Environmental Crimes N° 9605/1998).
While these protections are mostly
effective, very low levels of egg
poaching still exist (Thome´ et al. 2007).
Fisheries bycatch is the primary threat
to the SW Atlantic DPS. Although
regulations address this issue to some
extent, they do not do so adequately and
it continues to be a threat. In 2001,
Brazil established the National Plan for
the Reduction of Incidental Capture of
Sea Turtles in Fishing Activities
(Marcovaldi et al. 2005). However,
bycatch continues to be a major
problem. In Brazil, the use of TEDs in
trawl fisheries is mandatory (Instruc
¸a˜o
Normativa MMA No. 31; December 13,
2004), but most fishermen nevertheless
do not use such gear, and there is little
or no enforcement by authorities (IAC
Brazil Annual Report 2018). The UN
established a worldwide moratorium on
drift gillnet fishing effective in 1992, the
General Fisheries Commission for the
Mediterranean prohibited driftnet
fishing in 1997, and the International
Commission for the Conservation of
Atlantic Tunas (ICCAT) banned
driftnets in 2003. Despite these and
other numerous regulations and
international instruments to protect sea
turtles, significant bycatch still occurs
in artisanal and commercial fisheries
operating in the territorial waters of
Argentina, Uruguay, and Brazil and on
the high seas (Gonza´lez et al. 2012).
In summary, while numerous
regulatory mechanisms have been
enacted to provide some protections to
leatherback turtles, their eggs, and
nesting habitat throughout the range of
this DPS, they have been inadequate.
Many do not effectively reduce the
threat that they were designed to
address, generally as a result of limited
implementation or enforcement.
Fisheries bycatch, in particular, remains
a major threat to the DPS despite
regulatory mechanisms. We conclude
that the failure to implement and
enforce effective regulations is a threat
to the DPS.
2 Prohibition of the capture of sea turtles of the
species Caretta caretta, Dermochelys coriacea,
Eretmochelys imbricata, and Lepidochelys olivacea.
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Fisheries Bycatch
Fisheries bycatch is the primary threat
to the SW Atlantic DPS. Leatherback
turtles are captured as bycatch in
commercial and artisanal fisheries,
along coastal foraging and breeding
areas, and on the high seas. The
extensive foraging range of this DPS
makes it vulnerable to interactions with
fisheries off the coasts of Brazil,
Uruguay, and Argentina, in the pelagic
waters of the South Atlantic Ocean, and
along the coastal waters off western
Africa. Recoveries of females tagged in
Espı´rito Santo are scarce, however.
Three were found dead on the Brazilian
coast (incidentally captured in fisheries
around the Doce River mouth (TAMAR,
unpublished data)), one in Argentina
(Alvarez et al. 2009), and one in
Namibia, West Africa (Almeida et al.
2014). Fisheries interaction information
specific to this DPS is limited, because
the data do not differentiate among
individuals from this DPS and SE
Atlantic individuals that forage within
the same range. Because the SE Atlantic
DPS is much more abundant than the
SW Atlantic DPS, most fishery
interactions likely involve SE Atlantic
individuals. However, data about
bycatch involving the SE Atlantic DPS
is informative because the impact to the
SW Atlantic DPS individuals is likely to
be proportional to their relative
presence in the area. Bycatch in gillnets;
surface, deep-water longline hooks; and
trawls are the principal causes of sea
turtle deaths, with not only higher
interaction numbers, but higher
mortality rates than other fishery
interactions (Kotas et al. 2004; Pinedo
and Polacheck 2004; Tudela et al. 2005;
Giffoni et al. 2013).
Coastal gillnet fisheries interactions
are one of the largest threats to the
survival of the SW Atlantic DPS. In an
analysis of Brazilian fishery data from
1990 to 2012, Giffoni et al. (2013)
documented 237 leatherback turtle
interactions, and 31 percent mortality,
in coastal set, fixed, encircling, and
pelagic drift gillnets. The actual number
of interactions is likely substantially
higher, as many interactions go
unreported.
Smaller scale artisanal gillnet
fisheries occur in coastal waters that are
used by SW Atlantic individuals for
mating, access to nesting beaches, and
foraging. Thome´ et al. (2007) described
the occurrence of artisanal gillnet
fisheries close to the nesting beach but
indicated that Brazil was investing
resources in developing lower-impact
fishing techniques. However, such
fisheries occur throughout other
important coastal foraging areas off
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South America. Additionally, coastal
artisanal gillnet fishery interactions
with leatherback turtles are known to
occur off the western coast of Africa,
where some individuals from the SW
Atlantic DPS forage (Riskas and Tiwari
2013). The Rio de la Plata estuary, an
important foraging area off Uruguay, has
numerous documented instances of
leatherback turtle entanglements,
including mortalities from coastal
bottom-set gillnet fisheries (Fallabrino et
al. 2006; Lopez-Mendilaharsu et al.
2009; Velez-Rubio et al. 2013).
Larger-scale commercial ocean gillnet
fisheries are also a significant threat for
the SW Atlantic DPS, with high bycatch
rates reported off Brazil in drift and set
gillnets (Fiedler et al. 2012; Ramos and
Vasconcellos 2013). Drift gillnet fishing
off Brazil started in 1986, targeting
hammerhead sharks (Domingo et al.
2006). Marcovaldi et al. (2006) reported
that leatherback turtles comprised about
70 percent of all sea turtles captured in
Brazilian driftnet shark fisheries. From
2002 to 2008, 351 sea turtles were
incidentally caught in 41 fishing trips
and 371 sets. Leatherback turtles
accounted for 77.3 percent of the take (n
= 252 turtles, capture rate = 0.1405
turtles/km of net) with 22.2 to 29.4
percent of turtles dead upon retrieval
and no estimate of post-release mortality
for those released alive. The annual
catch by this fishery ranged from 1,212
to 6,160 leatherback turtles, as estimated
based on bootstrap procedures under
different fishing effort scenarios in the
1990s (Fiedler et al. 2012). In 1998, a
Brazilian Federal ordinance limited the
use and transport of bottom and drift
gillnets over 2.5 km long. Such
regulations were difficult to enforce,
and vessels from the ports of Itajaı´,
Navegantes and Porto Belo, Santa
Catarina, Brazil, deployed nets up to
7,846 m long between 2005 and 2006
(Kotas et al. 2008). In 2010 the
ordinance was suspended, permitting
unrestricted fishing with driftnets
(Fiedler 2012). The shark drift gillnet
fishery declined steeply in later years,
with no vessels operating in 2009
(UNIVALI/CTTMar 2010) likely because
of target species reduction, reduced
profitability, and IBAMA Normative
Instruction N166/2007 which
temporarily stopped the issuance of new
driftnet fishing licenses and established
a 2-year deadline by which vessels were
to replace driftnets with other gear.
Various other gillnet fisheries, such as
bottom gillnets for sharks and mollusks,
have reported leatherback mortalities as
well, such as that occurring off Uruguay
(Fallabrino et al. 2006; Laporta et al.
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2006; Eckert et al. 2009) and the western
coast of Africa (Riskas and Tiwari 2013).
Longline fisheries pose a significant
threat to the SW Atlantic DPS, as the
spatio-temporal distribution of
leatherback turtles overlaps with
longline fishing effort (Fossette et al.
2014). In a review of reported, observed
takes in hook and line fishery (primarily
longline) interactions with leatherback
turtles in all of Brazil from 1990 to 2012,
1061 takes were documented, with 3
percent of the taken turtles found dead
on the line and another 37.5 percent of
unknown condition after release
(Giffoni et al. 2013). High frequencies of
leatherback deaths from bycatch have
been documented on longline fishing
vessels from southern Brazil and
Uruguay (Kotas et al. 2004; Pinedo and
Polacheck 2004; Domingo et al. 2006;
Giffoni et al. 2008; Monteiro 2008).
Between 2004 and 2005, in a study off
southern Brazil, eight leatherback turtles
were captured, with a mean capture rate
of 0.03 turtles per 1,000 hooks
(Monteiro 2008). In 1999, there were 70
longliners in the Brazilian fleet, with 33
vessels operating out of southern Brazil
and fishing a total of 13,598,260 hooks
(ICCAT 2001). However, the overall
effort in the area was much higher, as
longliners from Uruguay, Chile, Japan,
Taiwan, and Spain fish in this area
(Folsom 1997; Weidner and Arocha
1999; Weidner et al. 1999). Scientific
observers documenting 10 trips by
longline vessels from the Uruguayan
fleet operating in the SW Atlantic Ocean
between 26° and 37° S between April
1998 and November 2000 observed 27
incidentally caught leatherback turtles
(Balestre et al. 2003). The prevalence of
leatherback interactions in pelagic
longline fisheries is likely a result of the
longline fleet fishing the productive
areas in the convergence zone of the
Brazilian Current and the cold waters
from the Falklands Current (Kotas et al.
2004), which coincides with important
sea turtle foraging and developmental
habitat. As with gillnets, the scope of
the longline threat to the SW Atlantic
DPS spans across the South Atlantic
Ocean in both coastal and oceanic
waters. In addition to exposure to
longline fisheries off South America,
coastal longline fisheries off Cameroon,
Angola, and Namibia, and pelagic
longlines in the Gulf of Guinea and the
eastern portion of the South Atlantic
Ocean have also been documented to
take leatherback turtles (Honig et al.
2007; Riskas and Tiwari 2013; Angel et
al. 2014; Huang 2015; Gray and Diaz
2017). Additional evidence of longline
interactions comes from the stranding
data, where flipper injuries on some of
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the stranded leatherback turtles could
have been caused by interactions with
pelagic longlines. Onboard observers in
longline fisheries off Brazil have
reported that leatherback turtles tend to
be foul-hooked in the flipper rather than
the mouth (Kotas et al. 2004; Pinedo and
Polacheck 2004; Lima 2007). In 2017,
Brazil enacted a law (PORTARIA
INTERMINISTERIAL No 74, DE 1o- DE
NOVEMBRO DE 2017) requiring the use
of circle hooks in the pelagic longline
fisheries as well as keeping specified
dehooking and gear removal equipment
on board any Brazilian longline vessel.
Specifically, the Brazilian government
required the use of 14/0 or larger circle
hooks for all longline vessels targeting
swordfish or tuna (https://
www.jusbrasil.com.br/diarios/
166677996/dou-secao-1-06-11-2017-pg81).
Trawl fisheries also impact the SW
Atlantic DPS, mainly along coastal
waters off southern Brazil, Argentina,
and Uruguay (Gonzalez Carman et al.
2011; Velez Rubio et al. 2013; Monteiro
et al. 2016). Although there are fewer
interactions with trawl fisheries relative
to other fisheries (i.e., gillnet and
longline fisheries), mortality rates in
trawl fisheries are far higher (Miller et
al. 2006; Laporta et al. 2013).
Observation of the Uruguayan bottom
trawl fishery, during a tagging and data
collection program designed to increase
the understanding of the fishery impacts
on sea turtles, documented 17
leatherback interactions from April 2002
to June 2005 (Laporta et al. 2013).
Coastal bottom trawl and artisanal
gillnet fisheries were the main causes of
death of leatherbacks found stranded in
Uruguay (Velez Rubio et al. 2013).
Recorded interactions in coastal trawl
fisheries are also known from Gabon,
Congo, and Namibia (Riskas and Tiwari
2013).
Other fisheries such as corrals, pound
nets, and pots appear to present a much
lower risk for leatherback turtles than to
other sea turtle species. From 1990 to
2012, Giffoni et al. (2013) documented
only two leatherback turtles (both alive)
of the 8,367 total sea turtles taken in
those fisheries.
While specific information is not
available to permit calculating an
estimate of overall bycatch and
mortality rates of SW Atlantic
leatherback turtles, it is clear that
fisheries bycatch, especially in gillnets
and longlines, is a major threat to the
DPS. Immature and adult individuals
are exposed to high fishing effort
throughout their foraging range and in
coastal waters near nesting beaches.
Bycatch mortality is also high, with
reported rates of up to 31 percent
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(Giffoni et al. 2013). Mortality reduces
abundance, by removing individuals
from the population; it also reduces
productivity, when nesting females are
incidentally captured and killed. Given
the small size of the DPS, the loss of
even a small number of individuals
from fishery interactions has the
potential to reduce abundance and
productivity. Therefore, we conclude
that fisheries bycatch is the primary
threat to the SW Atlantic DPS.
Vessel Strikes
There is little information regarding
vessel strikes for the SW Atlantic DPS.
Many of the primary foraging areas for
this DPS off the coasts of Argentina,
Uruguay, and Brazil are experiencing
increased vessel traffic from fishing
vessels, cargo transport, and tourism
(Lo´pez-Mendilaharsu et al. 2009;
Fossette et al. 2014), so leatherback
turtle interactions with vessels may
occur. Affected individuals likely
include immature and mature turtles.
Impacts range from injury to mortality.
We conclude from the best available
information that vessel strikes are likely
a threat to the DPS.
Pollution
As with all leatherback turtles,
entanglement in and ingestion of marine
debris and plastics is a threat that likely
kills several individuals a year. Multiple
studies have implicated the ingestion of
marine debris and/or entanglement in
cases of injury or death of turtles found
in waters occupied by the SW Atlantic
DPS (Bugoni et al. 2001; Eckert et al.
2009; Schulyer et al. 2013; Scherer et al.
2014). However, no individuals were
assigned to a particular population and
could have been members of the more
abundant SE Atlantic DPS, which is
known to occupy the same waters.
While there is no specific information
on effects to leatherback turtles of this
DPS, pollution from various economic
activities including maritime transport,
tourism, and domestic and industrial
waste discharges that are known to
occur within their range, may also have
an impact (Lo´pez-Mendilaharsu et al.
2009; Fossette et al. 2014). Events such
as the failure of a mining tailings dam
in 2015 that resulted in the discharge of
tons of mining mud contaminated with
heavy metals into the Doce River, and
subsequently into the waters off Espı´rito
Santo nesting beaches, are also a
concern, though no specific impacts to
leatherback turtles have so far been
noted from that event (Garcia et al.
2017). There is also concern about the
potential for increased oil and gas
exploration activities (Thome´ et al.
2007). The petroleum industry in Brazil
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has implemented a beach monitoring
program, along large stretches of the
Brazilian coast, including Espı´rito
Santo, to monitor for potential impacts
to sea turtles and their nesting beaches
from industry activities (Werneck et al.
2018)
Assigning impacts of pollution
specifically to individuals within the
SW Atlantic DPS is difficult, and the
best available information does not
quantify such impacts. However, given
its prevalence, we conclude that
pollution poses a threat to the DPS.
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Climate Change
Climate change poses a threat to the
SW Atlantic DPS. The impacts of
climate change include: Increases in
temperatures (air, sand, and sea
surface); sea level rise; increased coastal
erosion; more frequent and intense
storm events; and changes in ocean
currents.
Because leatherback turtles nest lower
on the beach than other sea turtles, their
eggs are more at risk of being exposed
and destroyed by increases in sea level
and coastal erosion (Boyes et al. 2010).
Additionally, given the limited
availability of suitable nesting habitat,
the loss of the current nesting habitat
with no buffer area to move into would
pose a major problem for the DPS. Thus,
rising sea level and beach erosion are
potential threats to the DPS.
While we do not have specific
information on pivotal temperatures and
temperature thresholds for egg mortality
for this DPS, sand temperatures
influence egg viability and sex
determination. Given the potential lack
of suitable nesting habitat outside the
area currently being utilized, there is
little opportunity for a spatial shift in
nesting in response to changing
temperatures. This DPS exhibits some
year-round nesting, which provides a
small measure of resilience to
counteract increasing temperatures.
However, it is not likely to be sufficient
to make up for the loss of nesting habitat
and opportunity resulting from sea level
rise and temperature increases. The
impacts on productivity and
survivorship for such shifts in nesting
are unknown.
The threat of climate change is likely
to modify the nesting conditions for the
DPS. Adverse impacts on turtles of the
SW Atlantic DPS would be inescapable
because the entire DPS is confined to a
limited nesting area. Impacts are likely
to range from small, temporal changes
in nesting season to large losses of
productivity. Therefore, we conclude
that climate change is a threat to the
DPS.
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Channel Dredging
There is evidence of interactions with
hopper dredges associated with channel
dredging and maintenance. Between
2008 and 2014, four leatherback turtles
were killed by hopper dredges in Rio de
Janeiro (Goldberg et al. 2015).
Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply
turtles of the SW Atlantic DPS (for a
description of each effort, please see the
section on conservation efforts for the
overall species): Southwest Atlantic Sea
Turtle Network, Convention on the
Conservation of Migratory Species of
Wild Animals, Convention on Biological
Diversity, Convention on International
Trade in Endangered Species of Wild
Fauna and Flora, Convention
Concerning the Protection of the World
Cultural and Natural Heritage (World
Heritage Convention), FAO Technical
Consultation on Sea Turtle-Fishery
Interactions, IAC, MARPOL, IUCN,
Ramsar Convention on Wetlands,
RFMOs, South Atlantic Association,
UNCLOS, and UN Resolution 44/225 on
Large-Scale Pelagic Driftnet Fishing.
Although numerous conservation efforts
apply to the turtles of this DPS, they do
not adequately reduce its risk of
extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the SW Atlantic DPS is at ‘‘high’’ risk of
extinction. The DPS exhibits a total
index of nesting female abundance of 27
females at the index beach. Such a
nesting population size places this DPS
at risk of stochastic or catastrophic
events that increase its extinction risk.
Although there has been substantial
variability in nesting at the index
nesting beach since 1986, the nest trend
shows a strong, nearly five percent
annual increase through 2017, with the
largest increase occurring in the past
decade. However, nesting has declined
in the past 3 years. There is only one
nesting aggregation, limited to a
relatively small stretch (47 km) of beach
along a single coast. Some nesting also
occurs outside that area, but is mostly
sporadic and limited by sand and
temperatures unsuited for nesting. Thus,
stochastic events have the potential to
have catastrophic effects on the entire
DPS, with no distant subpopulations
serving as a buffer or source of
additional individuals or diversity.
Based on these factors, we find the DPS
to be at risk of extinction as a result of
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its limited abundance, spatial structure,
and resilience.
Current threats place this DPS at
further risk of extinction. The primary
threat to this DPS is bycatch in
commercial and artisanal, pelagic and
coastal fisheries, especially gillnet and
longline fisheries. Fisheries bycatch
reduces abundance by removing
individuals from the population.
Because several fisheries operate near
nesting beaches, productivity is also
reduced when nesting females are
prevented from returning to nesting
beaches. Exposure to and impact of this
threat are high. Additional threats
include: Habitat modification,
overutilization, predation, pollution,
vessel strikes, and climate change.
Habitat modification includes incidents
such as the mining dam breach
upstream of the Doce River, which flows
into the ocean through the middle of the
primary nesting beach. Overutilization
and predation are threats for this DPS as
well, though some protective measures
exist. While many laws are in place to
protect sea turtles from fishery impacts,
the continued impact of bycatch
indicates that regulatory mechanisms
are inadequate to sufficiently address
the threat. Pollution and vessel strikes
are potentially increasing threats to the
DPS. Climate change is another threat
that is likely to increase, resulting in
reduced productivity due to greater
rates of coastal erosion and nest
inundation, and in some areas, nest
failure or skewed sex ratios due to
increased sand temperatures.
We conclude, consistent with the
Team’s findings, that the SW Atlantic
DPS is currently in danger of extinction.
The total index of nesting female
abundance make the DPS highly
vulnerable to threats despite the
apparent increasing nesting trend. In
addition, this DPS consists of only one
small nesting aggregation with limited
potential nesting beaches to the north
and south for expansion. The limited
nesting range and small size makes the
DPS highly vulnerable to stochastic
impacts in the natural environment as
well as singular, large-scale,
anthropogenic events such as oil spills.
Some degree of resilience is provided by
the use of multiple foraging areas across
a vast geographic area. However, that
expansive foraging range also exposes
the DPS to numerous fisheries (which
are coastal and on the high seas,
artisanal and commercial, off both
South America and western Africa),
making fisheries bycatch by far the
biggest threat to the DPS. Although
numerous conservation efforts apply to
the turtles of this DPS, they do not
adequately reduce the risk of extinction.
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We conclude that the SW Atlantic DPS
is currently in danger of extinction
throughout its range and thus meets the
definition of an endangered species. The
threatened species definition does not
apply because the DPS is at risk of
extinction now (i.e., at present), rather
than on a trajectory to become so within
the foreseeable future.
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SE Atlantic DPS
The Team defined the SE Atlantic
DPS as leatherback turtles originating
from the SE Atlantic Ocean, north of
47° S, east of 20° W, and west of 20° E;
the NW boundary is a diagonal line
between 12.084620° N, 20° W and
16.063° N, 16.51° W. The eastern
boundary occurs at the southern tip of
Africa, where the Agulhas and Benguela
Currents meet. Along with the cold
waters of the Antarctic Circumpolar
Current, these currents likely restrict the
nesting range of this DPS. We placed the
western boundary at the 20° W meridian
as an approximate midpoint between SE
Atlantic and SW Atlantic (i.e., turtles
that nest in Brazil) nesting beaches and
to reflect the DPS’s wide foraging range
throughout the South Atlantic Ocean;
this DPS is more likely to be
encountered in these waters compared
to individuals from the less abundant
SW Atlantic DPS. The northern
boundary is a diagonal line between the
elbow of Brazil and the northern
boundary of Senegal because the SE
Atlantic DPS does not appear to nest
above this boundary (Fretey et al. 2007).
The range of the SE Atlantic DPS is
extensive, mirroring that of the SW
Atlantic DPS. While nesting occurs
along the western coast of Africa, data
indicate that foraging areas and
migratory paths stretch along the
Atlantic coast of Africa from Senegal to
South Africa, across the South Atlantic
Ocean, and into the coastal waters of
Brazil, Uruguay, and Argentina. As with
the SW Atlantic DPS, this DPS does not
appear to forage in northern latitudes.
All nesting for the SE Atlantic DPS
occurs along the Atlantic coast of
western Africa, from Senegal to Angola,
a nesting range of over 7,500 km.
However, the vast majority of nesting
occurs in Gabon, Equatorial Guinea
(including Bioko Island), and the
Republic of Congo (TEWG 2007; Fretey
et al. 2007, Witt et al. 2009; Tiwari et
al. 2013). Gabon may have once hosted
the largest nesting aggregation in the
world when it was discovered in the
early 2000s (Witt et al. 2009), but
current data indicate much lower levels
of nesting (Formia et al. in prep)
compared to those described in Witt et
al. (2009).
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While nesting occurs along the
western coast of Africa, foraging
grounds and migratory paths stretch
across the South Atlantic Ocean to the
coastal waters of Brazil, Uruguay, and
Argentina. Because of the greater
abundance of this DPS, most
individuals found in the western South
Atlantic along the coast of South
America, and on the high seas, belong
to the SE Atlantic DPS. Prosdocimi et al.
(2014) found 84 to 86 percent of
leatherback turtles sampled from the
foraging grounds off Argentina and
Elevac¸a˜o do Rio Grande (an elevated
offshore area across from Brazil) to
originate from western African beaches.
Abundance
The total index of nesting female
abundance for the SE Atlantic DPS is
9,198 females. We based this total index
on nine nesting aggregations in Gabon
(n = 8,495 nesting females), Equatorial
Guinea (n = 457), Republic of Congo (n
= 69), Sierra Leone (n = 39), Liberia (n
= 45), Ivory Coast (n = 40), Ghana (n =
4), Cameroon (n = 3), and Sao Tome and
Principe (n = 46). Our total index does
not include 10 unquantified nesting
aggregations in Guinea-Bissau, Angola,
and other nations. For more information
on data sources and calculations, please
see the Status Review Report.
Our total index of nesting female
abundance is an index because we do
not have consistent data from much of
the nesting range of the DPS, which
extends from Senegal to Angola.
However, the largest nesting
aggregations occur in Gabon, Equatorial
Guinea (including Bioko Island), and
the Republic of Congo (TEWG 2007;
Fretey et al. 2007; Witt et al. 2009;
Tiwari et al. 2013), which are
represented in our total index. The
IUCN Red List assessment did not
provide an estimate of population size
but instead concluded that the
subpopulation was ‘‘data deficient’’
(Tiwari et al. 2013).
To calculate the index of nesting
female abundance in Gabon, where
annual aerial surveys of 600 km of
nesting beaches gather emergence data,
we used a remigration interval of 3
years, a clutch frequency of 7.8 clutches
per season per female, and estimated
that 95 percent of emergences resulted
in nesting (Formia et al. in prep). Our
index of nesting female abundance for
Gabon (i.e., 8,495 nesting females) is
lower than previous estimates.
According to Witt et al. (2009), Gabon
once hosted the largest leatherback
nesting aggregation in the world, with
an estimated 36,185 to 126,480 clutches
per year (approximately 15,730 to
41,373 nesting females). These estimates
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were based on a combination of aerial
surveys and ground-truthing surveys,
conducted during the 2002/2003, 2005/
2006, and 2006/2007 nesting seasons.
More recent aerial surveys indicate a
steep decline in nesting since the early
2000s, with a high of 108,588 estimated
nests in 2002/03, a low of 4,275
estimated nests in 2009/10, and fewer
than 25,000 nests in the final year of
available data (2015/16; Formia et al. in
prep).
Nesting is scattered on continental
Equatorial Guinea (Fretey 2001), but it
occurs on several beaches of Bioko
Island and is monitored at the Gran
Caldera Scientific Reserve (n = 457
nesting females, based on body pit data
from the 2000/2001 through 2017/2018
nesting seasons; D. Venditti et al.,
Drexel University, pers. comm., 2018).
Rader et al. (2006) documented an
average of 3,896 nests annually between
the 2000/2001 to 2004/2005 nesting
seasons, which equates to
approximately 2,338 nesting females
(i.e., using a 3-year remigration interval
and a clutch frequency of 5 nests
annually). Based on the data available
on nesting in the Republic of Congo
from the 2003/2004 to 2016/2017
nesting seasons (N. Breheret, SWOT,
pers. comm., 2018), we estimated 69
nesting females. In an analysis of older
data (1999 to 2008), Girard et al. (2016)
estimated 933 nests per year on the
monitored beaches, which equates to
approximately 560 nesting females.
In Guinea-Bissau, only one beach is
monitored regularly, in Orango National
Park, Bijagos Archipelago, where
occasional leatherback nesting tracks are
recorded. Each season, a few nests are
reported elsewhere throughout the
nation (Barbosa et al. 1998; Fretey et al.
2007).
In the Ivory Coast (n = 40 nesting
females), Gomez (2005) counted 218
nests over 41 km of beach in February
2001. Pen˜ate et al. (2007) reported 189
nests reported from non-exhaustive
surveys of 27 km of coastline during the
2001/2002 nesting season.
In Ghana, nest monitoring occurs on
three beaches: Mankoadze (n = 4 nesting
females), Ada, and Keta. We were
unable to calculate the index for Ada
and Keta beaches because we only
received information on nest averages.
From 2000 to 2017, an annual average
of 34 nests were observed on Ada Beach
(D. Agyeman, pers. comm., 2018).
During the 2006/2007 nesting season,
481 leatherback nests were counted on
Ada Beach (Allman and Armah 2010).
Over an unspecified time frame, an
annual average of 80 nests were
observed on Keta Beach (A. Fuseini,
pers. comm., 2018).
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In Cameroon (n = 3 nesting females;
Fretey and Nibam unpublished data
2018), Girard et al. (2016) estimated an
average of 43 leatherback nests
annually, which would equate to 26
nesting females, from 1999 to 2008. In
Sa˜o Tome´ and Principe (n = 46 nesting
females), Girard et al. (2016) estimated
an average of 78 nests annually from
1999 to 2008, which is similar to our
estimate.
Nesting occurs on other beaches
throughout western Africa. However,
recent consistent and standardized
monitoring data are not available.
Sporadic nesting occurs in Senegal
(Maigret 1978; Dupuy 1986), Republic of
The Gambia (Barnett et al. 2004,
Hawkes et al. 2006), Togo (Segniagbeto
2004), Nigeria (Fretey 2001; Mojisola et
al. 2015), Democratic Republic of
Congo, (OCPE-ONG 2006), and Angola
(Carr and Carr 1991; Weir et al. 2007).
The total index of nesting female
abundance of the SE Atlantic DPS
(9,198 females) does not reduce the risk
for environmental variation, genetic
complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). Such
abundance provides little resilience to
buffer losses of individuals. We
conclude that the nesting female
abundance, as estimated, does not
reduce the extinction risk of this DPS.
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Productivity
Based on data collected from the
largest nesting aggregation (i.e., Gabon),
the SE Atlantic DPS exhibits a declining
nesting trend. Data collected between
the 2002/2003 and 2015/2016 nesting
seasons (with two years of missing data)
indicated a median trend in nesting
activity of ¥8.6 percent annually (sd =
21.9 percent; 95 percent CI = –52.6 to
36.9 percent; f = 0.676; mean annual
nesting activities = 35,204). The trend in
Gabon is likely representative of the
entire DPS, because the majority of
nesting occurs there. Additional nest
trend data are available from the Gran
Caldera Scientific Reserve of Bioko
Island, where the number of body pits
increased 2.8 percent annually (sd =
15.6 percent; 95 percent CI = –27.2 to
36.0 percent) from 1996/1997 to 2017/
2018.
Regarding productivity parameters,
available information is often from a
limited area and may not be
representative of the entire DPS.
However, based on available data, the
size of nesting females, clutch size,
hatching success, and incubation period
appear to be similar to the species’
averages. We conclude that the
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declining nesting trend contributes to
the extinction risk of this DPS.
Spatial Distribution
The SE Atlantic DPS has a broad
spatial distribution. The nesting range is
centered on Gabon, with nesting
occurring from Senegal to Angola.
Genetic data available for Gabon and
Ghana indicate significant genetic
differentiation based on mtDNA data,
but weak differentiation based on
analysis of nuclear DNA, likely
indicating demographically
independent subpopulations connected
by limited gene flow (Dutton et al.
2013).
In addition to the extensive nesting
range, this DPS also has an expansive
foraging and migratory range, from the
coastal waters of Atlantic Africa, across
the pelagic waters of the South Atlantic,
and along the South American coast
from Brazil to Argentina. While nesting
along the coast of Africa extends only to
Angola, recent tag returns and satellite
telemetry indicate that turtles utilize the
waters in Namibia as well (Almeida et
al. 2014). Transatlantic movements were
first recorded from tag returns of four
leatherback turtles tagged on the nesting
beaches of Gabon and recaptured in the
waters of Argentina and Brazil (Billes et
al. 2006). Satellite telemetry confirmed
that nesting females from Gabon follow
three different post-nesting movement
trajectories towards the equatorial
Atlantic Ocean, South America, or
southern Africa (Witt et al. 2011). For
combined foraging areas off Argentina
and Elevac¸a˜o do Rio Grande (an
elevated offshore area across from
Brazil), the mean estimate from western
Africa was 84 to 86 percent (45 percent
Gabon, 41 percent Ghana; Prosdocimi et
al. 2014).
The wide distribution of foraging
areas likely buffers the DPS against local
catastrophes or environmental changes
that could limit prey availability. The
expansive nesting range may buffer the
DPS from acute environmental impacts
(e.g., storms and singular events) and to
some degree, chronic impacts (e.g., sea
level rise and temperature changes).
Thus, the combination of extensive
nesting range, widely distributed
foraging areas, and population structure
reduces the extinction risk of the SE
Atlantic DPS.
Diversity
Genetic analyses for the SE Atlantic
DPS are limited, but Dutton et al. (2013)
found moderate genetic diversity in
samples from Gabon and Ghana,
including four new haplotypes unique
to western African nesting females.
Nesting occurs on continental and
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insular beaches. There are multiple
foraging strategies, including pelagic
and coastal, along either side of the
Atlantic Ocean. The genetic diversity,
along with multiple and diverse
foraging sites (i.e., coastal and pelagic),
and combination of insular and
mainland nesting provide diversity and
resilience that may reduce the
extinction risk of this DPS.
Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
Modification and loss of habitat is a
threat to the SE Atlantic DPS. Present
threats include obstructions, erosion,
and light pollution at nesting beaches.
Future threats include coastal
construction and development in the
region.
Nesting beach obstruction due to logs
is a problem in Gabon, Equatorial
Guinea, and Cameroon (Formia et al.
2003). Logs that have broken loose from
timber rafts of industrial logging
operations wash up on the beaches of
Gabon at densities of up to 247 logs/km;
logs blocked 30.5 percent of the beach
in Pongara, Gabon, resulting in an
estimated 2,111 disrupted or aborted
nesting attempts (Laurance et al. 2008).
In addition, several leatherback turtles
have died as result of being trapped by
logs (Laurance et al. 2008). Pikesley et
al. (2013) determined that between 1.6
percent and 4.4 percent of nesting
females could be trapped at beaches
with high log- and turtle-densities.
However, Gabon has since banned the
export of whole logs. The Gabon Sea
Turtle Partnership has carried out log
removal efforts for at least one highdensity nesting beach in Pongara
National Park (Kingere Beach), and a 3
km stretch of nesting beach is now
virtually free of logs; at the other main
monitored beaches in Gabon, such as
Mayumba and Gamba, logs are not a
major threat (A. Formia, WCS, pers.
comm. 2019).
Habitat loss from coastal erosion due
to sand mining, harbor building, and
irregular current flows has
compromised the suitability of long
stretches of coastal areas as nesting
sites. This issue is especially prevalent
between Ghana and Nigeria (Formia et
al. 2003). Ikaran (2010) found low
hatching/emergence success rates at
three nesting sites in Gabon: Pointe
Denis (17/16 percent), Mayumba (43/40
percent), and Kingere (16/16 percent).In
addition to predation, the main
identified sources of egg mortality were
beach erosion and inundation (Ikaran
2010).
Light pollution modifies nesting
beach habitat, deterring nesting females
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and disorienting both hatchlings and
nesting females. Bourgeois (2009) found
that artificial lighting disoriented
leatherback hatchlings in Pongara
National Park, Gabon: Hatchlings in 27
of the 41 nests (66 percent) studied
crawled towards artificial lights. Deem
et al. (2007) documented 71 disoriented
females that crawled directly into the
savannah behind the beach and towards
the artificial lights. Bourgeois et al.
(2009) concluded that light pollution
from Libreville and Pointe Denis, Gabon
is a major threat to nesting females and
hatchlings, which become disoriented
and die in the surrounding savannah.
Urbanization and coastal
development are rapidly growing threats
at some nesting beaches (Girard and
Honarvar 2017). There is a high
potential for coastal development in
Gabon, including the beaches near
Pointe Denis, an important and growing
tourist area (Ikaran 2010). Along with
direct habitat loss from coastal
development and urbanization, impacts
from pollution and litter are expected to
increase.
In Gabon, a network of marine
protected areas was created by decree
00161/PR in 2017, covering 26 percent
of Gabon’s territorial seas, including a
vast area in front of the most important
nesting beach in Gabon (Mayumba
National Park) that stretches to the outer
limits of the EEZ.
We conclude that a large portion of
nesting females, hatchlings, and eggs are
exposed to the reduction and
modification of nesting habitat, as a
result of logging, erosion, coastal
development, and artificial lighting.
These threats impact the DPS by
reducing nesting and hatching success,
thus lowering the productivity of the
DPS. Logging also results in the death of
nesting females, reducing the
abundance of the population by
removing its most reproductively
important individuals. Based on the
information presented above, we
conclude that habitat loss and
modification are major and increasing
threats to the DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization is a threat to the SE
Atlantic DPS. Although receiving some
legal protections, eggs and turtles
nevertheless are poached for
consumption, traditional medicine, and
religious practices.
In Gabon, poaching is limited because
78 percent of nesting occurs within
national parks and human population
density along the coast is low (A.
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Formia, Gabon Sea Turtle Partnership,
pers. comm., 2018). However, elsewhere
in the region, poaching occurs at a high
rate, or would be reasonably expected to
return to high levels, if not limited by
activities funded through the USFWS’
Marine Turtle Conservation Fund
enacted under the MTCA. These
activities reduce poaching through
increased presence on nesting beaches,
beach monitoring, hiring of local
citizens for participation in the projects,
and raising awareness and providing
education to local communities (M.
Tiwari, NMFS, pers. comm. 2018).
Conflicting beliefs about sea turtles
exist throughout the region. In some
communities sea turtles are considered
divinely provided food, while in others
they have been historically protected by
indigenous custom, often based on
stories passed down by ancestors
(Barbosa and Regalla 2016; Alexander et
al. 2017). In general, however, poaching
is a significant problem throughout the
region. Catry et al. (2009) concluded
that, in addition to fisheries bycatch,
poaching of eggs and nesting females is
the main threat to sea turtles, including
leatherback turtles, in Guinea-Bissau. In
many cases ‘‘few if any turtles or nests
are left alone when found by locals’’
(Catry et al. 2009). The fat of leatherback
turtles is often used for various
purported medicinal applications,
including: Treatment of convulsions
and malaria (Togo), fever, fainting
spells, liver problems, tetanus (Benin),
and to induce vomiting (Togo, Benin).
In one community in the Ivory Coast
and parts of Cameroon, leatherback
turtle fat is applied to wounds in the
mouth and is used to massage into
painful joints. In northwestern and
southern Cameroon, it is applied to
bruises (Fretey et al. 1999). In Togo,
some mothers add turtle bones daily to
the baby’s bath water; some believe that
the power of the turtle (especially the
leatherback) will be transmitted to the
child through this practice (Segniagbeto
2004).
Turtles and eggs are poached
throughout the nesting range of the DPS.
Though most nesting females and eggs
are protected in Gabon, poaching is
widespread in other areas. Poaching of
nesting females reduces both abundance
(through loss of nesting females) and
productivity (through loss of
reproductive potential). Such impacts
are high because they directly remove
the most productive individuals from
DPS, reducing current and/or future
reproductive potential. Egg poaching
reduces productivity. Given the
moderate exposure and high impact, we
conclude that the poaching of turtles
and eggs poses a threat to the DPS.
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Disease or Predation
Information on diseases among
leatherback turtles originating in the SE
Atlantic is minimal, but an analysis of
samples from nesting females in Gabon
indicated normal blood chemistry
parameters (Deem et al. 2006). Predation
may occur at high rates in some areas,
but information is limited.
Predation of leatherback eggs and/or
hatchlings has been documented for a
variety of predators, including: Various
ants, ghost crabs, monitor lizards
(Varanus niloticius), crows (Corvus
albus), mongoose, porcupine (Atherurus
africanus), domestic dogs, African civet
cat (Civettictis civetta and Viverra
civetta), and drills (Mandrillus
leucophaeus) (summarized from Eckert
et al. 2012). In Kingere, Gabon, Ikaran
(2010) noted high predation rates of eggs
by crabs, lizards, mongooses, small cat
species, and ants. Predation was the
main source of egg mortality at three
nesting sites in Gabon: Pointe Denis (43
percent), Mayumba (44 percent), and
Kingere (51 to 56 percent; Ikaran 2010).
As is common for all sea turtle
species, leatherback hatchlings likely
experience predation from various fish
species as they enter the water and
swim towards the open ocean. In-water
predation of juveniles and adults is not
well-documented, but there is evidence
of shark and killer whale predation.
Shark predation was determined to be
the cause of one leatherback stranding
reported from Central Africa (Parnell et
al. 2007), while interactions between
killer whales and leatherback turtles
resulting in possible predation has been
observed in Namibian waters (Elwen
and Leeney 2011).
While all eggs and hatchlings have
some exposure to predation, the species
compensates for a certain level of
natural predation by producing a large
number of eggs and hatchlings. For this
DPS, the primary impact is to
productivity (i.e., reduced egg and
hatching success). We conclude that
predation poses a threat to the SE
Atlantic DPS.
Inadequacy of Existing Regulatory
Mechanisms
The SE Atlantic DPS is protected by
various regulatory mechanisms. For
each, the Team reviewed the objectives
of the regulation and to what extent it
adequately addresses the targeted threat.
The harvest of turtles and eggs is
illegal in most of the nations where the
DPS nests. In some cases, however,
these protective mechanisms are
inadequate. In addition, many nesting
beaches are not protected.
In Gabon, the harvest of turtles and
eggs is illegal (2011 decree 0164/PR/
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MEF) and much of the nesting beach
habitat (and turtles utilizing that
habitat) is protected because of
inclusion in parks as well as being far
from any city or town. However, low
levels of poaching occurs, and the
threats from encroaching development
and associated impacts are increasing.
In Congo, wildlife laws prohibit the
hunting and collection of wildlife and
their products, including eggs, between
November 1 and April 31. Turtles are
also protected in the Conkaouati-Douli
National Park. However, in areas
without permanent beach monitoring,
almost all eggs and nesting individuals
are collected and eaten (Bal et al. 2007).
In the Democratic Republic of Congo,
leatherback turtles are cited under the
1982 Hunting Act for protection.
However, there is no post-independence
legislation protecting sea turtles, and
there is little commitment to the
legislated protections (Fretey 2001).
Since 1988, Equatorial Guinea has
protected all sea turtles under Law 8/
1988 and Decree 183/87 on fishing
(Toma´s et al. 2010). However, the
poaching of eggs and females for local
consumption and sale has occurred
(Castroviejo et al. 1994).
In Ghana, the Wildlife Regulations
Act of 1974 prohibits all harvest of eggs
and turtles. However, poverty is
prevalent, and eggs and sea turtles are
poached at nesting beaches (Tanner
2013). Enforcement is likely inadequate
because of funding issues, the
remoteness of some nesting beaches,
and cultural practices.
Fishery bycatch is the primary threat
to this DPS. While most nations in the
region have some form of legal
protection for sea turtles, many
leatherback turtles die from fisheries
bycatch throughout the range of the
DPS. Examples of fisheries legislation
include Brazil’s gear restrictions and
Nigeria’s requirement to use TEDs in
bottom trawls.
In summary, numerous regulatory
mechanisms provide some protection to
leatherback turtles, their eggs, and
nesting habitat throughout the range of
this DPS. Though the regulatory
mechanisms provide some protection to
the turtles, many do not adequately
reduce the threat that they were
designed to address, generally as a
result of limited implementation or
enforcement. Fisheries bycatch,
poaching, and habitat loss remain major
threats to the DPS despite regulatory
mechanisms. We conclude that
inadequacy of the regulatory
mechanisms are a threat to the SE
Atlantic DPS.
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Fisheries Bycatch
Fisheries bycatch is the primary threat
to the SE Atlantic DPS. Leatherback
turtles are captured as bycatch in
commercial and artisanal fisheries along
coastal foraging and breeding areas as
well as on the high seas. Because of the
overlapping range with the SW Atlantic
DPS, this DPS is vulnerable to
interactions with fisheries off the coasts
of Brazil, Uruguay, and Argentina, in
the pelagic waters of the South Atlantic
Ocean, and along the coastal waters off
western Africa. Therefore, the
information presented on the fisheries
bycatch for the SW Atlantic is
applicable to this DPS.
One of the biggest threats for
leatherback turtles in Atlantic waters is
bycatch in artisanal and commercial
fisheries (Wallace et al. 2010; Riskas and
Tiwari 2013;). Lewison et al. (2004)
estimated that 30,000 to 60,000
leatherback turtles were taken as
longline fisheries bycatch in the entire
Atlantic Ocean in 2000. Stewart et al.
(2010) estimated that in West Africa,
Benin, Togo, and Cameroon had the
highest average fishing densities,
ranging from 11.1 to 6.5 boat-meters/
km2, and gillnet densities ranked among
the highest on a global scale. Despite
very active artisanal and industrial
fisheries in the region, overall bycatch
data are quite sparse or qualitative
(rather than quantitative) in nature, and
Africa still represents a significant gap
in bycatch evaluation studies (Wallace
et al. 2010, 2013). Accurate and reliable
bycatch data are difficult to achieve, as
direct observation rates are low (<1
percent of total fleets) and statistics
from the region’s many small-scale
fisheries are largely incomplete
(Kelleher 2005; Moore et al. 2010;
Wallace et al. 2010). However, several
studies have concluded that bycatch
rates in the region are high, given the
degree of fishing activity near nesting
and foraging areas (Lewison et al. 2004;
Moore et al. 2010; Wallace et al. 2010).
Along the coasts of Angola, Namibia,
and South Africa, Honig et al. (2007)
evaluated turtle bycatch by longline
fisheries in the Benguela Large Marine
Ecosystem by using data from observer
reports, surveys, and specialized trips
from the coastal nations of South Africa,
Namibia and Angola. They estimated
bycatch at 672 leatherback turtles
annually (based on an annual bycatch
estimate of 4,200 turtles, of which
approximately 16 percent are
leatherback turtles) in the southern and
central regions and as many as 5,600
leatherback turtles (based on an annual
bycatch estimate of 35,000 turtles) for
the entire Benguela Large Marine
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48371
Ecosystem (Honig et al. 2007). Mortality
rates were not provided in this study
but may range from 25 to 75 percent
(Aguilar et al. 1995). The estimates
mostly include turtles from the SE
Atlantic DPS, but telemetry studies
indicate that the turtles of the much
smaller SW Indian DPS also use this
foraging area (Luschi et al. 2006;
Robinson et al. 2016). Evaluating ICCAT
data, Angel et al. (2014) confirm
exposure to high longline fishing effort
and some purse seine effort for the
population originating from the SE
Atlantic Ocean.
The limited bycatch data available for
waters of the western coast of Africa
show that other fisheries interact with
leatherback turtles. Between 2005 and
2015, artisanal fishing nets in Loango
Bay in the Republic of Congo killed a
total of 45 leatherback turtles; 0 to 628
leatherback turtles were captured or
recaptured annually over that time
period (Bre´heret et al. 2017). An
assessment of bycatch in the trawling
fisheries in Gabon found that
leatherback turtles represented only 2
percent of the bycatch despite being the
most abundant sea turtle species in
Gabonese waters; the low rate is
possibly because leatherback turtles do
not occur in the section of the water
column where the trawl net is towed
(Casale et al. 2017). Trawl bycatch in the
waters around Sa˜o Tome´ and Principe
included 4 juvenile leatherback turtles
(17 to 21 cm in carapace length) in
March 1994 (Fretey et al. 1999).
While specific information to estimate
overall capture and mortality rates of SE
Atlantic leatherback turtles in fisheries
is not available, it is clear that bycatch
in fisheries, especially gillnets and
longlines, are a threat to the DPS across
its range. Immature and mature
individuals are exposed to high fishing
effort throughout their foraging range
and in coastal waters near nesting
beaches. Mortality is also high.
Mortality reduces abundance, by
removing individuals from the
population; it also reduces productivity,
when nesting females are incidentally
captured and killed. We conclude that
fisheries bycatch is a major, and the
primary, threat to the SE Atlantic DPS.
Vessel Strikes
There is little information regarding
vessel strikes for the SE Atlantic DPS,
but such interactions are a potential,
and possibly increasing, threat across at
least a portion of this DPS’s range. In the
western South Atlantic foraging grounds
off Brazil, Uruguay, and Argentina,
increasing vessel traffic from fishing
vessels, cargo transport, and tourism has
been noted (Lo´pez-Mendilaharsu et al.
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2009; Fossette et al. 2014), potentially
increasing the likelihood of vessel
strikes on leatherback turtles. Although
no specific information is available for
the waters off western Africa, any
economic development along the coast
is likely to result in an increase in vessel
traffic. We conclude that vessel strikes
are a threat to the SE Atlantic DPS.
Pollution
The SE Atlantic DPS faces the threat
of pollution across its extensive range
throughout the South Atlantic Ocean,
from Africa to South America. As the
ranges of the SW Atlantic and SE
Atlantic DPSs overlap, they are exposed
to the same pollutants, which include
contaminants, marine debris, and ghost
fishing gear. Throughout Africa, marine
and coastal pollution is widespread in
industrial and urban areas, and garbage
litters many developed beaches (Formia
et al. 2003; Agyekumhene et al. 2017).
Off the coast of South America, the
Argentine and Brazilian coastal waters
are increasingly impacted by economic
activities, such as maritime cargo
transport, tourism, and the discharge of
domestic and industrial waste (Lo´pezMendilaharsu et al. 2009; Fossette et al.
2014).
The Gulf of Guinea has increasingly
been the focus of extensive oil
exploitation activities, following the
discovery of large oil reserves. Drilling
activities by large oil corporations, with
associated pollution and habitat
destruction, are threats to nesting
aggregations in the area (Formia et al.
2003; Agyekumhene et al. 2017). In
2012/2013, oil spills following the
dredging of the Port of Pointe-Noire in
the Republic of Congo significantly
degraded the fauna and flora of Loango
Bay, where leatherback turtles occur.
However, the ecosystem is believed to
be slowly recovering (Bre´heret et al.
2017). In 2005, a moderate slick of oil
on the beaches of Mayumba National
Park in Gabon was observed, although
its impacts on turtles are unknown
(Parnell et al. 2007).
In Nigeria, the main sources of
pollution include industrial waste, raw/
untreated sewage, and pesticides. Oil
exploration, exploitation, and
transportation have a significant effect
on the environment. Spills of crude and
refined oil are frequent in the coastal
and marine environment, especially
during periods of very strong ocean
currents, when they can spread to cover
the entire 853 km coastline of Nigeria.
It is clear that individuals from the SE
Atlantic DPS have a high probability of
encountering pollution across their
range and throughout their lifecycle.
Although the best available information
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does not quantify such impacts, ample
information demonstrates that these
threats are ongoing. We conclude that
pollution is a threat to the DPS.
Climate Change
Climate change is a threat to the SE
Atlantic DPS. The impacts of climate
change include: Increases in
temperatures (air, sand, and sea
surface); sea level rise; increased coastal
erosion; more frequent and intense
storm events; and changes in ocean
currents.
Sea level rise resulting from climate
change negatively impacts sea turtle
nesting. Erosion of important nesting
beaches in Gabon may be at least
partially attributable to sea level rise.
From 1983 through the 2000s, some
areas have lost up to 100 m of beach
width, reducing the availability of
suitable nesting beach (Gabon Sea
Turtle Partnership 2018; https://
www.seaturtle.org/groups/gabon/
erosion.html). Because leatherback
turtles nest lower on the beach than
other sea turtles, their eggs are more at
risk of being inundated and destroyed
by increases in sea level and coastal
erosion (Boyes et al. 2010).
Changes in sand temperatures are
likely to impact egg viability and sex
determination. Ikaran (2010) found the
thermal range of sand over the nesting
season to be adequate for hatchling sex
ratios to be mixed or even male
dominated. In Gabon, the early rainy
months tend to produce males, while
the later, warmer months produce
females, with a tendency towards a net
higher production of males. Ikaran
(2010) considered the nesting beaches of
Gabon to be an important male
producing area. However, based on
predictions of warming trends, he found
that within two decades the ratio could
skew towards 100 percent female.
The threat of climate change is likely
to modify the nesting conditions for
turtles of the DPS, and it is unclear
whether they have or can develop the
ability to nest in different locations
along existing beaches, or on new
beaches. Impacts from climate change
are likely to range from small, temporal
changes in nesting season to large losses
of productivity. Therefore, we conclude
that climate change is a threat to the
DPS.
Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply
within the range of the SE Atlantic DPS
(for a description of each effort, please
see the section on conservation efforts
for the overall species): Convention on
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the Conservation of Migratory Species of
Wild Animals, Convention on Biological
Diversity, Convention on International
Trade in Endangered Species of Wild
Fauna and Flora, Convention
Concerning the Protection of the World
Cultural and Natural Heritage (World
Heritage Convention), FAO Technical
Consultation on Sea Turtle-Fishery
Interactions, IAC, MARPOL, IUCN,
Memorandum of Understanding
Concerning Conservation Measures for
Marine Turtles of the Atlantic Coast of
Africa, Ramsar Convention on
Wetlands, South-East Atlantic Fisheries
Organization, UNCLOS, and UN
Resolution 44/225 on Large-Scale
Pelagic Driftnet Fishing. Although
numerous conservation efforts apply to
the turtles of this DPS, they do not
adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded
overall that the SE Atlantic DPS is at
high risk of extinction. The total index
of nesting female abundance is 9,198
females. Since 2002, the first year that
aerial survey data was collected, nesting
activity has declined by ¥8.6 percent
annually in Gabon, the largest nesting
aggregation of the DPS, and what was,
in 2002, the largest nesting aggregation
in the world. This declining trend has
the potential to further lower abundance
and increase the risk of extinction.
Nesting and foraging is broadly
distributed; thus, the population is
somewhat buffered from stochastic
events that could otherwise have
catastrophic effects on the entire DPS.
There is a metapopulation structure
within this DPS, with fine-scale genetic
differentiation between Gabon and
Ghana. Genetic diversity also appears to
be moderate. Based on the reduced
nesting female abundance and declining
nest trend, we find the DPS to be at risk
of extinction, likely as a result of past
threats.
Current threats place the DPS at
further risk of extinction. The primary
threat to this DPS is bycatch in
commercial and artisanal, pelagic and
coastal, fisheries, especially coastal
gillnet and pelagic longline fisheries.
Fisheries bycatch reduces abundance by
removing individuals from the
population. Because several fisheries
operate near nesting beaches,
productivity is also reduced when
nesting females are prevented from
returning to nesting beaches. Thus,
exposure and impact of this threat are
high. Habitat loss or modification is a
threat that reduces abundance and
productivity and includes the impacts
of logs, which block access to the
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beaches or trap nesting females and
hatchlings. Poaching of turtles and eggs
is also a threat to this DPS, although
most nesting beaches in Gabon are
somewhat protected because they occur
in parks or are far from any towns.
Many of the beaches outside Gabon
(e.g., Guinea-Bissau) have limited or no
protection. The degree of overutilization
is highly varied across locations, but
quite extensive in some areas. Funding
from the MTCA has resulted in some
reduction of this threat as conservation
activities, research, and community
involvement results in lower poaching
on those beaches. However, poaching
continues at high levels in other areas.
Additional threats include: predation
and disease, inadequate regulatory
mechanisms, pollution, and climate
change. Predation can be extensive at
some specific beaches, but overall it
does not occur at a high level. Pollution
is a persistent and potentially increasing
threat. Ingestion of plastics and
entanglement in marine debris result in
injury and reduced health, and
sometimes mortality. Climate change is
likely to result in reduced productivity
due to greater rates of coastal erosion
and nest inundation, and in some areas,
nest failure or skewed sex ratios due to
increased sand temperatures. Vessel
strikes are a threat that is likely to
increase over time as recreational and
commercial vessel activity increases,
resulting in more opportunity for
interactions. Though many regulatory
mechanisms are in place, they do not
adequately reduce the impact of logs,
poaching, and fisheries. Additionally,
many areas in the region have little or
no enforcement of laws protecting
turtles or nests on the beach.
The DPS is relatively data-poor,
reducing our ability to quantify threats
for more than a small portion of the
population. For this reason, the Status
Review Team did not come to
consensus regarding the extinction risk
analysis for the SE Atlantic DPS. All
Team members were present to vote on
the level of extinction risk. Nine Team
members concluded with moderate
confidence that the DPS is at high
extinction risk due to threats and loss of
abundance; their confidence was
moderate due to the lack of data on this
DPS. Two team members concluded
with low confidence that the DPS is at
moderate extinction risk; their
confidence in this conclusion is low due
to the lack of data on this DPS.
We conclude, consistent with the
Team’s overall conclusion, that the SE
Atlantic DPS is currently in danger of
extinction. The decreasing nesting trend
(i.e., 8.6 percent annually since 2002) is
at or near a level that make the DPS
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highly vulnerable to threats, given the
total index of nesting female abundance
of 9,198 females. It faces present,
ongoing threats that are likely to create
imminent and substantial demographic
risks (i.e., declining trends and reduced
abundance). Though numerous
conservation efforts apply within the
range of this DPS, they do not
adequately reduce the risk of extinction.
We conclude that the SE Atlantic DPS
is currently in danger of extinction
throughout its range and therefore meets
the definition of an endangered species.
The threatened species definition does
not apply because the DPS is at risk of
extinction currently (i.e., at present),
rather than on a trajectory to become so
in the foreseeable future.
SW Indian DPS
The Team defined the SW Indian DPS
as leatherback turtles originating from
the SW Indian Ocean, north of 47° S,
east of 20° E, and west of 61.577° E. The
western boundary occurs at the
southern tip of Africa, approximately
where the Agulhas and Benguela
Currents meet. The eastern boundary
occurs at the border between Iran and
Pakistan, where the Somali Current
begins. These currents, and the cold
waters of the Antarctic Circumpolar
Current, likely restrict the nesting range
of this DPS.
The range of the DPS (i.e., all
documented areas of occurrence)
extends into the SE Atlantic Ocean,
where leatherback turtles forage in the
highly productive Benguela Current
Large Marine Ecosystem, which occurs
along the western coast of Africa, from
Angola to South Africa. Leatherback
turtles also range throughout the waters
of eastern Africa (Ross 1985) and
possibly into the Red Sea (Gasparetti et
al. 1993). Records indicate that the
species has been observed in the waters
of the following nations: Djibouti;
Eritrea; French Territories (Reunion
Island, Mayotte, and Iles Eparses);
Kenya; Madagascar; Mozambique;
Seychelles; Somalia; South Africa;
Tanzania; and Yemen (Hamann et al.
2006). Leatherback turtles may occur in
the waters of the following nations:
Bahrain, Kuwait; United Arab Emirates;
Oman; and Sudan (Hamann et al. 2006).
Leatherback turtles of the SW Indian
DPS nest over a distance of
approximately 900 km, from Cape Vidal,
South Africa to Bazaruto Islands,
Mozambique (Videira et al. 2011; Nel et
al. 2015). The vast majority of nesting
(80 to 90 percent) occurs in South
Africa, between Bhanga Nek and
Leifeld’s Rock (Nel et al. 2015). In
Mozambique, most nesting occurs from
the southern border to Inhaca Island,
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Mozambique, with low levels of nesting
farther north at Bilene Beach and
Bazaruto Islands (Nel et al. 2015). This
DPS nests at the highest latitude (and
southernmost location) of all
leatherback turtles (Saba et al. 2015).
Nesting occurs on long (5 to 15 km),
broad (50 to 100 m), silica sand beaches
with little vegetation (Botha 2010; Nel et
al. 2015; Robinson et al. 2017). The
beaches are characterized by pristine,
intact dunes that rise up to 100 m above
sea level, interspersed with a few
dynamic dunes and small, primary
dunes (Nel et al. 2015). The beaches are
separated by short rocky headlands
(Robinson et al. 2017). Subtidal rock
formations are dispersed throughout the
high energy coastline. Nesting females
approach the beach using strong ripcurrents through obstruction-free areas
(Hughes 1974; Hughes 1996; Botha
2010; Nel et al. 2015).
Foraging areas of the SW Indian DPS
include coastal and pelagic waters of the
SW Indian Ocean and the SE Atlantic
Ocean. The DPS is somewhat unique in
that turtles forage in two ocean basins
and do not need to undergo long
migrations between nesting and foraging
areas because highly productive
foraging areas are available adjacent to
nesting beaches or connected to nesting
beaches via fast-moving currents. For
example, the warm, fast-flowing
Agulhas Current (Lutjeharms 2001; Nel
et al. 2015) results in high productivity
foraging areas near nesting beaches and
provides a migratory corridor to distant
foraging areas. As a result, the SW
Indian turtles have the largest body size,
largest clutch size, and highest
reproductive output of all leatherback
turtles (Saba et al. 2015).
Satellite tracking of post-nesting
females (n = 27) reveals the use of one
of three post-nesting migratory
corridors: north into the nearby coastal
waters of the Mozambique channel;
south and west (via the Agulhas and
Benguela Currents) into the pelagic
waters of the South Atlantic Ocean; or
south and east (via the Agulhas Current
and Retroflection) into the oceanic
eddies in the SW Indian Ocean (Luschi
et al. 2006; Robinson et al. 2016; Harris
et al. 2018). Luschi et al. (2006)
reviewed satellite telemetry data of 11
post-nesting females tagged between
1996 and 2003 (Hughes et al. 1998;
Luschi et al. 2003; Sale et al. 2006); and
Robinson et al. (2016) satellite tracked
16 post-nesting females tagged between
2011 and 2013. Evaluating tracking data
for 14 post-nesting females between
2006 and 2014, Harris et al. (2018)
found that leatherback turtles equally
used all three migration corridors. In the
other studies, a total of 11 post-nesting
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females migrated a relatively short
distance (approximately 500 km) to the
shallow (less than 50 m depth), coastal
waters of the Sofala Banks (i.e., the
Mozambique Channel), where net
primary productivity and sea surface
temperatures remain elevated yearround (n = 4, Sale et al. 2006; n = 7,
Robinson et al. 2016). One post-nesting
female migrated to the similarly
hospitable coastal waters of Madagascar
(Robinson et al. 2016). Ten post-nesting
females tracked to pelagic waters of the
Atlantic Ocean (n = 6, Sale et al. 2006;
n = 4, Robinson et al. 2016). These
waters are among the most productive
in the world, as a result of strong
upwelling (caused by the southeast
trade winds) and the area’s unique
bathymetry, hydrography, chemistry,
and trophodynamics (Honig et al. 2007).
Five post-nesting females appeared to
track oceanic eddies into the SW Indian
Ocean (n = 1, Sale et al. 2006; n = 4,
Robinson et al. 2016). Luschi et al.
(2003 and 2006) characterized
leatherback turtles using this latter
strategy as ‘‘wanderers, ranging over
vast oceanic areas while searching for
their planktonic prey.’’
Opportunistically encountered and
highly productive eddies likely shaped
the circuitous routes of these foraging
turtles, which resemble drifters more
than active swimmers (Luschi et al.
2006; Robinson et al. 2016; Harris et al.
2018). Thus, this DPS benefits from the
use of three migratory corridors that all
provide highly productive foraging
opportunities, with minimal energetic
cost required to return to waters off
nesting beaches.
Abundance
The total index of nesting female
abundance of the SW Indian DPS is 149
females. We based this index on two
nesting aggregations: South Africa
(Ezemvelo KwaZulu-Natal Wildlife
(Ezemvelo), unpublished data, 2018)
and Mozambique (Centro Terra Viva
Estudos e Advocacia Ambiental (CTV),
unpublished data, 2018). Our total
index does not include two
unquantified nesting aggregations in
Mozambique. To calculate the index of
nesting female abundance (i.e., 134
females) for the South Africa
‘‘monitoring area’’ (i.e., a 52.8 km
stretch of beach that has been monitored
for decades), we divided the total
number of nests between the 2014/2015
and 2016/2017 nesting seasons (i.e., a 3year remigration interval; Hughes 1996;
Lambardi et al. 2008; Nel et al. 2013;
Saba et al. 2015) by the clutch frequency
(7 clutches/season; Nel et al. 2013; Saba
et al. 2015). To calculate the index of
nesting female abundance in
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Mozambique (i.e., 15 females), we
divided the total number of nests
between the 2015/2016 and 2017/2018
nesting seasons (i.e., a 3-year
remigration interval) by the clutch
frequency for South Africa (7 clutches/
season; Nel et al. 2013; Saba et al. 2015).
This is an index for the DPS because
it only includes available data from
recently and consistently monitored
nesting beaches. While nesting occurs
on beaches that stretch across 900 km of
South Africa and Mozambique,
consistent and standardized monitoring
occurs only across approximately 300
km of beaches across the two nations
(Nel et al. 2013; Nel et al. 2015).
Furthermore, while nesting is known to
occur at low levels at Inhaca Island and
Bazaruto Archipelago in Mozambique,
we did not include these sites because
we did not have data from the most
recent 3 years.
Other estimates of total or annual
nesting female abundance have been
published. The IUCN Red List
assessment estimated the total number
of mature individuals (males and
females) at 148 individuals, based on an
average of 259 annual nests (Nel et al.
2013), a 3-year remigration interval (Nel
et al. 2013), and a 3:1 sex ratio (Wallace
et al. 2013). Their estimates are based on
nesting surveys conducted in South
Africa, which hosts approximately 80 to
90 percent of nesting, and Mozambique
(Wallace et al. 2013; Nel et al. 2015).
Their estimate is less than our index,
despite including mature males and
females. The reason for this difference is
because they used an average annual
number of nests that was lower than
recent nest counts over the 3-year
remigration interval. Nel et al. (2015)
estimated the size of the total nesting
population at approximately 100
females per season (Nel et al. 2015),
based on 2010 data: 375 emergences and
336 nests in South Africa; and 61
emergences in Mozambique (Videira et
al. 2011). This estimate (n = 300, based
on a 3 year remigration interval) is
greater than our index because there
were more nests in 2010 compared to
more recent years (2014 to 2016).
Hamann et al. (2006) estimated
approximately 20 to 40 nesting females
annually in South Africa and
approximately 10 nesting females
annually in southern Mozambique. This
estimate (n = 90 to 150, based on a 3
year remigration interval) is less than
our index, likely as a result of using data
collected over a different time-frame.
The difference in estimates likely results
from using different methods of
calculation and different time frames
and reflects some uncertainty in the
precise number of nesting females. Our
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total index of nesting female abundance
falls within the range of other estimates
and is based on the best available data
for the DPS at this time.
There are additional published
estimates for the South Africa
monitoring area. Nel et al. (2013)
identified 2,578 nesting females over 45
years (1965 to 2009), with a mean of
69.4 ± 38.1 nesting females per season
(or 209 total nesting females) in the
monitoring area. Hughes (1996) reported
an annual average of 24 nesting females
in the first decade (1976 to 1985) and an
annual average of 86 nesting females in
the second decade (1986 to 1995) in the
monitoring area. Hughes (1996) also
reported an annual average of 113
nesting females from 1986 to 1995 in an
extended protected area that includes
the monitoring area plus another 93 km
in the St. Lucia Marine Reserve, which
is surveyed periodically. The difference
between these two averages reflects that
most estimates of nesting female
abundance in South Africa are
minimum estimates because nesting
occurs outside the monitoring area.
Thorson et al. (2012) found that annual
resightings for leatherback turtles
decreased from the 1960s to 2009, and
their modeling indicated that this
decline was due to decreased detection
probabilities (i.e., decreased probability
of returning to the monitored portion of
the KwaZulu-Natal nesting beach),
rather than decreased survival. Based on
satellite tracking of 17 post-nesting
females, Harris et al. (2015) estimates
that approximately 66 percent of
leatherback nesting activity occurs
outside the monitoring area. However,
considerable inter-annual variability
exists, ranging from less than 30 percent
to over 80 percent, with a median of
approximately 49 percent (Harris et al.
2015). Thus, incomplete beach
monitoring is a source of uncertainty for
this DPS and for our total index of
nesting female abundance.
For Mozambique, our index of nesting
females is similar to other published
estimates, which are generally less than
20 nesting females (Hamann et al. 2006;
Louro 2014; Pereira et al. 2014;
Fernandes et al. 2018). If we use the
clutch frequency for Ponta Malongane
(2.25 clutches per season; Louro et al.
2006), which is low for the species, our
index of nesting female abundance is 45
females. This clutch frequency may be
underestimated due to females nesting
in distant areas where monitoring does
not regularly occur. If we use the clutch
frequency for South Africa, (7 clutches/
season; Nel et al. 2013; Saba et al. 2015),
the resulting index of nesting female
abundance for Mozambique (i.e., 15
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nesting females) is closer to published
estimates.
The total index of nesting female
abundance of 149 females places the
DPS at risk for environmental variation,
genetic complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). These
processes, working alone or in concert,
place small populations at a greater
extinction risk than large populations,
which are better able to absorb losses in
individuals. Due to its small size, the
DPS has restricted capacity to buffer
such losses. Given the intrinsic
problems of small population size, we
conclude that the limited nesting female
abundance is a major factor in the
extinction risk of this DPS.
Productivity
The SW Indian DPS exhibits a slightly
decreasing nesting trend. We base our
conclusion on data consistently
collected in a standardized approach in
the 56 km South African monitoring
area (Ezemvelo, unpublished data,
2018), where nest counts decreased by
¥0.3 percent annually (sd = 2.1 percent;
95 percent CI = ¥4.5 to 4.1 percent; f
= 0.557; mean annual nests = 301)
between the 1973/1974 and 2016/2017
nesting seasons. The trend in South
Africa is likely representative of the
entire DPS, as 80 to 90 percent of
nesting is estimated to occur there
(Wallace et al. 2013; Nel et al. 2015) and
the 44-year time series is quite robust.
Our trend estimates yield similar
results to other published findings for
the population. The IUCN concluded
that this population has declined
slightly, by 5.6 percent over the past
three generations, with an annual
decline of ¥0.1 percent in South Africa
and ¥0.7 percent in Mozambique
(Wallace et al. 2013). Hamann et al.
(2006) also identified a declining trend
in the nesting population of the SW
Indian Ocean. Studies focused on the
South African monitoring area (i.e., the
source of data for our trend analysis),
however, disagree on the whether the
trend has declined recently (Hamann et
al. 2006; Nel et al. 2013) or is stable (Nel
et al. 2015; Saba et al. 2015). The nest
trend may be stable if nesting in
unmonitored areas has increased over
time (Thorson et al. 2012; Harris et al.
2015). Different datasets lead to
different conclusions due to different
methods of calculation, different time
frames, incomplete monitoring of all
nesting areas, and therefore uncertainty
in the precise number of nesting
females. We find that Nel et al. (2013)
provide the best available published
data, which are based on the most
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recent, primary data, and we agree with
their characterization of the trend as
declining or recently declining.
Despite the recent decline in nesting,
productivity parameters remain
relatively high for the SW Indian DPS,
which has the largest body size, largest
clutch size, and highest reproductive
output of all leatherback turtles, likely
due to the close proximity between their
nesting beaches and highly productive
foraging areas (Saba et al. 2015). Nel et
al. (2015) reports that most metrics (i.e.,
female size, egg size, incubation time,
and hatching success) are above average
for this DPS. Nesting females produced
1,171 to 53,139 hatchlings each season
in the South Africa monitoring area
between 1965 and 2009, with an average
of 36,583 to 51,610 hatchlings per
season, which was calculated by
multiplying 480 hatchlings per nesting
female by 69.4 ± 38.1 nesting females
per season (Nel et al. 2013).
The recent nesting decline may reflect
the effects of past and current threats
that overwhelm the population’s high
productivity metrics. We conclude that
the slightly declining nest trend places
the DPS at risk of extinction, which is
further exacerbated by the limited
nesting female abundance.
Spatial Distribution
The SW Indian DPS comprises, in
essence, a single nesting aggregation,
with nesting females moving freely
between South African and
Mozambican beaches (Hughes 1996;
Luschi et al. 2006; Nel et al. 2015).
Nesting is limited to a total distance of
approximately 900 km along South
African and Mozambican coasts (Nel et
al. 2015). While 80 to 90 percent of
nesting is concentrated in South Africa,
nesting is somewhat concentrated in the
southern section of the South African
monitoring area, although most
characterize nesting as low density
throughout South Africa (Hughes 1974;
Lambardi et al. 2008; Botha 2010; Nel et
al. 2013; Harris et al. 2015; Nel et al.
2015).
The DPS exhibits a broad foraging
range that extends into coastal and
pelagic waters of the eastern Atlantic
and western Indian Oceans (Luschi et
al. 2006; Lambardi et al. 2008; Girondot
2015). There is limited evidence that
leatherback turtles may remain in South
African waters throughout the year, as
suggested by year-round fisheries
bycatch records (Luschi et al. 2003,
2006; Petersen et al. 2009). Some forage
off the coast of Madagascar (Robinson et
al. 2016; Harris et al. 2018). Some
turtles follow the Agulhas and Benguela
Currents into foraging areas in the
southeast Atlantic Ocean, off the coasts
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of Angola and Namibia (Girondot 2015;
Robinson et al. 2016; Harris et al. 2018).
Others follow the Agulhas Retroflection
and deep-sea eddies into the SW Indian
Ocean (Luschi et al. 2006; Lambardi et
al. 2008; Robinson et al. 2016; Harris et
al. 2018). Leatherback turtles, possibly
from this DPS, have also been observed
in the Red Sea, presumably foraging
(Hamann et al. 2006). The use of various
foraging areas may be influenced by the
prevalent currents encountered off the
nesting beaches (Luschi et al. 2006;
Lambardi et al. 2008; Robinson et al.
2016).
The wide distribution of foraging
areas likely buffers the DPS somewhat
against local catastrophes or
environmental changes that would limit
prey availability. Nesting occurs along
one coastline, which is 3,000 km in
length and may be similarly affected by
environmental variation and directional
changes (e.g., sea level rise). Because the
DPS is essentially a single nesting
aggregation, it has limited capacity to
withstand other catastrophic events.
Thus, spatial distribution likely has
little net effect on the extinction risk of
the SW Indian DPS.
Diversity
Within the SW Indian DPS, genetic
diversity is low, with only two mtDNA
haplotypes found in 41 nesting females
in South Africa (haplotype diversity =
0.298 ± 0.078 and nucleotide diversity
= 0.0004 ± 0.0004; Dutton et al. 2013).
Nesting habitat is mainly restricted to
beaches along the same coast, with a
few nests on Mozambican islands. The
DPS does not exhibit temporal or
seasonal nesting diversity, with most
nesting occurring between October and
March. The foraging strategies are
diverse, however, with turtles using
coastal and pelagic waters in the
Atlantic and Indian Oceans. Diverse
foraging strategies may provide some
resilience against local reductions in
prey availability or catastrophic events,
such as oil spills, by limiting exposure.
Low genetic diversity indicates the DPS
may lack the raw material necessary for
adapting to long-term environmental
changes, such as cyclic or directional
changes in ocean environments due to
natural and human causes (McElhany et
al. 2000; NMFS 2017). We conclude that
limited overall diversity increases the
extinction risk of this DPS by reducing
its resilience to threats.
Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
Coastal erosion, foot and vehicle
traffic, and artificial lighting modify the
available, suitable nesting habitat and
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thus are threats to the SW Indian DPS.
Angel et al. (2014) identifies coastal
erosion as the main beach-based threat
to this population and one that is likely
to increase with climate change.
Coastal erosion removes sand from
nesting beaches, inundating nests and
destroying eggs. Because leatherback
turtles nest lower on the beach than
other sea turtles, they have greater
exposure to tidal erosion and deposition
(Boyes et al. 2010). At South African
nesting beaches over a duration of 70
days, Boyes et al. (2010) found an
average of 0.62 m deposition (S.D. 0.15
m; range 0.34–0.85 m) and 0.42 m
erosion (S.D. 0.17 m; range 0.14– 0.71
m). Because the average depth of
leatherback nests was 0.66 m (S.D. 0.19
m; range 0.15–1.07 m), eggs are at some
risk of being exposed and destroyed
(Boyes et al. 2010). Nel et al. (2006)
concludes that coastal erosion is a threat
in South Africa, where the high-energy
coastline varies seasonally. During two
nesting seasons (2009/2010 and 2010/
2011), de Wet (2012) found that 6.3
percent of nests in the South African
monitoring area were destroyed by
erosion. In Bazaruto Archipelago,
Mozambique, coastal erosion and rising
sea levels destroyed approximately 12
percent of nests over 10 seasons of
monitoring (Videira and Louro 2005;
Louro 2006). Despite nest loss due to
erosion, hatching success remains high
in South Africa (70 to 80 percent; Nel
et al. 2015; Santidria´n Tomillo et al.
2015). Though the introduction of
Casuarina trees do not necessarily
increase the risk of erosion, they
obstruct nesting females’ access to and
from beaches and alter nest incubation
environments (de Vos et al. 2019).
Evolving in a high-energy coastline
environment with seasonal variation has
likely provided the DPS with some
resilience to nesting losses due to
coastal erosion. Sea level rise as a result
of climate change, however, is likely to
increase the rate and magnitude of this
natural process.
In Mozambique, Louro (2006)
describes beach driving as a ‘‘very
serious problem.’’ Tourism and beach
driving are increasing in Ponta
Malongane and Bazaruto Island, nesting
areas in Mozambique, where there is no
legislation regarding beach driving
(Louro 2006). Foot and vehicular traffic,
for tourism and recreational purposes,
have been found to impact nesting
beach habitat and turtles in several
ways. Beach activities can deter females
from using a nesting beach. Beach
driving causes sand compaction, which
may lower nest success. It also creates
ruts that slow hatchlings’ crawl to the
surf, increasing their vulnerability to
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predators. Beach driving occurs to a
lesser extent in South Africa.
Recreational beach driving is allowed
on a 1.5 km stretch of beach, and
tourism driving (for concession,
management, and media) involves a
maximum of 10 vehicles per night
across 40 km of beach (Nel 2006).
Artificial lighting modifies the quality
of nesting beaches because lights over
land disorient nesting females and
hatchlings. Instead of crawling toward
the surf and their marine habitat, they
crawl further inland, where they may
become dehydrated and die or become
susceptible to predation. Within the 280
km of coastline within the iSimangaliso
Wetland Park, South Africa, there are
only four areas of less than 100 m each
that contain artificial lighting (Nel
2006). We were unable to find data on
artificial lighting in Mozambique.
The majority of nesting habitat occurs
within the 280 km coastline of the
iSimangaliso Wetland Park in South
Africa, which has been a World Heritage
Site since 1999 (UN Educational,
Scientific and Cultural Organization
1999; Hughes 2010; Robinson et al.
2016). From 1979 to 1999, much of the
nesting habitat and nearshore marine
habitat was protected, first as the St.
Lucia Marine Reserve, then the
Maputaland Marine Reserve (Hughes
1996). Such protections contributed to
the prevention of dredging a deep water
harbor through turtle nesting beaches
and mining heavy minerals in the
adjacent dunes (Hughes 2009, 2010). In
Mozambique, the Ponta do Ouro Partial
Marine Reserve has provided beach and
marine habitat protection since 2009.
Additional protection is provided to
Mozambican nesting beaches in: The
Ponto du Ouro—Kosi Bay Transfrontier
Marine Conservation Area; the Maputo
Special Reserve; the Bazaruto
Archipelago National Park; and the
Quirimbas Archipelago National Park.
However, nest protection only occurs
over nine percent of the Mozambique
coastline (Videira et al. 2008; Garnier et
al. 2012). Such protections have
minimized vehicular traffic at nesting
beaches in South Africa, but beach
driving remains a threat in
Mozambique. Erosion is a threat to
nesting beaches in both South Africa
and Mozambique. Thus, we conclude
that the present modification of nesting
habitat is a threat to the SW Indian DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization is a threat to the SW
Indian DPS (Bourjea 2015; Williams et
al. 2016; Williams 2017). Two of nine
leatherback turtles equipped with
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satellite tags between 1996 and 2006
were incidentally or intentionally
captured in Mozambique and
Madagascar and likely retained for food
or sale (de Wet 2012). In Mozambique,
eggs and turtles were once legally
harvested and are now illegally poached
for consumption (Nel 2012; Wallace et
al. 2013; Fernandes et al. 2018). Turtle
poaching includes turtles taken on the
beaches and at sea (Williams et al. 2016;
Williams 2017). We do not have recent,
quantitative estimates of egg or turtle
poaching in Mozambique. However,
significant usage has been documented
at various points in time. Hughes (1995)
reported that nearly every nesting
female was killed during the civil war
(1977 to 1992). An estimated 32
loggerhead and leatherback turtles were
killed at Ponta Malongane in 11 years
(Louro 2006). Recent egg and turtle
poaching rates in Mozambique have
been qualitatively described as
‘‘alarming,’’ ‘‘significant,’’
‘‘widespread,’’ ‘‘prominent,’’ and
‘‘prevalent’’ (Fernandes et al. 2015;
Williams et al. 2016; Williams 2017;
Pereira and Louro 2017; Fernandes et al.
2017; Fernandes et al. 2018). Nest
monitoring programs in Mozambique
have provided some protection since the
1990s (Garnier et al. 2012). Pereira et al.
(2014) reports that as a result of the
monitoring program at the Ponta do
Ouro Partial Marine Reserve, where the
majority of nesting in Mozambique
occurs, turtle mortalities are very rare.
Egg poaching has been reduced in the
Bazaruto Archipelago, where it was
previously prevalent (Louro 2006).
National legislation in Mozambique
include: Diploma Legislativo 2627 (7
August 1965), Forest and Wildlife
Regulation (Decree 12/2002 of 6 June
2002) and Conservation Law (Law 5/
2017 of 11 May). These laws protect
turtles and eggs and impose fines for
poaching or possession. However, the
laws are poorly implemented and
enforced (Costa et al. 2007; Louro 2006;
Williams et al. 2016; Fernandes et al.
2018). We conclude that the poaching of
turtles and eggs remains a significant
threat in Mozambique.
Poaching of turtles is also a threat in
Madagascar, where leatherback turtles
caught in gillnets are taken back to local
villages and consumed, which is
documented to have occurred twice in
2016 (Williams 2017). Leatherback
turtles were caught and consumed or
sold in five of eight Malagasy villages
surveyed between October 2004 and
March 2004. Fishers reported that
leatherback turtles were uncommon but
large, possibly indicative of mature
individuals (Walker and Roberts 2005).
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No leatherback turtles were reported
caught during a 2007 Malagasy village
survey (Humber et al. 2010). Although
protected by Presidential Decree (2006–
400), fishers target turtles at sea for
consumption (Ratsimbazafy 2003; Epps
2006; Humber et al. 2010). Humber et al.
(2010) report that the Malagasy law is
not adequately implemented due to lack
of enforcement, a reluctance to manage
the local, cultural fishery, and the size
of the coastline (Rakotonirina and Cooke
1994; Okemwa et al. 2005). We
conclude that the poaching of turtles
remains a significant threat in
Madagascar.
Egg and turtle poaching does not
appear to be a significant threat in South
Africa. Prior to the ban on egg harvest
in 1963, substantial numbers of
leatherback eggs in South Africa were
harvested, likely contributing to the
critically low number of nesting females
at that time (Nel et al. 2015). Hughes et
al. (1996) concluded that nesting
females were not harvested. As a result
of the ban, and with a lucrative tourism
industry centered on the nesting turtles,
egg and turtle harvest has been nearly
eliminated (Hughes et al. 1996). Nesting
females and hatchlings receive
‘‘intensive and effective’’ protection, as
most nesting beaches fall within the
iSimangaliso Wetland Park (Nel et al.
2015). Such beach protections have
been key to recovering the number of
nesting females to current levels
(Hughes et al. 1996; Saba et al. 2015;
Nel et al. 2015). We conclude that the
poaching of turtles and eggs is not a
significant threat in South Africa.
Exposure to poaching is low in South
Africa, where the majority of females
nest. Few females nest in Mozambique,
reducing the DPS’s overall exposure to
egg and nesting female poaching during
nesting. However, turtles regularly
forage in the Mozambique Channel,
where they may be poached along the
coasts of Mozambique and Madagascar.
Poaching of nesting females or postnesting females (i.e., on land or at sea)
reduces both abundance (through loss of
nesting females) and productivity
(through loss of reproductive potential).
Such impacts are high because they
directly remove the most productive
individuals from DPS, reducing current
and/or future reproductive potential.
Egg poaching reduces productivity. We
conclude that overutilization, as a result
of poaching of turtles and eggs, poses a
threat to the DPS.
Disease or Predation
While we could not find any
information on disease for this DPS,
predation is a threat to the SW Indian
DPS. In South Africa, nest predators
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include feral dogs, side-striped jackals,
honey badgers, and ghost crabs (Hughes
1996; Nel 2006). In the 1960s, the
removal of feral dogs greatly reduced
nest predation. Similarly, jackals were
once a threat (Hughes 1996). However,
nest predation by jackals has not been
observed for 17 years (R. Nel, pers.
comm. April 15, 2019). Nel (2006)
reports current rates of predation as
relatively low. Nel et al. (2013) reports
that there is no evidence for significant
beach predation on South African
beaches. Describing nest predation as
minimal in South Africa, de Wet (2012)
found that 15.7 percent of nests were
depredated in the 2009/2010 and 2010/
2011 nesting seasons; ants and ghost
crabs were the main cause of egg
mortality. During the two seasons, ghost
crabs consumed 3.2 percent of
hatchlings as they made their way to the
sea (de Wet 2012).
While all eggs and hatchlings have
some exposure to predation, the species
compensated for a certain level of
natural predation by producing a large
number of eggs and hatchlings. For this
DPS, the primary impact is to
productivity (i.e., reduced egg and
hatching success). We conclude that,
though much reduced, predation still
poses a threat to the SW Indian DPS.
Inadequacy of Existing Regulatory
Mechanisms
The SW Indian DPS is protected to
some degree by several regulatory
mechanisms. For each, we review the
objectives of the regulation and to what
extent it adequately addresses the
targeted threat.
Despite efforts to reduce impacts,
fisheries bycatch continues to be the
primary threat to this DPS (Petersen et
al. 2009; Nel et al. 2013; Wallace et al.
2013; Fossette et al. 2014; Angel et al.
2014; Nel et al. 2015; Harris et al. 2018).
To minimize the impacts from longline
fisheries, the FAO published guidelines
for sea turtle protection, entitled
Technical Consultation on Sea TurtleFishery Interactions (FAO 2004; Huang
and Liu 2010). The UN 1995 Code of
Conduct for Responsible Fisheries (FAO
2004) provides guidelines for the
development and implementation of
national fisheries policies, including
gear modification (e.g., circle hooks, fish
bait, deeper sets, and reduced soak
time), new technologies, and
management of areas where fishery and
sea turtle interactions are more severe.
The guidelines stress the need for
mitigation measures, data on all
fisheries, fishing industry involvement,
and education for fishers, observers,
managers, and compliance officers (FAO
2004; Honig et al. 2007). These
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guidelines, however, are rarely enacted
in full. The ICCAT has adopted a
resolution for the reduction of sea turtle
mortality (Resolution 03–11),
encouraging States to submit data on sea
turtle interactions, release sea turtles
alive wherever possible, and conduct
research on mitigation measures. The
responsibility to implement mitigation
measures remains within each nation,
and many nations have not
implemented such measures (Honig et
al. 2007). South Africa, Namibia, and
Angola signed the Memoranda of
Understanding concerning Conservation
Measures for Marine Turtles of the
Atlantic Coast of Africa. Though South
African vessels are required to carry a
dehooker and line-cutter (Honig et al.
2007) and has instituted an observer
program (Petersen et al. 2009), few other
at-sea conservation measures have been
implemented (Honig et al. 2007). For
Taiwanese fishing vessels operating
within the range of this DPS, Taiwan
has regulations to limit the number of
vessels in the area and to require vessels
to carry de-hookers. However, bycatch
and mortality remain high (Huang and
Liu 2010). Similarly, though the extent
of shark nets off South African beaches
has been reduced from 44 km in the
early 1990s to 23 km in 2007, bycatch
and mortality continue to occur (Brazier
et al. 2012), and Nel et al. (2015)
identify bather protection nets, together
with boat strikes, as the second greatest
threat to the DPS, after longline
fisheries. Regarding shark nets, Brazier
et al. (2012) concludes that bycatch is
low and rates are stable, but because the
leatherback population is small, a
further reduction in bycatch is
desirable. Because the offshore longline
fishery contributes more than the shark
nets to leatherback mortality, Brazier et
al. (2012) also recommends further
introduction of bycatch reduction
techniques in the longline fishery.
Because longline threats are
proportionally large and possibly
increasing, Harris et al. (2018)
concludes that bycatch mitigation
measures in this industry remain first
and most important management action.
Thus, existing regulations have been
inadequate to meet their objectives.
Beach habitat is protected throughout
a portion of the nesting range of this
DPS. In South Africa, approximately
280 km of nesting beaches benefit from
intensive and effective protection as
part of the iSimangaliso Wetland Park,
a World Heritage Site since 1999 (UN
Educational, Scientific and Cultural
Organization 1999; Nel et al. 2015).
iSimangaliso includes 280 km of
beaches, rocky shores, mangroves, lakes,
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estuaries, and coastal waters out to three
nautical miles (5 km) and 200 m depth.
Regulations prevent coastal
development and commercial fishing
within this area. However, Harris et al.
(2015) estimated that 66 percent of
leatherback turtles nest outside of the
protected monitoring area (i.e., only 300
km of the 900 km nesting area is
monitored and protected). In addition,
leatherback turtles use coastal waters
that are not protected under the marine
reserve. In Mozambique, much of the
nesting habitat is protected, including:
The Ponto du Ouro—Kosi Bay
Transfrontier Marine Conservation Area;
the Maputo Special Reserve; the
Bazaruto Archipelago National Park;
and the Quirimbas Archipelago National
Park. However, nest protection only
occurs over nine percent of the
Mozambique coastline (Videira et al.
2008; Garnier et al. 2012). Thus,
regulations to protect the nesting habitat
of the DPS have been successful.
However, leatherback turtles nesting
outside these areas receive no
protection.
In addition, South Africa hosts several
marine protected areas and has
proposed to add 20 new marine
protected areas to expand protection to
five percent of its EEZ (https://
www.marineprotectedareas.org.za/).
Two of these were proposed in order to
protect leatherback marine habitat: The
1200 km2 iSimangaliso Marine
Protected Area (off nesting beaches);
and the 6200 km2 Agulhas Front Marine
Protected Area (encompassing core
foraging habitat). These initiatives are
likely to protect leatherback turtles
within the proposed areas. However, the
DPS has a large range that extends well
beyond protected areas. Harris et al.
(2018) identifies the Mozambique
Channel as an additional key priority
area to protect.
In South Africa, a 1963 ban on egg
and turtle harvest has been effective in
virtually eliminating overutilization
(Hughes 1996). The current law,
Regulation 58(7) of the MLRA (1998),
provides full protection to sea turtles
and their products. In Mozambique,
national legislation includes: Diploma
Legislativo 2627 (7 August 1965), Forest
and Wildlife Regulation (Decree 12/
2002 of 6 June 2002) and Conservation
Law (Law 5/2017 of 11 May). These
laws protect turtles and eggs and impose
fines for poaching or possession. For
example, the Forest and Wildlife
regulation prohibits the killing of turtles
and the possession of their eggs, with
fines up to US $1,000 (Decree 12/2002
of 6 June 2002; Costa et al. 2007). In
2008, there were at least 13 conservation
programs focusing on protection and
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education. Despite these efforts, illegal
poaching of eggs and turtles remains
prevalent in Mozambique (Fernandes et
al. 2014) due to limited implementation
and enforcement of the environmental
legislation (Costa et al. 2007; Louro
2006; Williams et al. 2016; Fernandes et
al. 2018). In Madagascar, all sea turtles
are protected from exploitation by
Presidential Decree (2006–400).
However, fishers continue to target and
consume turtles captured at sea
(Ratsimbazafy 2003; Epps 2006; Humber
et al. 2010). The effectiveness of the
Malagasy law is limited due to lack of
enforcement, a reluctance to manage the
local, cultural fishery, and the size of
the coastline (Rakotonirina and Cooke
1994; Okemwa et al. 2005; Humber et al.
2010). Thus, while regulations to
prevent the harvest of turtles and eggs
have been adequate in South Africa,
regulatory protections in Mozambique
and Madagascar are inadequate.
In summary, numerous regulatory
mechanisms protect leatherback turtles,
eggs, and nesting habitat throughout the
range of this DPS. Though the regulatory
mechanisms provide some protection to
the species, many do not adequately
reduce the threat that they were
designed to address, generally as a
result of limited implementation or
enforcement. As a result, bycatch,
incomplete nesting habitat protection,
and poaching in Mozambique and
Madagascar remain threats to the DPS.
In summary, we consider the
inadequacy of the regulatory
mechanisms to be a threat to the SW
Indian DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat
to the SW Indian DPS (Wallace et al.
2013; Fossette et al. 2014; Angel et al.
2014; Nel et al. 2015; Harris et al. 2018).
Bycatch occurs in commercial and
artisanal, coastal and pelagic fisheries.
Gear types include: Longline, purse
seine, pelagic trawl, shrimp trawl,
gillnets, and beach seines (Honig et al.
2007; Petersen et al. 2009; Nel et al.
2013; Nel et al. 2015).
Of all gear types, longline fisheries
likely have the largest impact on the
DPS (Petersen et al. 2009; Nel et al.
2013; Angel et al. 2014; Nel et al. 2015;
Harris et al. 2018). Leatherback turtles
are exposed to longline fisheries
throughout their foraging range,
including the Benguela Current in the
Atlantic Ocean, the Agulhas Current in
the Indian Ocean, and coastal waters off
South Africa, Mozambique, and
Madagascar (Honig et al. 2007; Peterson
et al. 2009; Huang and Liu 2010; Harris
et al. 2018). Flag states include: South
Africa, Mozambique, Japan, and Taiwan
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(Honig et al. 2007; Peterson et al. 2009;
Huang and Liu 2010).
Harris et al. (2018) found a positive,
significant relationship between the
longline fisheries’ extent of overlap with
leatherback migratory corridors and
threat intensity (F1,8 = 184.7, P <0.001,
R2 = 0.95), which was defined as a
product of the turtles utilization
distribution and the normalized fishing
effort. They concluded that incidental
capture in longline fisheries was the
most important offshore threat to
leatherbacks and supports the
hypothesis that longlining is
suppressing growth of this DPS (Nel et
al. 2013; Harris et al. 2018). Harris et al.
(2018) calculated longline bycatch rates,
around Southern Africa, to be 1,500
leatherback turtles annually. Though
this estimate likely includes turtles from
other DPSs (SE Atlantic and NE Indian),
the authors concluded that even low
absolute bycatch has a
disproportionately large effect in
slowing population growth rates, due to
the small nesting female abundance of
the SW Indian DPS (Harris et al. 2018).
Additional reason for concern is that the
threat intensity of longlining was
especially high in the last 5 years of the
study (ICCAT and IOTC data from 2004
to 2013), suggesting that the threat and
its impacts on the DPS are increasing
(Harris et al. 2018). Throughout the SE
Atlantic and SW Indian Oceans, Harris
et al. (2018), Wallace et al. (2013),
deWet (2012), Thorson et al. (2012), and
Peterson et al. (2009) analyze longline
bycatch over a large portion of the DPS’s
foraging range. Wallace et al. (2013)
categorize the longline fishing effort as
medium to high and conclude that such
effort leads to a high risk and high
bycatch impact for the SW Indian DPS.
Thorson et al. (2012) used data from the
IOTC (1954 to 2009) and South African
fishery (2006 to 2009) in a model of
leatherback turtle survival and
availability. Their model did not find
that leatherback survival declined
during the period when longline fishing
effort increase. However, the authors
state that their null result could be
explained by an imprecise index of
longline effort or using newer bycatch
rates for the South African longline
fishery (i.e., Petersen et al. 2009). For
example, based on fisheries data from
30 South African and Asian pelagic
longline vessels operating in the South
African EEZ between 2006 and 2010, De
Wet (2012) estimates the mean annual
bycatch to be 7.8 (±7.8 S.D.) leatherback
turtles, based on 39 leatherback turtle
captures reported over 5 years. Other
studies estimate bycatch to be higher.
Based on extrapolations from
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independent observer bycatch reports
from 1998 to 2005 (n = 2,256 sets),
Peterson et al. (2009) estimates that the
South African pelagic longline fishery
for tunas and swordfish captures 50
leatherback turtles annually, many of
which likely belong to the SW Indian
DPS (the remainder belong to the SE
Atlantic DPS). Though most (84 percent)
were caught alive, Peterson et al. (2009)
estimates the long-term survival of
affected turtles at 50 percent (based on
an estimated range of 25 to 75 percent;
Aguilar et al. 1995). Peterson et al.
(2009) thus estimates total mortality
from the South African pelagic longline
fishery to be 25 turtles annually, or
around two percent of the total
population (based on a total population
size of 1,200 leatherback turtles), which
they conclude is enough to hamper
recovery of the SW Indian population.
Nel et al. (2013) agrees with this
conclusion, citing a 30 year (1965 to
1995) increasing trend in nesting female
abundance that stalled as the longline
fishery expanded from 1990 to 1995.
Huang and Liu (2010) come to a similar
conclusion. They report that the
longline fishery operated at a relatively
low level until 1995, when South
Africa, Japan, and Taiwan started a joint
venture fishing program.
In the Indian Ocean, Huang and Liu
(2010) evaluated the Taiwanese longline
fishery bycatch, and Louro (2006)
described illegal longlining in
Mozambique waters. Huang and Liu
(2010) evaluated observer data from 77
trips (4,409 sets) on Taiwanese largescale longline fishing vessels. They
identified 84 leatherback turtles
captured from 2004 to 2008, with 48
mortalities (57 percent; Huang and Liu
2010). Extrapolating to the entire
Taiwanese longline fishery in the Indian
Ocean, they estimated an average
bycatch of 173 leatherback turtles
between 2004 and 2007. This number
likely included individuals from the SW
and NE Indian DPSs. In addition to
commercial longlining, artisanal
longlining also occurs in the SW Indian
Ocean. Illegal longlining off
Mozambique targets sharks and
leatherback turtles. The level of take and
mortality is unknown. A program called
Eyes on the Horizon reports such
events, when observed (Louro 2006).
In the SE Atlantic Ocean, Honig et al.
(2007) and Angel et al. (2014) evaluate
longline bycatch. Honig et al. (2007)
evaluated turtle bycatch by longline
fisheries in the Benguela Large Marine
Ecosystem by using data from observer
reports, surveys, and specialized trips
from the coastal nations of South Africa,
Namibia and Angola. They estimated
bycatch at 672 leatherback turtles
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annually (based on an annual bycatch
estimate of 4,200 turtles, of which
approximately 16 percent are
leatherback turtles) in the southern and
central regions and as many as 5,600
leatherback turtles (based on an annual
bycatch estimate of 35,000 turtles) for
the entire Benguela Large Marine
Ecosystem (Honig et al. 2007). These
estimates likely include many
leatherback turtles from the much larger
SE Atlantic DPS, but telemetry studies
indicate that the turtles of the SW
Indian DPS use this foraging area too
(Luschi et al. 2006; Robinson et al.
2016). Evaluating ICCAT data, Angel et
al. (2014) confirms exposure to high
longline fishing effort but reports that
bycatch of this population is low
relative to other leatherback
populations. Although Thorson et al.
(2012) found that increased fishing
effort had no explanatory power
regarding changes in leatherback
survival, other studies identify longline
fisheries as the primary threat to the
DPS (Petersen et al. 2009; Nel et al.
2013; Angel et al. 2014; Nel et al. 2015;
Harris et al. 2018). Based on the weight
of evidence, we agree with the latter and
conclude that longline fisheries pose a
major threat to the DPS throughout its
foraging range.
Other fisheries also impact the SW
Indian DPS, possibly resulting in
substantial mortalities. However, these
fisheries are not as well studied, and
mortality estimates are not available
(Honig et al. 2007; Nel et al. 2013).
Leatherback turtles are caught in
artisanal and commercial shrimp trawl,
pelagic trawl, gillnet, purse seine, and
beach seine fisheries (Honig et al. 2007;
Petersen et al. 2009; Nel et al. 2013).
Citing Walker (2005) and Rakotonirina
(1994), Nel (2013) reports that the
number of sea turtles (all species)
caught in artisanal fisheries of the
Mozambique Channel could exceed
commercial fishery catches. Honig et al.
(2007) echoes this concern for the
Benguela Current Large Marine
Ecosystem, citing high mortality rates
for these fisheries in other regions. The
Mozambican shrimp trawl fishery
operates in the Sofala Bank of the
Mozambique Channel, near leatherback
nesting, migrating, and foraging areas
(Luschi et al. 2006; Robinson et al.
2016). The fishery supports 50 to 96
vessels that employ standard otter trawl
nets in a single or quad-net
configuration with an average tow-time
of three hours (Brito 2012). It does not
employ TEDs and incidentally captures
several (i.e., at least two to six but
possibly many more) leatherback turtles
annually (Louro 2006; Videira et al.
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2010; SWOT 2017). In 2001, one shrimp
trawler captain reported capturing more
than six leatherback turtles since fishing
season opened; all were captured alive
(Gove et al. 2001). Based on 39
interviews with observers, enforcement
officers, and vessel operators, the fleet
(n = 50) captures approximately 56 (±40)
leatherback turtles; the overall estimated
mortality rate for bycaught turtles is 14
percent (Brito 2012). Given the overlap
between the fishery and an important
foraging area, M. Pereira (CTV, pers.
comm., 2019) concludes that the
Mozambican shrimp trawl fishery may
be one of the main threats to this DPS.
The South African shrimp trawl fishery
has been reduced to two vessels, with
an average annual bycatch of less than
one leatherback (Honig et al. 2007;
Petersen et al. 2009; Nel et al. 2013).
Domestic shrimp trawling in Eritrea is
considered a major threat to sea turtles,
and bycatch is underreported. However,
leatherback turtles are relatively rare in
these waters, as demonstrated by the
foreign trawl fleet, which has 100
percent observer coverage and bycatch
records indicating 39 leatherback turtles
between 1996 and 2005 (Pilcher et al.
2006).
During a small random sampling
exercise in 2013 by onboard observers
from the Research Division of Eritrea,
one leatherback turtle (of 48 sea turtles
total) was captured and released
(Mebrahtu 2015). On June 20, 2019, the
European Union passed a regulation
(PE–CONS 59/1/19 Rev 1) that requires
shrimp trawl fisheries to use a turtle
excluder device in European Union
waters of the Indian and West Atlantic
Oceans.
Gillnets in Macaneta, Mozambique,
killed two leatherback turtles during the
2010 nesting season (Videira et al. 2010)
and captured one in the 2003 nesting
season (Louro 2006). In Madagascar,
leatherback turtles are a ‘‘common’’
bycatch of the set gillnet shark fishery
(Robinson and Sauer 2013); mortality is
likely high given the 24-hour soak time
and propensity for consuming turtle
meat. Purse seine fisheries have a much
lower impact than longline fisheries
(Angel et al. 2014); two leatherback
turtles were captured (alive) between
1995 and 2010 in the Indian Ocean
(Clermont et al. 2012). In the EEZ of all
Indian Ocean French Territories (mostly
from the Mozambique Channel), 40
leatherback turtles were captured in
unspecified fisheries from 1996 to 1999;
92 percent were released alive (Ciccione
2006).
Shark or bather nets, which are
gillnets installed off beaches in South
Africa to limit human-shark
interactions, incidentally capture
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leatherback turtles. According to Nel et
al. (2015), bather protection nets and
boat strikes together present the second
greatest threat to the DPS, after fisheries.
Three of nine leatherback turtles
equipped with satellite tags between
1996 and 2006 were caught in shark
nets (de Wet 2012). Between 1981 and
2008, 150 leatherback turtles were
captured (mean = 5.36; SE = 0.60), of
which 20 were mature females and 39
were mature males (Brazier et al. 2012).
Total mortality was 62.7 percent, with
an annual range of 1 to 12 mortalities
(mean = 3.4; SE = 0.47; Brazier et al.
2012). Most turtles were captured in
December, the peak month for nesting,
which together with the prevalence of
mature individuals, suggests that
bycatch is dominated by adults from
nearby nesting and breeding areas
(Brazier et al. 2012). Analyzing these
data over an additional 2 years (1981 to
2010), de Wet (2012) found that 157
leatherback turtles (mean = 5.26; SD =
2.7) were captured in the nets, with a
62.4 percent mortality rate (mean = 3.3;
SD = 1.8).
To reduce bycatch mortality in
longlines, South African regulations
require vessels to carry a dehooker and
line cutter (Honig et al. 2007). To reduce
bycatch in the shark nets, effort was
reduced from 44 km of nets in the early
1990s to 23 km in 2007 (Brazier et al.
2012). Despite these efforts, a previously
increasing trend in nesting female
abundance has stalled and ‘‘declined
recently’’ (Nel et al. 2013).
Individuals (immature and adult
turtles) of this DPS are exposed to high
fishing effort throughout their foraging
range. Estimates of bycatch rates, when
available, range considerably. For
example, Harris et al. (2018) estimated
the annual longline bycatch rates
around Southern Africa to be 1,500
leatherback turtles annually; whereas,
de Wet (2012) estimated the mean
annual bycatch to be 7.8 (±7.8 S.D.)
leatherback turtles. We have annual
mortality estimates for few individual
fisheries: n = 25 for South African
longline (Peterson et al. 2009); n = 12 for
Taiwanese longline (Huang and Liu
2010); n = 1 to 12 for shark nets (Brazier
et al. 2012). Adding in other longline
fisheries and additional gear types may
result in more than 100 mortalities
annually. These estimates likely include
individuals from other DPSs (i.e., the SE
Atlantic and NE Indian). However,
because of the small nesting population,
even small levels of mortality have the
potential to slow population growth
(Harris et al. 2018). Mortality reduces
abundance, by removing individuals
from the population; it also reduces
productivity, when potential nesting
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females are killed. Several studies
conclude that bycatch has prevented
continued population growth and/or
contributed to the recent slight decline
in nesting (Petersen et al. 2009; Huang
and Liu 2010; Brazier et al. 2012; Nel et
al. 2013; Harris et al. 2018). We
conclude that fisheries bycatch is the
primary threat to the SW Indian DPS.
Vessel Strikes
Vessel strikes are a threat to the SW
Indian DPS. According to Nel et al.
(2015), vessel strikes and bather
protection nets together present the
second greatest threat to the DPS, after
fisheries. Together these threats kill up
to 10 leatherback turtles annually (Nel
et al. 2015). One of 24 leatherback
turtles stranded along the South African
coastline between 1972 and 2010 was
struck by a boat propeller (Nel 2008).
However, additional mortalities or
injuries may go unnoticed or
unreported. Vessel strikes affect adult
females returning to nest, removing
individuals and their future
reproductive potential. Thus, this threat
reduces the abundance and productivity
of the DPS. We conclude that vessel
strikes pose a threat to the DPS.
Pollution
Pollution includes contaminants,
marine debris, and ghost fishing gear.
As with all leatherback turtles,
entanglement in and ingestion of marine
debris and plastics are threats that likely
kill several individuals a year. For six
stranded hatchlings and 24 stranded
adults over the past 40 years, the cause
of death was generally unknown.
However, fishery-related injuries, ghostfishing (i.e., entanglement in discarded
fishing gear), disease, or pollution may
be responsible (de Wet 2012). Plastic
pollution may be a main threat in the
waters off Mozambique (M. Pereira,
pers. comm., 2019). Outer accumulation
of the Indian Ocean ‘‘garbage patch’’
(Cozar et al. 2014) overlaps with
foraging areas in the Mozambique
Channel and occurs in waters offshore
from nesting areas in South Africa and
Mozambique. Though we were unable
to find ingestion or entanglement data
for SW Indian leatherback turtles, 51.4
percent of gut and fecal samples from
loggerhead turtles (n = 74) captured as
bycatch in the Reunion Island longline
fishery contained marine debris, of
which plastic comprised 96.2 percent
(Hoarau et al. 2014). Ryan et al. (2016)
found that 24 of 40 loggerhead turtle
post-hatchlings had ingested plastics or
other anthropogenic debris. Based on
the foraging behavior of leatherback
turtles and the proximity of the ‘‘garbage
patch,’’ we conclude that the ingestion
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and entanglement of marine debris are
threats to this DPS.
In addition, State of the World’s Sea
Turtles (SWOT 2017) identifies
hydrocarbon extraction along the
eastern African seaboard, including
northern Mozambique, as the greatest
emerging concern for this DPS. They
report that the impact of such activities
remain to be seen (SWOT 2017).
However associated oil spills are likely
to modify habitat off nesting beaches
and reduce prey availability for all life
stages. Harris et al. (2018) found that the
hydrocarbon industry poses a moderate
threat to the DPS because of its spatial
overlap with migratory corridors
(second in extent, after longline
fisheries). They expressed concern over
the expansion of the hydrocarbon
extraction along the coasts of southern
Mozambique and northeastern South
African and the possibility of an oil spill
in these areas (Harris et al. 2018).
Pretorius (2018) identified 28 significant
impacts to sea turtles as a result of
hydrocarbon exploration and
production; these included: Potential
water pollution, light pollution, noise
pollution, and habitat destruction.
However, Du Preez et al. (2018) reports
that metal and metalloid contaminants
do not appear to be a problem for this
DPS. We conclude that pollution poses
a threat to the DPS.
Climate Change
Climate change is a threat to the SW
Indian DPS. The impacts of climate
change include: Increases in
temperatures (air, sand, and sea
surface); sea level rise; increased coastal
erosion; more frequent and intense
storm events; and changes in ocean
currents.
Angel et al. (2014) identifies coastal
erosion as the main beach-based threat
to this population and one that is likely
to increase with climate change. Though
coastal erosion is a natural process, sea
level rise (as a result of climate change)
increases the rate of erosion and the
amount of beach affected. In Bazaruto
Archipelago, Mozambique, coastal
erosion and rising sea levels destroyed
approximately 12 percent of nests over
10 seasons of monitoring (Videira and
Louro 2005; Louro 2006). Because
leatherback turtles nest lower on the
beach than other sea turtles, their eggs
are more at risk of being exposed and
destroyed by increases in sea level and
coastal erosion (Boyes et al. 2010).
Thus, erosion and rising sea level as a
result of climate change are a threat to
the DPS.
Sand temperatures influence
leatherbacks’ egg viability and sex
determination. Temperatures over 32 °C
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result in death and temperatures below
29.2 °C produce only males (Rimblot et
al. 1985; Rimblot-Baly et al. 1986).
Temperature probes on South African
beaches reveal that nests are already
close to pivotal temperatures, with an
average of 29.04 °C (S.D. 0.86 °C; range
27.62 to 29.69 °C; Boyes et al. 2010). A
modeling study suggests that even if
South African beaches experience a
temperature increase of 5 °C, hatching
success and emergence success may not
be significantly reduced (Santidria´n
Tomillo et al. 2015). Instead, nesting
females may shift their nesting season to
months (e.g., July through October)
when temperature and precipitation
would be similar to current conditions
of the current nesting season (i.e.,
October through January). However, the
authors cautioned that because nesting
females do not change their nesting
habits in response to oceanographic
conditions, they may not change their
nesting habits in response to climate
change either (Santidria´n Tomillo et al.
2015). In addition, a shift in the nesting
season could have impacts beyond
hatching success, such as reduced posthatchling survival and suboptimal
foraging conditions for post-nesting
females. We therefore conclude that
increased temperatures may be a threat
to the DPS, and will likely result in
impacts ranging from nesting season
shifts to significant nest losses.
The threat of climate change may
modify the nesting conditions for the
entire DPS. Impacts likely range from
small, temporal changes in nesting
season to large losses of productivity.
Because we are already seeing small
impacts due to coastal erosion and sea
level rise, we conclude that climate
change is a threat to the SW Indian DPS.
Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply to
the SW Indian DPS (for a description of
each effort, please see the section on
conservation efforts for the overall
taxonomic species): African Convention
on the Conservation of Nature and
Natural Resources (Algiers Convention),
Convention on the Conservation of
Migratory Species of Wild Animals,
Convention on Biological Diversity,
Convention on International Trade in
Endangered Species of Wild Fauna and
Flora, Convention on the Conservation
of European Wildlife and Natural
Habitats, Convention for the Cooperation in the Protection and
Development of the Marine and Coastal
Environment of the West and Central
African Region (Abidjan Convention)
and Memorandum of Understanding
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Concerning Conservation Measures for
Marine Turtles of the Atlantic Coast of
Africa (Abidjan Memorandum),
Convention Concerning the Protection
of the World Cultural and Natural
Heritage (World Heritage Convention),
FAO Technical Consultation on Sea
Turtle-Fishery Interactions, Indian
Ocean Tuna Commission, The Indian
Ocean Tuna Commission, Indian
Ocean—South-East Asian Marine Turtle
Memorandum of Understanding,
MARPOL, IUCN, Nairobi Convention for
the Protection, Management and
Development of the Marine and Coastal
Environment of the Eastern African
Region, Ramsar Convention on
Wetlands, UNCLOS, and UN Resolution
44/225 on Large-Scale Pelagic Driftnet
Fishing. Although numerous
conservation efforts apply to the turtles
of this DPS, they do not adequately
reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the SW Indian DPS is at high risk of
extinction. The DPS exhibits a total
index of nesting female abundance of
149 females. Such a limited nesting
population size places this DPS in
danger of stochastic or catastrophic
events that increase its extinction risk.
This DPS exhibits a slightly decreasing
nest trend at monitored nesting beaches
in South Africa. This declining trend
has the potential to further lower
abundance and thereby increase the risk
of extinction. With only one nesting
aggregation, the DPS lacks spatial
structure, and its genetic diversity is
low. Thus, stochastic events could have
catastrophic effects on nesting for the
entire DPS, with no potential source
subpopulations to buffer losses or
provide additional diversity. However,
the DPS uses multiple, distant, and
diverse foraging areas, providing some
resilience against reduced prey
availability. Based on these factors, we
find the DPS to be at risk of extinction,
likely as a result of past threats.
Current (ongoing) threats further
contribute the risk of extinction of this
DPS. The primary threat to this DPS is
bycatch in commercial and artisanal,
pelagic and coastal, fisheries. Longline
fisheries constitute the greatest threat.
Though poorly studied, other fisheries
together may have overall mortality
rates for affected turtles from this DPS
that rival those from longline fisheries.
Fisheries bycatch reduces abundance by
removing individuals from the
population. Because several fisheries
operate near nesting beaches,
productivity is also reduced when
nesting females are prevented from
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48381
returning to nesting beaches. Exposure
and impact of this threat are high.
Poaching is also a threat to the DPS. Egg
and turtle poaching, while no longer a
threat in South Africa, likely continues
in Mozambique. In Madagascar, turtles
are illegally captured at sea and
consumed in local villages. Vessel
strikes also pose a threat. Vessel strikes
kill several leatherback turtles each
year, including females returning to
beaches to nest. While exposure is low,
impacts are high, affecting both
abundance and productivity. Coastal
erosion and beach driving in
Mozambique modify nesting habitat and
are believed to result in minor
reductions in productivity currently.
However, these threats are likely to
increase over time as climate change
and tourism increases. Climate change
is likely to result in reduced
productivity due to greater rates of
coastal erosion and nest inundation.
Predation of eggs and hatchlings is also
a threat. However, although predation
has the potential to reduce productivity,
the DPS has likely adapted to predation
by native species, which account for
most of the predation at present.
Ingestion of plastics and entanglement
in marine debris are threats to all
leatherback turtles, most likely resulting
in injury and reduced health, though
sometimes mortality. Though many
regulatory mechanisms are in place,
they do not reduce the impact of these
threats to levels that allow the DPS to
continue its previous increasing nesting
trend.
Thus, the Team unanimously
concluded, that the SW Indian DPS is at
high risk of extinction. The total index
of nesting female abundance of 149
females makes the DPS highly
vulnerable to threats. We determine,
consistent with the team’s findings, that
the DPS is currently ‘‘in danger of
extinction.’’ The slightly declining nest
trend and lack of spatial structure and
diversity further contribute to its risk of
extinction. While this small population
had an increasing or stable nesting trend
for decades, the lack of continued
population growth and recent decline
may indicate that threats have outpaced
productivity. Past egg and turtle harvest
initially reduced the nesting female
abundance of this DPS and likely
confined its nesting habitat to a
relatively small geographic area, with
little diversity or spatial structure.
Currently, fisheries bycatch is the
primary present, ongoing threat. It
reduces abundance and productivity
(i.e., imminent and substantial
demographic risks) by removing mature
and immature individuals from the
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population at rates exceeding
replacement. Though numerous
conservation efforts apply to this DPS,
they do not adequately reduce the risk
of extinction. We conclude that the SW
Indian DPS is in danger of extinction
throughout its range and therefore meets
the definition of an endangered species.
The threatened species definition does
not apply because the DPS is at risk of
extinction currently (i.e., at present),
rather than on a trajectory to become so
within the foreseeable future.
NE Indian DPS
The Team defined the NE Indian DPS
as leatherback turtles originating from
the NE Indian Ocean, south of 71° N,
east of 61.577° E, and west of 120° E.
The western boundary occurs at the
border between Iran and Pakistan,
where the Somali Current begins. This
current, and the cold waters of the
Antarctic Circumpolar Current, likely
restrict the nesting range of this DPS.
We placed the eastern boundary at 120°
E to approximate the Wallace and
Huxley lines, which are established
biogeographic barriers to gene flow
between Indian and Pacific Ocean
populations of numerous species. While
the genetic differences between the NE
Indian and West Pacific DPSs
demonstrate discreteness, genetic
sampling is unavailable from areas
where the nesting range of the DPSs
likely meet, preventing us from defining
the boundary more specifically.
The range of the DPS (i.e., all areas of
documented occurrence) extends
throughout the Indian Ocean and
possibly into the Pacific Ocean. Records
indicate that the species occurs in the
waters of the following nations: India,
Sri Lanka, Bangladesh, Myanmar,
Thailand, Malaysia, Indonesia, Vietnam,
China, and Philippines (Hamann et al.
2006). Given the range of the DPS,
leatherbacks may also occur in the
waters of Pakistan, Australia, Brunei,
Cambodia, Philippines, and Taiwan.
Leatherback turtles of the NE Indian
DPS nest on beaches scattered
throughout the NE Indian Ocean. The
largest abundance of nesting occurs on
beaches of the Andaman and Nicobar
Islands in India. The sandy beaches of
the Andaman and Nicobar Islands
consist of soft limestone formed of coral
and shell (Lal 1976; Bandopadhyay and
Carter 2017). A moderate amount of
nesting occurs in Sri Lanka, and even
less occurs in Thailand and Sumatra,
Indonesia (Hamann et al. 2006; Nel
2015).
Information on this DPS is limited,
but foraging appears to occur
throughout the Indian Ocean (Andrews
et al. 2006; Hamann et al. 2006). The
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foraging range extends throughout the
Bay of Bengal, south of Sri Lanka, and
along the west coast of Sumatra,
Indonesia, as indicated by satellite
telemetry data and fisheries reports
(NMFS and FWS 2013). Nesting females
at Little Andaman Island likely use a
variety of foraging areas and have been
tracked to: South and east of the
Andaman and Nicobar Islands; along
the coast of Sumatra; beyond Cocos
(Keeling) Island towards Western
Australia; and across the Indian Ocean
towards Madagascar and the African
continent (Namboothri et al. 2012;
Swaminathan et al. 2017; Swaminathan
et al. 2019). Stranding data also indicate
the use of diverse foraging areas: 15
individuals stranded or were caught in
fishing gear along the mainland coast of
India (Shanker 2013). Leatherback
turtles have also stranded along the
coasts of Mindanao, Philippines and
Pakistan (Firdous 2006; Lucero et al.
2011).
Abundance
The total index of nesting female
abundance of the NE Indian DPS is 109
females. We based this total index on
the nesting aggregations at South and
West Bays, Little Andaman Island, India
(K. Shanker pers. comm., 2018). Our
total index does not include 14
unquantified nesting aggregations in
Bangladesh, India, Indonesia, Malaysia,
Myanmar, Sri Lanka, Thailand,
Philippines, and Vietnam. To calculate
the index of nesting female abundance,
we divided the total number of nests at
South and West Bays, Little Andaman
Island between the 2015/2016 and 2017/
2018 nesting seasons (i.e., a 3-year
remigration interval; Andrews 2002) by
the clutch frequency (3.8 clutches/
season; Andrews 2002; Eckert et al.
2015). This number represents an index
of abundance for this DPS, and is likely
to be an underestimate, because it only
includes available data from recently
and consistently monitored nesting
beaches. Additional nesting occurs at
other locations but is unquantified.
Published estimates of total nesting
female abundance are not available for
this DPS. The IUCN Red List assessment
did not provide an estimate of the total
number of mature individuals because
monitoring was not sufficient (Tiwari et
al. 2013). Currently, the largest nesting
aggregations occur in the Andaman and
Nicobar Islands of India. Nesting in Sri
Lanka may consist of about 100 to 200
nesting females per year, and low levels
of nesting occur in Thailand and
Sumatra, Indonesia (Hamann et al.
2006; Nel 2012). Low and scattered
nesting occurs in Indonesia: 1 to 14
nesting females annually at Alas Purwo
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in East Java; and one to three nesting
females annually on three beaches in
Bali. There are also rare reports of
nesting in the Philippines (Lucero et al.
2011; Arguelles 2013), Vietnam, and
Malaysia. In Myanmar, nesting is rare,
and only one confirmed nesting event
has been recorded in recent years (i.e.,
December 2016; Platt et al. 2017).
Historically, there may have been
nesting in Bangladesh, but no current
reports exist (Hamann et al. 2006).
Malaysia once hosted the DPS’s
largest nesting aggregation (Chan and
Liew 1996). It is now considered
functionally extinct or extirpated
(Pilcher et al. 2013), as a result of
continuous, large-scale egg harvest and
fisheries bycatch (Chan and Liew 1996;
Eckert et al. 2012). The major nesting
site in Malaysia, Rantau Bang in
Terengganu, decreased drastically from
10,000 nests in the 1950s to 10 or fewer
nests in the 2010s (reviewed by Eckert
et al. 2012), and to one or no nests
annually, more recently. The number of
nesting females in Vietnam has also
decreased dramatically, from
approximately 500 nesting females in
the 1960s to two to three nests in 2005
and 2007 (The Chu and Nguyen 2015).
In the late 1970s, females nested in
multiple locations of Thailand,
including: along the airport beach in
Changwat Phuket; in the Laem Phan Wa
marine reserve; and in coastal
Changwan Phangnga (Bain and
Humphry 1980). Settle (1995) recorded
about 30 nests on the Phuket and
Phangnga coastlines from 1992 to 1993.
Aureggi et al. (1999) found nine nests
between 1997 and 1998, during a survey
of Phra Thong Island in the south.
Our total index of nesting female
abundance (109 females) places the DPS
at risk for environmental variation,
genetic complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). These
processes, working alone or in concert,
place small populations at a greater
extinction risk than large populations,
which are better able to absorb losses in
individuals. Due to its small size, the
DPS has restricted capacity to buffer
such losses. Historic abundance
estimates indicate that the DPS was
once much larger. The current
abundance is likely a result of past and
current threats, which we describe
below. Given the intrinsic problems of
small population size, we conclude that
the limited nesting female abundance is
a major factor in the extinction risk of
this DPS.
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Productivity
Diversity
The NE Indian DPS has exhibited a
drastic population decline with
extirpation of its largest nesting
aggregation in Malaysia. Nest counts
from Malaysia exhibited a steep decline
of 17.9 percent annually (sd = 4.2
percent; 95 percent CI = ¥25.5 to ¥8.4
percent; f = 0.998; mean annual nests =
1,166) over the 44-year period of data
collection (1967 to 2010). The drastic
decline of nests observed in Malaysia is
likely representative of the overall trend
for the DPS given the magnitude of
historical abundance for that site and
the high confidence in the trend
estimate.
Despite the dramatic population
decline, driven by the extirpation of the
largest nesting aggregation (i.e.,
Malaysia), productivity parameters are
similar to the species averages.
However, we have a low degree of
confidence in these estimates due to
limited monitoring of existing nesting
aggregations. We conclude that the NE
Indian DPS exhibits a declining nesting
trend, which increases its extinction
risk.
Genetic diversity of the NE Indian
DPS is potentially relatively high, based
on analyses of samples collected from
the previously large, but now
functionally extinct, nesting aggregation
in Malaysia (Dutton et al. 1999, 2007);
genetic diversity has not been assessed
at other nesting sites. The diversity of
nesting sites is low, given that the
majority of the nesting currently occurs
on islands (Sivasundar and Prasad
1996). We conclude that existing
diversity provides little resilience to the
DPS.
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Spatial Distribution
For the NE Indian DPS, nesting is
limited to a few, scattered nesting
beaches. Currently, the majority of the
nesting occurs on beaches of the
Andaman and Nicobar Islands and Sri
Lanka, with small numbers of nests on
the western coast of Thailand, Sumatra,
and Java (Nel et al. 2015).
Spatial structure is unknown but
presumed to be low. There are no
estimates of genetic population
structure within this DPS because
published genotypes only exist for
Malaysia (Dutton et al. 1999, 2007).
Genetic samples were taken from
nesting females at Little Andaman
Island from 2008 through 2010, but the
results are not yet available (Namboothri
et al. 2010).
The wide distribution of foraging
areas likely buffers the DPS somewhat
against local catastrophes or
environmental changes that would limit
prey availability. Remaining nesting is
limited to a few, scattered but broadly
distributed nesting sites. The largest
nesting aggregations are clustered, thus
rendering the DPS susceptible to
environmental catastrophes (e.g.,
tsunamis), and directional changes (e.g.,
sea level rise). Thus, despite widely
distributed foraging areas, the somewhat
limited nesting distribution increases
the extinction risk of the NE Indian
DPS.
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Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
Erosion, coastal development, and
artificial lighting have destroyed or
modified the available, suitable nesting
habitat and thus are threats to the NE
Indian DPS.
Erosion reduces the available nesting
habitat for the DPS. Some erosion
occurs as a result of natural disasters. In
2004, a major earthquake occurred off
the west coast of Sumatra, Indonesia,
resulting in the 2004 tsunami, which
destroyed many of the beaches that once
hosted over 1,000 nests (Subramaniam
et al. 2009). As a result of the tsunami,
the width of the coastline was reduced
by one meter, severely modifying the
beaches of South Bay, Little Andaman
Island, and resulting in very low
leatherback nesting in 2005 and 2006
(Namboothri 2010). The tsunami also
caused drastic changes at other
leatherback nesting beaches (Alfred et
al. 2005; Ramachandran et al. 2005;
Murugan 2005; Andrews et al. 2006). It
caused erosion at some beaches and
accretion at others, especially in the
Andaman and Nicobar Islands, which
lie closest to the epi-center of the
earthquake and host the largest numbers
of nesting females in the DPS
(Subramaniam et al. 2009). In addition,
the beaches in Indonesia are being lost
due to erosion from high tides and
monsoons (Obermeier 2002).
Sand mining and tourism-related
development are the main threats to
nesting habitat (Fatima et al. 2011).
While we were unable to find specific
information regarding sand mining,
coastal development is increasing in Sri
Lanka, India, and Bangladesh. The
beaches of Sri Lanka are under high
threat from tourism development (e.g.,
large hotels and restaurants), urban and
industrial development, and artificial
lighting (Kapurisinghe 2006). Along the
mainland of India, granite blocks and
embankments prevent turtles from
approaching many beaches (Andrews et
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48383
al. 2006). Intense coastal development,
stemming from the tourism industry,
occurs in Bangladesh without
environmental review (Pilcher 2006),
resulting in the alteration of sand dunes
and other activities that reduce the
quality of nesting habitat (Islam 2002;
Islam et al. 2011). In Vietnam,
increasing tourism is expected to result
in coastal development on the beaches
of Son Tra Peninsula, QuanLan, and
Minh Chau (Ministry of Fisheries 2003).
In addition, most Vietnam beaches are
affected by a large amount of marine
debris (e.g., glass, plastics, polystyrenes,
floats, nets, and light bulbs), which can
entrap turtles and impede nesting
activity.
Artificial lighting modifies the quality
of nesting beaches because lights over
land disorient nesting females and
hatchlings. Instead of crawling toward
the surf and their marine habitat, they
crawl further inland, where they may
become dehydrated and die or are
susceptible to predation. Nests moved to
hatcheries as part of conservation efforts
may be subject to inadequate hatchery
practices, which have resulted in
skewed sex ratios and low hatching
success (Chan and Liew 1996;
Kapurisinghe 2006; Rajakaruna et al.
2013; Phillott et al. 2018).
Some areas are protected. Of the 306
islands in the Andaman and Nicobar
Islands of India, 94 are designated as
wildlife sanctuaries, six of which are
national parks, and two of which are
marine national parks (Andrews et al.
2006). In Sri Lanka, in 2006, sea turtle
sanctuaries were established at Rekawa
(4.5 km stretch) and Godawaya (3.8 km
stretch), where high frequency
leatherback nesting is observed; the area
is bounded 500 meters towards the sea
and 100 meters towards the land from
the high tide level in both sites (Phillott
et al. 2018). Although laws protect sea
turtles throughout Sri Lanka, most
nesting areas are not protected and
hence, local communities can disturb
nesting beaches by removing sand,
lighting the beaches, and cutting the
beach vegetation (Phillott et al. 2018). In
Malaysia, turtle sanctuaries have been
established in Terengganu, Sabah, and
Sarawak. However, nesting habitat
modification and destruction continue
in many areas.
We conclude that nesting females,
hatchlings, and eggs are exposed to the
reduction and modification of nesting
habitat, as a result of erosion, coastal
development, and artificial lighting.
These threats impact the DPS by
reducing nesting and hatching success,
thus lowering its productivity. The most
abundant remaining nesting
aggregations are protected from
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development, but they experience high
rates of erosion; other nesting beaches
are subject to anthropogenic threats.
Thus, we conclude that habitat loss and
modification pose a threat to the NE
Indian DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Overutilization is a threat to the NE
Indian DPS. The harvest of turtles and
eggs led to the historical decline of the
DPS, and poaching continues in several
areas (Phillott et al. 2018).
Regular, nearly complete egg harvest
caused the functional extinction of the
once large nesting aggregation in
Malaysia (Chan and Liew 1996). In the
early 1960s, the Terengganu, Malaysia
nesting beaches were leased to the
highest bidder, and nearly all
leatherback eggs were harvested. In the
1980s, the State Fisheries Department
tried to buy back about 10 percent of the
harvested eggs to be incubated in a
hatchery (Siow and Moll 1982; Chan
and Liew 1996; Stiles 2009). However,
such efforts could not prevent the
extirpation. Excessive egg harvest, both
legal and illegal, also caused declines in
India, Sri Lanka, and Thailand (Ross
1982).
The harvest of turtles and eggs
continues but has not been quantified
(Nel 2012). In Sri Lanka, almost all eggs
are taken from the beach and sold at
markets or to hatcheries for ecotourism
(Kapurusinghe 2000, 2006; Rajakaruna
et al. 2013; Phillott et al. 2018). The
conservation benefit provided by
hatcheries in Sri Lanka is debatable
(Phillott et al. 2018) because they do not
follow the hatchery practices
established by the IUCN (Hewavisenthi
1994; IUCN 2005; Namboothri et al.
2012; Rajakaruna et al. 2013; Phillott et
al. 2018). Egg harvest also continues in
Thailand. Commercial egg harvest is
illegal in the Andaman and Nicobar
Islands, and in the Andaman Islands, a
ban on hunting and harvesting of turtles
came into force in 1977. However, the
original inhabitants of the Andaman and
Nicobar Islands are exempt from the
Indian Wildlife Protection Act (Shanker
and Andrews 2004), and Namboothri et
al. (2012) observed intense egg harvest
at Delgarno, Trilby, and East Turtle
Islands. In Myanmar, despite
regulations prohibiting the consumption
of turtle meat and eggs (Hamann et al.
2006), there is illegal trade of turtles
caught at sea, including leatherback
turtles (Murugan 2007). In Sri Lanka,
the historically high direct take of
turtles at sea has been reduced
(Kapurushinghe 2006). Records indicate
that turtle meat and parts were once
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regularly exported from Tamil Nadu,
India, to Sri Lanka, and then to other
nations such as the United States,
Singapore, and Belgium (Kuriyan 1950;
Chari 1964; Shanmughasundarun 1968
as cited in Agastheesapillai and
Thiagarajan 1979).
Exposure to egg and turtle poaching
remains high throughout the range of
the DPS. Poaching of nesting females or
post-nesting females at sea reduces both
abundance (through loss of nesting
females) and productivity (through loss
of reproductive potential). Such impacts
are high because they directly remove
the most productive individuals from
the DPS, reducing current and future
reproductive potential. Egg harvest
reduces productivity only, but, as
previously demonstrated within this
DPS, can have devastating populationlevel impacts. We conclude that
overutilization, as a result of egg and
turtle harvest, poses a major threat to
the NE Indian DPS.
Disease or Predation
While we could not find any
information on disease for this DPS, the
best available data indicate that
predation is a threat to the NE Indian
DPS. Multiple predators prey on eggs
and hatchlings at several nesting
beaches (Andrews 2000). During a 2016
survey of the Nicobar Islands,
approximately 57 percent (n = 1,223) of
leatherback nests were lost to
depredation by feral dogs, water
monitor lizards, or feral pigs (Sus
domesticus; Swaminathan et al. 2017).
In the South Bay of Great Nicobar
Island, wild boars and dogs prey on
eggs, while fiddler crabs, dogs, and
raptors prey on hatchlings (Sivakumar
2002). Sivasundar and Prasad (1996)
documented that Asian water monitor
lizards took 68.6 percent of leatherback
nests in the Andaman Islands. In Sri
Lanka, egg predators include feral dogs,
water and land monitor lizards, jackals,
wild boars, mongooses, and ants. Egg
predation by feral pigs is a major threat
in Indonesia (Maturbongs et al. 1993;
Maturbongs 1995, 1996; Sivasundar and
Prasad 1996).
A large number of eggs and hatchlings
are exposed to predation. Though
leatherback turtles produce a large
number of eggs and hatchlings,
published rates of predation (57 to 69
percent) are high. The predation of eggs
and hatchlings mainly impacts
productivity. We conclude that
predation poses a threat to the NE
Indian DPS.
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Inadequacy of Existing Regulatory
Mechanisms
Turtles of the NE Indian DPS are
protected by several regulatory
mechanisms. For each, we review the
objectives of the regulation and to what
extent it adequately addresses the
targeted threat. Nearly all nations that
host nesting aggregations have
legislation to protect sea turtles.
In India, the leatherback turtle is
included on Schedule I, Part II of the
Wildlife (Protection) Act, 1972 (Entry
No. 11) updated by Wild Life
(Protection) Amendment Act, 2002 (No.
16 of 2003). India also bans the hunting
and trade of wild animals (India
National Report to CMS, 1991 and
1994). However, the indigenous people
of the Andaman and Nicobar Islands are
exempt from these laws. India has
regulations to require TEDs and
minimize fisheries interactions; and
much of the Andaman and Nicobar
Islands are protected as wildlife
sanctuaries, including two marine
national parks (Andrews et al. 2006).
In Indonesia, Order No. 301/1991 lists
leatherback turtles as a protected
species. Pursuant to the Act of 10
August 1990 on the Conservation of
Living Resources and Their Ecosystems,
it is prohibited to kill, capture, possess,
transport, trade in or export protected
animals whether alive or dead, or parts
of such animals. The taking,
destruction, trade or possession of the
eggs or nests of protected animals are
also prohibited (ECOLEX 2003). There
are no habitat protections and no
regulations to minimize fisheries
interactions or require TEDs in
Indonesia.
In Sabah, Malaysia, the leatherback
turtle is not listed as a totally protected
or partially protected species in the
Wildlife Conservation Enactment (No. 6
of 1997). In Sarawak, Malaysia,
leatherback turtles have been fully
protected since 1958. Under the
Wildlife Protection Ordinance 1998, all
marine turtles in Malaysia are protected
from hunting, killing, capture, sale,
import, export, possession of any
animal, recognizable part or derivative
or any nest, except in accordance with
the permission in writing of the
Controller of Wildlife for scientific or
educational purposes or for the
protection or conservation of a species
(Tisen and Bali 2002). The nesting
beach at Rantau Abang, Terengganu is
protected. However, the nesting
aggregation that once used this beach
has been extirpated. In 1994, the waters
surrounding 38 offshore islands of
Peninsular Malaysia and Labuan
became protected as marine parks. In
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addition, one national park in Sarawak,
three in Sabah, and one state park in
Terengganu protect coastal and marine
ecosystems (Malaysia National
Biodiversity Policy 1998). Additional
habitat protections include: The Turtle
Trust Ordinance 1957; Land Code 1958;
Turtle Protection Rules 1962; Fisheries
Prohibited Areas under section 61 of the
Fisheries Act 1985; and the Wildlife
Protection Ordinance 1998 (Tisen and
Bali 2002). The use of TEDs will be
required in Malaysia by 2020.
In Myanmar, the Burma Wildlife
Protection Act 1936 (Act No. VII of
1936) requires licenses to hunt, possess,
sell, or buy wild animals with closed
hunting seasons (FAOLEX 2003). The
Burma Wildlife Protection Rules of 1941
states that the import or export of any
reptile (including parts or products) into
or from Myanmar is prohibited.
In Pakistan, the leatherback turtle is
protected in Baluchistan, Azad Kashmir
and Sind (Baluchistan Wildlife
Protection Act 1974 No.19/1974; The
Azad Jammu and Kashmir Wildlife Act
1975 No.23/1975; The Sindh Wildlife
Protection Ordinance 1972 No.5/1972).
Possession, transport, and/or national
trade are prohibited or regulated
(ECOLEX 2003).
In Sri Lanka, the leatherback turtle is
protected under the Fauna and Flora
Protection Ordinance (Sri Lanka
National Report to CMS 1994), which
makes it an offense to kill, wound, harm
or take a turtle, or to use a noose, net,
trap, explosive or any other device for
those purposes, to keep in possession a
turtle (dead or alive) or any part of a
turtle, to sell or expose for sale a turtle
or part of a turtle, or to destroy or take
turtle eggs. The minister of Fisheries
and Aquatic Resources may also
prohibit or regulate the import and
export of turtles or their derivatives
(Parliament of the Democratic Socialist
Republic of Sri Lanka 1993). The
nesting beach in Yala Reserve is also
protected.
In Thailand, the Leatherback Turtle is
protected under the Animals Protection
Act B.D 2535 (The Zoological Park
Organization 2003).
In summary, numerous regulatory
mechanisms protect leatherback turtles,
their eggs, and nesting habitat
throughout the range of this DPS.
Although these regulatory mechanisms
provide some protection, many do not
adequately reduce the threat that they
were designed to address, generally as a
result of limited implementation or
enforcement. As a result, bycatch,
nesting habitat protection, and legal and
illegal harvest remain threats to the
DPS. We conclude that the inadequacy
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of the regulatory mechanisms is a threat
to the NE Indian DPS.
Fisheries Bycatch
Fisheries bycatch is a threat to the NE
Indian DPS. Capture in gillnet, trawl,
purse seine, and longline fisheries is a
significant cause of leatherback
mortality for this DPS (Wright and
Mohanty 2002; Hamann et al. 2006;
Project GloBAL 2007; Bourjea et al.
2008; Abdulqader 2010; Wallace et al.
2010).
Gillnet fisheries pose a major threat to
the DPS. A survey conducted at 16 main
fishing ports in Sri Lanka estimated that
431 leatherback turtles were caught in
gillnets between 1999 and 2000
(Kapurusinghe and Cooray 2002). In
Malaysia, Chan et al. (1988) reported an
average of 742 and 422 sea turtles, most
of which were leatherback turtles,
caught in drift gillnets and bottom
longlines, respectively. In Bangladesh,
gillnets, set bag nets, trawl nets, seine
nets, hook and line and other net types
of gear capture turtles (Hossain and Hoq
2010). Gillnet and purse seine fisheries
are common off the coasts of the
Andaman and Nicobar Islands, where
the largest nesting aggregations occur
(Shanker and Pilcher 2003; Chandi et al.
2012).
Trawl fisheries also pose a large threat
to the DPS. In India, TEDs are required
for trawl nets. However, fishers are
reluctant to use them (Murugan 2007).
Trawl fishing is also common in
Bangladesh, and the use of TEDs is not
required (Ahmed et al. 2006).
Longline fisheries occur in coastal
and pelagic waters. Huang and Liu
(2010) evaluated observer data from 77
trips (4,409 sets) on Taiwanese largescale longline fishing vessels in the
Indian Ocean. They identified 84
leatherback turtles captured from 2004
to 2008, with 48 mortalities (57 percent;
Huang and Liu 2010). Extrapolating to
the entire Taiwanese longline fishery in
the Indian Ocean, they estimated an
average bycatch of 173 leatherback
turtles between 2004 and 2007. This
number likely includes individuals from
both the SW and NE Indian DPSs (Louro
2006). In Vietnam, longline fisheries
continue to capture leatherback turtles.
However, a circle hook program has
been implemented to minimize the
impact (WWF 2013).
Purse seine fisheries have a much
lower impact than longline fisheries
(Angel et al. 2014); two leatherback
turtles were captured (alive) between
1995 and 2010 in the Indian Ocean
(Clermont et al. 2012). In the EEZ of all
Indian Ocean French Territories (mostly
from the Mozambique Channel), 40
leatherback turtles were captured in
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48385
unspecified fisheries from 1996 to 1999;
92 percent were released alive (Ciccione
2006).
In Thailand, one of the main causes
of decline in the turtle population is
bycatch in trawl, drift gillnet, and purse
seine fisheries. The rapid expansion of
fishing operations is largely responsible
for the increase in adult turtle mortality
due to bycatch (Settle 1995).
In Malaysia, the Fisheries Act of 1985
prohibited capture of sea turtles by any
type of fishery. However, this merely
reduced the reporting of interactions
(Yeo et al. 2011 in Dutton et al. 2011).
The 1991 Regulations prohibit fishing in
waters adjacent to Rantau Abang during
the leatherback nesting season (Chan
1993).
We conclude that juveniles and adults
are exposed to high fishing effort
throughout their foraging range and in
coastal waters near nesting beaches.
Mortality rates are likely high,
especially in areas where turtle meat is
consumed. Mortality reduces
abundance, by removing individuals
from the population. It also reduces
productivity, when nesting females are
incidentally captured and killed. We
conclude that fisheries bycatch is a
major threat to the NE Indian DPS.
Pollution
Pollution includes contaminants,
marine debris, and ghost fishing gear.
Ghost fishing gear can drift in the ocean
and fish unattended for decades and kill
numerous individuals (Wilcox et al.
2013). The main sources of ghost fishing
gear are gillnet, purse seine, and trawl
fisheries (Stelfox et al. 2016). In one
collection event, volunteers collected
over 600 nets, ropes, and buoys from
India, Maldives, Oman, Pakistan, Sri
Lanka, and Thailand (Stelfox et al.
2016). Though educational programs
created in 2014 focus on reusing and
recycling fishing gear, the threat
continues throughout the range of the
DPS. Ghost nets in the Maldives
primarily drift from fisheries in the Bay
of Bengal (e.g., Sri Lanka and India;
Stelfox et al. 2016). Around the
Andaman and Nicobar Islands and Sri
Lanka, plastics and other garbage are
washed from polluted beaches and
inland waters to the sea, where they can
kill or harm sea turtles through
ingestion or entanglement
(Kapurusinghe 2006; Das et al. 2016).
Pollution has been identified as a main
threat to sea turtles in Iran (Mobaraki
2007) and Pakistan (Firdous 2001).
However, no specific information about
the type of pollution was provided. In
Gujarat, India, increased port and
shipping traffic have resulted in oil
spills and the release of other
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pollutants, such as fertilizers and
cement (Sunderraj et al. 2006). Heavy
metals and E. coli were found at
relatively high levels in the waters of
Malaysia (including Terengganu) and in
the pancreases and livers of leatherback
turtles (Caurant et al. 1999; Ngah et al.
2012). It is not known how these
pollutants affect leatherback physiology
(Jakimska et al. 2011).
As with all leatherback turtles,
entanglement in and ingestion of marine
debris and plastics are threats that likely
kill several individuals a year. However,
data specific to this DPS were not
available. We conclude that pollution is
a threat to the NE Indian DPS, albeit
with effects that are unquantifiable on
the basis of the best available
information.
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Climate Change
Climate change is a threat to the NE
Indian DPS. A significant rise in sea
level would further reduce nesting
habitat, which is already affected by
erosion. The DPS is also likely to be
affected by increases in sand
temperatures (Hawkes et al. 2009;
Poloczanska et al. 2009). Sand
temperatures prevailing during the
middle third of the incubation period
determine the sex of hatchling sea
turtles (Mrosovsky and Yntema 1980).
Incubation temperatures near the upper
end of the tolerable range produce only
female hatchlings, while incubation
temperatures near the lower end of the
tolerable range produce only males. As
temperatures increase, incubation
temperatures may exceed the thermal
tolerance for embryonic development,
thus increasing embryo and hatchling
mortality.
In addition, the frequency and
intensity of severe storm events and
cyclones in the Bay of Bengal are
predicted to increase with climate
change (Balaguru et al. 2014).
Climate change is likely to modify
nesting conditions for the entire DPS.
Impacts likely range from small changes
in nesting metrics to large losses of
productivity. As the DPS is already
experiencing nesting habitat loss due to
coastal erosion, we conclude that
climate change is a threat to the NE
Indian DPS.
Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply to
the NE Indian DPS (for a description of
each effort, please see the section on
conservation efforts for the overall
species): Association of Southeast Asian
Nations Ministers on Agriculture and
Forestry, Andaman and Nicobar Island
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Environmental Team, The Centre for
Herpetology/Madras Crocodile Bank
Trust, Convention on the Conservation
of Migratory Species of Wild Animals,
Convention on Biological Diversity,
Convention on International Trade in
Endangered Species of Wild Fauna and
Flora, Convention Concerning the
Protection of the World Cultural and
Natural Heritage (World Heritage
Convention), FAO Technical
Consultation on Sea Turtle-Fishery
Interactions, The Indian Ocean Tuna
Commission, Indian Ocean—South-East
Asian Marine Turtle Memorandum of
Understanding, MARPOL, IUCN,
Memorandum of Agreement between
the Government of the Republic of the
Philippines and the Government of
Malaysia on the Establishment of the
Turtle Island Heritage Protected Area,
Memorandum of Understanding on
Association of South East Asian Nations
Sea Turtle Conservation and Protection,
The Memorandum of Understanding of
a Tri-National Partnership between the
Government of the Republic of
Indonesia, the Independent State of
Papua New Guinea and the Government
of Solomon Islands, National Sea Turtle
Conservation Project in India, Ramsar
Convention on Wetlands, UNCLOS, and
UN Resolution 44/225 on Large-Scale
Pelagic Driftnet Fishing. Although
numerous conservation efforts apply to
the turtles of this DPS, they do not
adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the NE Indian DPS is at high risk of
extinction. The once large nesting
aggregation in Malaysia is now
functionally extirpated. The total index
of nesting female abundance is 109
females at all monitored beaches. This
estimate is likely low because several
nesting sites were not included in the
calculation due to lack of consistent,
standardized monitoring over multiple
and entire nesting seasons. Still, the low
nesting female abundance places this
DPS at risk of stochastic or catastrophic
events that increase its extinction risk.
The DPS once exhibited much greater
nesting female abundance, which has
dramatically declined in recent decades.
It currently exhibits a slightly declining
nest trend at monitored nesting beaches
in India. The DPS exhibits average
productivity metrics, such as body size,
clutch size and frequency. Though it
exhibits some spatial distribution and
diversity, with multiple foraging sites
and relatively high genetic diversity at
the sampled locations, nesting only
occurs on islands. Based on these
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factors, we find the DPS to be at risk of
extinction as a result of past threats.
Current threats further contribute to
the risk of extinction of this DPS. Major
threats to the DPS include fisheries
bycatch and the harvest of turtles and
eggs. There are not many nests to
exploit, but evidence suggests that if
such nests are found by humans, the
eggs are at risk of being harvested. Egg
harvest led to the extirpation of the
largest nesting aggregation (i.e.,
Malaysia), and current overexploitation
occurs in Thailand, Vietnam, and Sri
Lanka. The poaching of turtles is also a
threat in Myanmar. Fisheries bycatch is
a major threat, with turtles being
captured in trawl and gillnet fisheries in
Malaysia, India, Thailand, Sri Lanka,
Bangladesh, and Indonesia. Erosion on
the Andaman and Nicobar Islands, as a
result of tsunami damage, has
significantly reduced available nesting
habitat. Additional habitat
modifications include coastal
development and artificial lighting, as a
result of increases in tourism. Pollution
and climate change are threats that
likely affect the DPS by reducing
abundance and productivity, though the
best available data do not allow for
quantification of those effects. Though
many regulatory mechanisms are in
place, they do not reduce the impact of
threats to levels that ensure the
continued existence of the DPS.
We conclude, consistent with the
team’s findings, that the NE Indian DPS
is currently in danger of extinction. Its
low nesting female abundance makes
the DPS highly vulnerable to threats.
Dramatic declines in over the past
several decades contribute to our
concern over the continued persistence
of the DPS. Past egg and turtle harvest
initially reduced the nesting female
abundance of this DPS and likely
confined its nesting habitat to a few
island beaches, with little diversity and
reduced spatial distribution. The
present, ongoing threats include:
overutilization (i.e., turtle and egg
harvest); fisheries bycatch; loss of
habitat; and predation. Overutilization
and fisheries bycatch reduces
abundance and productivity (i.e.,
imminent and substantial demographic
risks) by removing mature and
immature individuals from the
population at rates exceeding
replacement. The loss of nesting habitat
and predation (of eggs) reduces
productivity and the DPS’s ability to
recover to its previous abundance.
Though numerous conservation efforts
apply to this DPS, they do not
adequately reduce the risk of extinction.
We conclude that the NE Indian DPS is
in danger of extinction throughout its
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range and therefore meets the definition
of an endangered species. The
threatened species definition does not
apply because the DPS is at risk of
extinction currently (i.e., at present),
rather than on a trajectory to become so
within the foreseeable future.
West Pacific DPS
The Team defined the West Pacific
DPS as leatherback turtles originating
from the West Pacific Ocean, south of
71° N, north of 47° S, east of 120° E, and
west of 117.124° W. The northern and
southern boundaries reflect the highest
latitude occurrences of leatherback
turtles in each hemisphere (Goff and
Lien 1988; Carriol and Vader 2002;
McMahon and Hayes 2006; Shillinger et
al. 2008; Benson et al. 2011; Eckert et
al. 2012). We placed the western
boundary at 120° E to approximate the
Wallace and Huxley lines, which are
established biogeographic barriers to
gene flow between Indian and Pacific
Ocean populations of numerous species.
While the genetic differences between
the Northeast Indian and West Pacific
DPSs demonstrate discreteness, genetic
sampling is unavailable from areas
where the nesting ranges of the DPSs
likely meet, preventing us from defining
the boundary more specifically. We
placed the eastern boundary at the
border between the United States and
Mexico to reflect the DPS’s wide
foraging range throughout the Pacific
Ocean. We chose this border because
the West Pacific DPS crosses the ocean
to forage in the eastern Pacific Ocean,
including in waters of the United States,
whereas the East Pacific DPS forages
primarily off the coasts of Central and
South America. The two DPSs overlap
in foraging habitats off waters of Chile
and Peru (Donoso and Dutton 2010).
The range of the DPS (i.e., all areas of
occurrence) extends throughout the
Pacific Ocean with specific coastal and
pelagic areas in the Indo-Pacific basin
providing important foraging and
migratory habitats. Documented nesting
occurs on beaches of the following
nations: Indonesia, Papua New Guinea,
Solomon Islands, and Vanuatu.
Leatherback turtles of the West Pacific
DPS migrate through the EEZs of at least
32 nations including in the U.S. EEZs of
California and Hawaii, spending
between 45 and 78 percent of the year
on the high seas (Harrison et al. 2018).
Of the 32 nations, the West Pacific DPS
migrates through at least 18 nations or
territories of the western and
southwestern Pacific Ocean: Indonesia,
Papua New Guinea, Solomon Islands,
Philippines, Malaysia, Vietnam, Japan,
Palau, Micronesia, Marshall Islands,
Northern Mariana Islands and Guam,
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Fiji, Vanuatu, Australia, New Caledonia,
New Zealand, Line Islands, and Kiribati
(Harrison et al. 2018). Foraging occurs
in seven ecoregions: South China/Sulu
and Sulawesi Seas, Indonesian Seas,
East Australian Current Extension,
Tasman Front, Kuroshio Extension of
the Central North Pacific, equatorial
Eastern Pacific, and California Current
Extension (Benson et al. 2011).
Individuals demonstrate fidelity to these
foraging areas, likely as a result of their
post-hatchling dispersal patterns and
nesting season (Benson et al. 2011;
Gaspar et al. 2012; Gaspar and Lalire
2017; Harrison et al. 2018).
Leatherback turtles of the West Pacific
DPS nest in tropical and subtropical
latitudes primarily in Indonesia, Papua
New Guinea, and Solomon Islands, and
a lesser extent in Vanuatu (Dutton et al.
2007; Benson et al. 2007a; Benson et al.
2007b; Benson et al. 2011). The majority
of nesting occurs along the north coast
of the Bird’s Head Peninsula, Papua
Barat, Indonesia at Jamursba-Medi and
Wermon Beaches (Dutton et al. 2007). A
recent discovery of a previously
undocumented nesting area on Buru
Island, Maluku Province, Indonesia
(WWF 2018) suggests that additional
undocumented nesting habitats may
exist on other remote or infrequently
surveyed islands of the western Pacific
Ocean. This DPS nests year round, and
exhibits a bimodal nesting strategy
whereby a proportion of females nest
during November through February (i.e.,
‘‘winter’’ nesting females) and other
females nest May through September
(i.e., ‘‘summer’’ nesting females; Benson
et al. 2007a; Benson et al. 2007b; Dutton
et al. 2007; Tapilatu and Tiwari 2007;
Benson et al. 2011).
Nesting beach habitats throughout the
West Pacific are generally dynamic,
high profile beaches associated with
deep water approaches and strong
waves. Beaches can be quite narrow as
in parts of the Solomon Islands or Papua
New Guinea, or broad as in the case of
Jamursba-Medi, Indonesia during the
summer months. Nesting females appear
to prefer coarse-grained sand free of
rocks, coral, or other abrasive substrates
(reviewed by Eckert et al. 2012).
While West Pacific leatherback turtles
do not have distinct ‘‘migratory
corridors,’’ several areas are considered
‘‘areas of passage’’ used by turtles
traveling between nesting and foraging
locations, and there is clear separation
of migratory and foraging destinations
based on nesting season (Benson et al.
2007a, b; Benson et al. 2011; Harrison et
al. 2018). Post-nesting, winter nesting
females from Papua New Guinea,
Indonesia, and Solomon Islands migrate
through the Halmahera, Bismarck,
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Solomon, and Coral Seas, towards
Southern Hemisphere temperate and
tropical foraging areas in the Tasman
Sea, East Australian Current, and
western South Pacific Ocean (Benson et
al. 2011; Harrison et al. 2018; Jino et al.
2018). Genetic analyses of leatherback
turtles caught in fisheries off Peru and
Chile indicates that approximately 15
percent of sampled individuals originate
from the West Pacific DPS, likely winter
nesting females that have migrated
across the Southern Hemisphere to the
productive waters off South America
(Donoso and Dutton 2010; NMFS
unpublished data 2018). It is unclear
what proportion of the West Pacific DPS
might utilize this area and how
important it might be to this DPS.
Leatherback turtles migrate through
and forage in the waters of the
Philippines (Benson et al. 2007a, 2011;
MRF 2010, 2014). In 2005, Salinas et al.
(2009) found a female in San Fernando
(close to El Nido) that had been
previously tagged at Jamursba-Medi in
July 2003. The Marine Research
Foundation (MRF) utilized aerial
transects to assess leatherback foraging
area use in Palawan waters and off the
coast of Borneo (MRF 2010, 2014). They
found leatherback turtles (n = 28 in
2010 and 2013/2014) foraging in
nearshore waters around the NE and SE
coasts of Palawan, potentially linked to
large jellyfish aggregations from
February to May, and overlapping with
high density fishing activity in Taytay
Bay, off NE Palawan (MRF 2010, 2014).
Additionally, numerous leatherback
turtle marine sightings, strandings, and
fishery bycatch (typically entangled in
gillnet gear) exist for locations
throughout the Philippines including
Marine Wildlife Watch of the local
NGO, Marine Wildlife Watch of the
Philippines, from 2010 to 2018
(Bagarinao 2011; Cruz 2006; MRF 2010;
MWWP unpublished data 2018).
Abundance
The total index of nesting female
abundance of the West Pacific DPS is
1,277 females. We based this total index
on two nesting aggregations in
Jamursba-Medi and Wermon, Indonesia
(Tapilatu et al. 2013; Tiwari et al. in
prep). Our total index does not include
18 unquantified nesting aggregations in
Indonesia, Papua New Guinea, Solomon
Islands, and Vanuatu. To calculate the
index of nesting female abundance (723
females) for Jamursba-Medi (i.e., a 18
km stretch of beach that has been
monitored since 1981), we divided the
total number of nests between the 2015/
2016 and 2017/2018 nesting seasons
(i.e., a 3-year remigration interval) by
the clutch frequency (5.5 clutches per
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season; Tapilatu et al. 2013). We
performed a similar analysis for data
from Wermon (index = 554 females), a
6 km beach that has been monitored
since 2002.
Based on the Tapilatu et al. (2013)
study, the IUCN Red List assessment
estimated the total number of mature
individuals (including females and
males) utilizing Jamursba-Medi and
Wermon beaches to be 1,438 leatherback
turtles (Tiwari et al. 2013). The IUCN
estimate includes males and thus is
higher than ours. Curtis et al. (2015)
provided a minimum annual nesting
female estimate of 318 females (or 954
total nesting female abundance over a 3year remigration interval). Dutton et al.
(2007) estimated that 1,113 females may
have nested annually, or conservatively
2,700 total nesting females, in the entire
western Pacific population. At that time,
they estimated 75 percent of the
population originated from Bird’s Head
Peninsula (or approximately 2,025
females; Dutton et al. 2007). Our total
index is within the range of published
estimates of abundance for this DPS,
taking into account differences in
survey methods over time, and is based
on the best available data for the DPS at
this time.
Within the nesting range of this DPS,
nest monitoring activities have occurred
relatively recently, with standardized
methods in Papua Barat first
implemented in 2002 (Hitipeuw et al.
2007; Tapilatu et al. 2013). Outside the
Bird’s Head Peninsula, monitoring has
been sporadic, opportunistic, and
spatially limited because the region is
vast, remote, and logistically
challenging to access. Often nesting
beaches are located far from towns or
cities, where there are no roads to, or
electricity in, adjacent villages. Cultural
and socio-economic dynamics confound
monitoring programs, which are
dependent upon fiscal sponsorship,
incentives, community buy-in, and the
degree of familiarity of local
communities with concepts of
sustainability or conservation (Kinch
2006; Gjersten and Pakiding 2012).
While Jamursba-Medi and Wermon
beaches have been monitored fairly
consistently over time, less is known
about the status and trends of nesting
beaches in Papua New Guinea, Solomon
Islands, and Vanuatu. Records are
further confounded by changes in place
names and jurisdictional boundaries
over recent decades (e.g. the Indonesian
province formerly known as Irian Jaya is
currently two provinces of Papua and
Papua Barat). Village names or location
descriptions have also changed over
time, and geographic coordinates were
not recorded historically. Therefore, all
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estimates of abundance in this DPS
carry substantial uncertainty.
In Indonesia, aerial surveys provided
the first indication of leatherback
nesting in Papua (i.e., Irian Jaya; Salm
1982). At that time, Salm (1982) did not
provide location details out of concern
that public disclosure prior to
protection would be detrimental.
Follow-up studies during the 1980s and
1990s indicated that a large nesting
population was located along the coastal
beaches of northern Papua or Papua
Barat, Bird’s Head Peninsula (Bhaskar
1985). Systematic monitoring of
leatherback turtles began during the
early 1990s, primarily in the form of
annual nest counts (Hitipeuw et al.
2007). On the Bird’s Head Peninsula of
Papua Barat, nesting occurs mainly at
Jamursba-Medi and Wermon, where a
total of 1,371 nesting females were
tagged between 2002 and 2011 (Tapilatu
et al. 2013). The primary nesting season
at Jamursba-Medi occurs during the
summer (May to September), while
nesting occurs year round at Wermon
with a small peak in July and primary
nesting activity during the winter,
between November and February
(Hitipeuw et al. 2007). Historically,
approximately 60 percent of nesting
activity occurs at Jamursba-Medi with
40 percent of activity at Wermon
(Tapilatu et al. 2013). While a few
females have been documented nesting
at both beaches during a nesting season
(Tapilatu et al. 2013), the vast majority
of females do not appear to utilize both
Jamursba-Medi and Wermon Beaches
during a single nesting season (Tapilatu
and Tiwari 2007; Tapilatu et al. 2013;
Lontoh 2014). Based on nest counts and
clutch frequency per season (mean = 5.5
+/¥ 1.6 nests per female),
approximately 464 to 612 females
nested at Jamursba-Medi and Wermon
in 2011 (Tapilatu et al. 2013).
Additional low-level nesting activity in
Indonesia occurs in the Manokawari
region of the Bird’s Head Peninsula to
the east of the Jamursba-Medi and
Wermon Beaches (Suganuma et al.
2012). Between 2008 and 2011, 84 to
135 nests were recorded, or a mean of
about 117 nests annually (Suganuma et
al. 2012). However, survey effort was
limited and not consistent across years
and may underestimate total nesting
activity. Further it is unknown whether
interchange occurs between turtles
nesting in the Manokawari region and
those of the Bird’s Head Peninsula
index beaches. In 2016, nesting activity
was identified in Central Maluku at
Buru Island, west of Bird’s Head
Peninsula. In 2017, a monitoring
program to quantify nesting activity was
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initiated on three north coast beaches of
Buru Island (totaling 10 km) which
documented 203 nests, and preliminary
data indicates that there might be two
nesting peaks: May through July and
November through February (WWF
2018). Nesting activity in other areas of
Indonesia are known or suspected, but
unquantified (Dutton et al. 2007;
Tapilatu 2017).
In Papua New Guinea, the majority of
known nesting activity occurs during
the winter months (November to
February) along the Huon Coast on the
northeastern coast of the Morobe
Province, where 576 females were
tagged between 1999 and 2013 (Pilcher
2006, 2008, 2009, 2010, 2011, 2012,
2013; Pilcher and Chaloupka 2013).
Aerial surveys along the Huon Coast in
January and December between 2004
and 2006 documented 276 nests, with
an estimate of 500 nests per season
(Benson et al. 2007b; Dutton et al. 2007).
During the Huon Coast Leatherback
Turtle Project, which took place
between 2005 and 2012, an average of
258 nests were laid per season (range:
193 to 527) at seven beaches which
comprised approximately 35 km of
nesting habitat along the Huon Coast
(Pilcher 2013; WPRFMC 2015). One
challenge in estimating nesting activity
in Papua New Guinea is that leatherback
site fidelity appears to be variable, with
some satellite tagged animals seen
visiting a number of areas during one
nesting season (Benson et al. 2007b).
For example, a number of Huon Coast
nesting females visited other nearby
beaches and east-facing beaches of the
Huon Peninsula, including Bougainville
and Woodlark Islands during a single
nesting season (Benson et al. 2007b).
Therefore, for assessment purposes, we
consider the Huon Coast to be one
nesting beach complex.
Additional nesting activity occurs in
other areas of Papua New Guinea, such
as along the north coast of the Madang
Province and on several islands
including Manus, Long, New Britain,
Bougainville, New Ireland, and
Normanby (Prichard 1982; Spring 1982;
Benson et al. 2007b; Dutton et al. 2007).
In these areas nesting activity has not
been quantified via standardized or
consistent methods, but information has
been obtained via community surveys,
aerial surveys, or rapid assessments.
Nesting occurs primarily in the winter
months, although low-level year-round
nesting may also occur (Spring 1982;
Dutton et al. 2007). Approximately 50
nests may be laid annually along the
north coast of the Madang Province
(Benson et al. 2007b; TIRN 2017). The
Islands of New Britain and Bougainville
may host approximately 140 to 160
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nests per year, respectively (Benson et
al. 2007b; Dutton et al. 2007; Kinch et
al. 2009). On Bougainville Island, aerial
surveys conducted during the 2005 and
2007 nesting seasons documented a
mean of 68 nests (range: 41 to 107 nests)
or an extrapolated estimate of 160 to 415
nests per year (Dutton et al. 2007;
Benson et al. 2007b). In 2009, a one
week full-island ground survey
(conducted by boat and foot) recorded
46 leatherback nests (Kinch et al. 2009).
In the Solomon Islands nesting
activity is distributed throughout the
country with the majority of nesting
activity at Sasakolo and Litogarhira
beaches on Isabel Island, and on
Rendova and Tetepare Islands in the
Western Province (Pita 2005; Dutton et
al. 2007; Benson et al. 2018a). The
nesting season occurs primarily during
winter (November through February),
although some year-round nesting has
been documented (Pilcher 2010b;
Williams et al. 2014; Jino et al. 2018;
TNC-Solomon Islands 2018
unpublished). Leatherback turtle
monitoring was begun by the Solomon
Island Department of Fisheries in 1989
(Pita 2005). Between 1999 and 2006, an
estimated 640 to 700 nests were laid
annually in the Solomon Islands,
representing approximately eight
percent of the total western Pacific
leatherback nesting at that time (Dutton
et al. 2007). At Sasokolo Beach, Isabel
Island, during a 54 day monitoring
period between November 28, 2000 and
January 21, 2001, 132 nests were
documented with an additional 35 nests
present when monitoring began
(Ramohia et al. 2001). Between
December 27, 2006 and January 2, 2007,
aerial surveys provided seasonal
estimates of 207 nests laid on Isabel
Island, and an additional 312 nests on
other islands (Benson et al. 2018a). A
January 2011 site visit resulted in 315
nests identified at Sasakolo and
Litogahira (Tiwari 2011 unpublished).
Recently, nesting activity has also been
documented at the southeastern side of
Isabel, where approximately 52 females
may nest annually (TNC-Solomons 2018
unpublished). Since 2002, the Tetepare
Descendants’ Association (TDA) has
monitored nesting activity
opportunistically in the Solomon
Islands, where approximately 30 to 50
leatherback nests are laid seasonally on
two beaches (Goby et al. 2010). Between
July 1, 2012 and April 30, 2013, TDA
undertook 257 beach surveys and found
44 leatherback nests (TDA 2013). While
monitoring efforts may be ongoing, data
management and analysis remains a key
challenge for these isolated
communities (TDA 2013; Pilcher
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2010b). At Rendova Island during the
2003/2004 winter nesting season, 235
leatherback turtle nests were recorded,
and during the 2009/2010 season, 79
nests were laid (Pilcher 2010b; Goby et
al. 2010). Likely the most
comprehensive surveys occurred from
September 1, 2012 to April 30, 2013 (91
patrols, 3 days per week), which
documented a total of 74 nests (TDA
2013). During the 2017/2018 winter
nesting season, 29 nests were
documented (Solomon Islands
Community Conservation Partnership
2018 unpublished data). The
community on Vangunu Island
documented a total of 23 nests and 11
females between June 2011 and July
2014 (Jino et al. 2018). Nesting occurred
during two distinct seasons from May to
July and from November to January, and
of the females tagged, one nested
successfully six times and another
nested five times (Jino et al. 2018). The
other nine turtles were only observed
nesting once or twice, and it is likely
that either some nesting events were not
recorded or the females nested on
surrounding unmonitored beaches (Jino
et al. 2018). On Malaita Island at
Waisurione beach, nesting activity
occurs during the summer (June to
August), but only a few females were
determined to use the area, with five
and seven nests documented in 2014
and 2015, respectively (Williams et al.
2014).
Nesting occurs in low numbers at
other islands in the western Pacific
Ocean. In Vanuatu, 30 to 40 nests are
laid annually on Epi and Ambrym
Islands (Dutton et al. 2007; Petro et al.
2007; WSB 2011), although fewer nests
(n = 15) were documented during the
2014/2015 nesting season (WSB 2016).
Leatherback turtles have been reported
in Fiji (Rupeni et al. 2002; NMFS and
USFWS 2013; Jino et al. 2018), but these
accounts involved foraging or in-water
capture of animals, and it is unclear if
historic reports included nesting
activity (Guinea 1993; Benson et al.
2013). Historical nesting records also
exist for the eastern coast of
Queensland, in New South Wales, and
in the Northern Territories from
December to February (Dobbs 2002;
Limpus 2009). However, current
information was not available at the
time of the study, and no nests have
been observed since 1995 despite
regular monitoring (Flint et al. 2012).
Since the 1980s, there have also been
reports of leatherback turtles nesting in
the Philippines (Cruz 2006; MRF 2010).
Of recent reports, two documented cases
have been confirmed by sea turtle
experts (i.e., staff of the Marine Wildlife
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Watch of the Philippines). On July 15,
2013, at Barangay Yawah, Legazpi City,
Albay, NAVFORSOL (the Philippines
Naval facility) personnel observed a
leatherback nesting, but the eggs failed
to hatch. On August 6, 2013 at Camp
Picardo beach, Barangay, Eastern Samar,
a nesting event was aborted due to
disturbance on the beach, but according
to the social media report (i.e., a
Facebook post), the female was tagged
and led back to sea (MWWP
unpublished 2018). Given the low-site
fidelity of the turtles in this DPS
(Benson et al. 2007b), it is not surprising
that leatherbacks might distribute nests
among various areas throughout the
region.
The total index of nesting female
abundance of the West Pacific DPS (i.e.,
1,277 females) places it at risk for
environmental variation, genetic
complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). These
processes, working alone or in concert,
place small populations at a greater
extinction risk than large populations,
which are better able to absorb impacts
to habitat or losses in individuals. Due
to its small size, the DPS has restricted
capacity to buffer such losses. Given the
intrinsic problems of small population
size, we conclude that the nesting
female abundance is a major factor in
the extinction risk of this DPS.
Productivity
The West Pacific DPS exhibits a
declining nesting trend. We conducted
trend analyses for the two index beaches
in Indonesia, which were the only two
beaches with 9 or more recent years of
standardized data, with the most recent
data collection in 2014 or more recently
(the standards for conducting a trend
analysis in this report). The median
trend in annual nest counts estimated
for Jamursba-Medi (data collected from
2001 to 2017) was ¥5.7 percent
annually (sd = 5.4 percent; 95 percent
CI = ¥16.2 to 5.3 percent; f = 0.867;
mean annual nests = 2,063). While data
are available for the period starting in
1999, the best available information
indicates that beach monitoring and
nest protection practices improved in
2001; therefore, we used the time series
starting in 2001. For Wermon (data
collected from 2006 to 2017, excluding
2002–2005 and 2013–2015 due to low or
insufficient effort), the median trend
was ¥2.3 percent annually (sd = 8.4
percent; 95 percent CI = ¥19.8 to 14.9
percent; f = 0.643; mean annual nests =
1,010). As Jamursba-Medi and Wermon
currently represent approximately 75
percent of nesting for this DPS, we
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consider these declining trends to be
representative of the entire DPS.
Our trend data for Indonesia yield
similar results to other published
findings. The IUCN Red List assessment
found a decreasing trend of ¥7 percent
annually (Tiwari et al. 2013). Tapilatu et
al. (2013) identified a ¥5.5 percent
annual rate of decline at Jamursba-Medi
between 1984 and 2011 and a ¥11.6
percent annual rate of decline at
Wermon between 2002 and 2011.
Between 1986 and 2010, Benson et al.
(2013) highlighted drastic declines in
the annual number of nests at JamursbaMedi and Wermon. Additionally, a 27year aerial survey study indicates a
decline in the number of leatherback
turtles foraging off central California
(Benson et al. 2018b). From 1995 to
2003, an estimated 12 to 379 individuals
(mean = 178) foraged in this area
(Benson et al. 2007), while from 2004 to
2017, an estimated 23 to 112 individuals
foraged in this area, representing a
decline of 5.6 percent annually (Benson
et al. 2018b).
At Jamursba-Medi, nesting data have
been collected for some years since
1981. However, no data were collected
during many years in the mid-1980s and
late 1990s (Tapilatu et al. 2013). There
is considerable uncertainty in the early
estimates, with over 4,000 nests
estimated in 1981, 14,522 nests in 1984,
and a dramatic drop to 3,261 nests in
1985 (Tapilatu et al. 2013). It is unclear
if there was sampling inconsistency
between years or if there was an actual
decline in nesting activity. However, if
analyses are based on the 1984 data,
during which the greatest number of
nests was recorded, there was a 78.3
percent decline over the past 27 years
(1984 to 2011), or 5.5 percent annual
rate of decline (Tapilatu et al. 2013).
Alternatively, if analysis is based on
2005 to 2011 when the Tapilatu et al.
(2013) study ensued, nesting activity
declined 29 percent from 2,626 nests (in
2005) to 1,596 nests (in 2011; Tapilatu
et al. 2013). Since the Tapilatu et al.
(2013) study, University of Papua
scientists have continued to engage with
local communities to monitor nesting
activity. The overall nesting trend has
continued to decline by 5.6 percent per
year between 2003 and 2017. However,
there appears to be an increase in
nesting since 2013 (Tiwari et al. in
prep).
The first comprehensive surveys at
Wermon beach in 2002 found almost as
many nests laid on Wermon as on
Jamursba-Medi (Hitipeuw et al. 2007).
At that time, it was hypothesized that
the decline at Jamursba-Medi may have
been offset by an increase at Wermon
(Hitipeuw et al. 2007). However,
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Tapilatu et al. (2013) found a significant
decline in nesting at Wermon from
2,994 nests in 2002 to 1,096 nests in
2011 (62.8 percent total or 11.6 percent
annual rate of decline). Unfortunately,
no monitoring activities occurred at
Wermon between 2013 and 2015 due to
community discord, which prevented
beach access. Between 2006 and 2017,
nesting has continued to decline at
approximately 2.3 percent (Tiwari et al.
in prep). However, there may have been
a slight increase in recent nesting,
similar to Jamursba-Medi (Tiwari et al.
in prep).
Local residents stated that leatherback
turtles were the dominant sea turtle
species nesting in Maokawari prior to
the 1980s, but that the population has
declined significantly since the 1990s
due to village development and
exploitation of turtles and eggs (Tapilatu
et al. 2017).
Data collection in Papua New Guinea
spanned 8 years and ended prior to
2014. Because these data did not meet
our criteria for ‘‘recent,’’ we did not
perform a trend analysis, but included
a bar graph in the Status Review Report.
In Papua New Guinea, nesting activity
along the Huon Coast was relatively
stable between 2005 and 2013, with 193
to 527 nests per year (mean annual nests
= 258) and with most nesting activity
occurring at two primary areas, Busama
and Kamiali (Pilcher 2013; Benson et al.
2015; WPRFMC 2015). Given the
exchange of females and evidence of
multiple beach use among females in
Papua New Guinea (Benson et al.
2007b), we consider the Huon Coast to
be one nesting area and not individual
nesting beaches. Though there have
been several independent studies of
abundance over time, we determined
that these data are inadequate to
incorporate into a trend analysis
because these data do not meet our
criteria (i.e., nest count data consistently
collected in a standardized approach for
at least 9 years). For historical
perspective, leatherback turtle nesting
along the Huon Coast was first
identified south of the city of Lae near
the Buang River, at an area likely
between Labu Tale and Busama villages
(i.e., Maus Buang or Buang-Buassi;
Bedding and Lockhart 1989; Quinn and
Kojis 1985; Hirth et al. 1993). Estimates
of leatherback turtle nesting at Maus
Buang during the 1980s ranged from
five to 10 turtles per night from
November to January (Quinn and Kojis
1985) or 300 nests laid annually
(Bedding and Lockhart 1989). Quinn
and Kojis (1985) estimated that 300 to
500 females may nest annually in Papua
New Guinea, although it is unclear if
estimates were for the Maus Buang area
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specifically or the Huon Coast at large.
Hirth et al. (1993) undertook the most
standardized survey at that time and
recorded 76 nests and 34 females
nesting at ‘‘Piguwa’’ (i.e., Maus Buang)
on 725 meters of beach during a 15-day
period in December 1989. During the
Huon Coast leatherback turtle nesting
beach program, an average of 35 and 114
nests were laid annually during the 4month nesting season in this similar
area at Labu Tale and Busama beaches,
respectively (Pilcher 2013; WPRFMC
2015). Kamiali Beach lies approximately
30 km south of the city of Lae. In 1996,
the Kamiali Wildlife Management Area
was declared a protected area for
leatherback turtles, and the harvest of
nests was prohibited along 2 km of
beach. In 1999, village rangers began
opportunistic tagging of nesting females
at Kamiali. A community-based nesting
beach monitoring program was
established in 2003, which soon grew
into the Huon Coast Leatherback Turtle
Conservation Program (Benson et al.
2007b; Pilcher and Chaloupka 2013;
Kinch 2006). By 2005, monitoring
activities expanded from Kamiali Beach
(approximately 7 km) to seven beaches
encompassing approximately 35 km of
nesting beaches, which included an
agreement by participating villages to no
longer harvest eggs (Kinch 2006; Pilcher
2013). Of these seven beaches, Kamiali
was the nesting beach with the longest
running, most consistent monitoring
within the Huon Coast nesting beach
complex. At Kamiali, 194 females were
tagged between 1999 and 2012, and an
average of 77 nests laid per winter
nesting season between 2005/2006 and
2012/2013 (Pilcher 2010, 2011, 2012,
2013; Pilcher and Chaloupka 2013).
While we are unable to interpret an
overall trend from these studies,
anecdotal reports from villagers and
historic information indicates that
leatherback nesting activity was
significantly greater in past decades
(Benson et al. 2007b, 2015; Hirth et al.
1993; Kinch 2006; Bellagio Sea Turtle
Conservation Initiative 2008).
In the Solomon Islands, it is not
possible to estimate nesting trends due
to non-standardized methods and
opportunistic monitoring efforts over
time. Available datasets cannot be
compared due to differences in
methodology and do not meet our
criteria (i.e., nest count data consistently
collected in a standardized approach for
at least 9 years). Historically, nesting
was reported at more than 15 beaches in
the Solomon Islands, which may have
totaled several hundred nests per season
(McKeown 1977; Vaughan 1981).
Currently, nesting activity occurs
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primarily in eight locations (Pita 2005;
Dutton et al. 2007; Benson et al. 2018a;
Jino et al. 2018). However, due to the
remoteness of these areas and lack of
systematic surveys, and likely
additional undocumented nesting
beaches, additional low numbers of
nesting leatherback turtles are likely to
exist in Solomon Islands. For example,
nesting activity was recently identified
on Vanugnu Island, where 23 nests were
recorded and 11 females nested between
2011 and 2014 (Jino et al. 2018).
Additionally, it is unknown to what
extent females use multiple beaches
throughout the Solomon Islands, or
those in Papua New Guinea, and what
proportion of females nest in the
summer versus winter (Benson et al.
2007b; Jino et al. 2018; TNC-Solomons
2018 unpublished). While we are unable
to interpret an overall trend, local
villagers indicate that leatherback
nesting was greater in past decades
(Bellagio Sea Turtle Conservation
Initiative 2008; Benson et al. 2007b;
Benson et al. 2015).
In Vanuatu, anecdotal information
suggests that nesting has declined over
time (Petro et al. 2007). During the
2010/2011 winter nesting season, 41
nests were laid at Votlo Beach, Epi
Island, and, during the 2014/2015
nesting season, three females laid 15
nests (WSB 2011, 2016). It is not
possible to estimate nest trends due to
non-standardized methods and
opportunistic monitoring efforts over
time, which render existing data
incomparable and do not meet our
criteria (i.e., nest count data consistently
collected in a standardized approach for
at least 9 years).
In addition to an overall declining
nest trend, the West Pacific DPS
exhibits low reproductive output (i.e.,
low hatching success), due in part to a
combination of past and current threats
(e.g., beach erosion, predation, and
beach temperatures).
The DPS exhibits low productivity
(i.e., low hatching success), and the
overall nest trend is declining, likely
due to anthropogenic and
environmental impacts at nesting
beaches and in foraging habitats (Tiwari
et al. 2013). We conclude that the
declining nest trend and low
reproductive output place the DPS at
elevated extinction risk, especially
given the low nesting female
abundance.
Spatial Distribution
The West Pacific DPS nests
throughout four countries with a broad,
diverse foraging range. It exhibits
metapopulation dynamics and fine-scale
population structure.
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Aerial surveys conducted between
2004 and 2007 identified Indonesia,
Papua New Guinea and Solomon
Islands as the core nesting areas for the
DPS (Benson et al. 2007a; Benson et al.
2007b; Benson et al. 2011; Benson et al.
2018b). During the nesting season,
nesting females generally stayed within
300 km or less of these nesting beaches,
although a few females were
documented visiting multiple beaches
during a nesting season (Benson et al.
2007b). Distributing nesting activity
among various habitats may help to
buffer some of the population from
impacts at a single nesting area, but the
majority of females utilize one nesting
area during a nesting season (Benson et
al. 2011).
Migration and foraging strategies vary
based on nesting season, likely due to
prevailing offshore currents and
seasonal monsoon-related effects
experienced by the turtles as hatchlings
(Gaspar et al. 2012). The lack of
crossover among seasonal nesting
populations suggests that leatherback
turtles develop fidelity for specific
foraging regions, likely based on
juvenile dispersal patterns (Benson et
al. 2011; Gaspar et al. 2012; Gaspar and
Lalire 2017). Oceanic currents help to
structure the spatial and temporal
distribution of juveniles and lead them
to foraging and developmental habitats
(e.g., the North Pacific Transition Zone)
and to undertake seasonal migrations
seeking favorable oceanic habitats/
temperatures and abundant foraging
resources, such as the central California
ecoregion (Gaspar and Lalire 2017).
Inter-annual or long-term variability in
dispersal patterns can influence
population impacts or resilience to
regional or Pacific Ocean perturbations
(e.g., exposure to fisheries, ENSO
events, etc.). Stable isotopes, linked to
particular foraging regions, confirm
nesting season fidelity to specific
foraging regions (Seminoff et al. 2012).
Size differences are also apparent, with
slightly larger adults appearing to
exploit distant temperate foraging
habitats regardless of nesting season
(Benson et al. 2011; Lontoh 2014).
Summer nesting females forage in
Northern Hemisphere habitats in Asia
and the Central North Pacific Ocean,
while winter nesting females forage in
tropical waters of the Southern
Hemisphere in the South Pacific Ocean
(Benson et al. 2011; Harrison et al.
2018). This variance in foraging strategy
results in a foraging range that covers
much of the Pacific Ocean: Tasman Sea;
East Australian Current; eastern and
western South Pacific Ocean;
Indonesian, Sulu and Sulawesi, and
South China Seas; North Pacific
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Transition Zone; equatorial currents;
and central California ecoregion
(Benson et al. 2011; Lontoh 2014;
Harrison et al. 2018; Jino et al. 2018).
Different strategies result in
demographic differences within the DPS
which may affect productivity and
reproductive output. For example,
leatherback turtles that exploit distant
temperate foraging habitats (e.g., central
California) may require multiple years
of seasonal foraging before returning to
nesting beaches, due to greater energetic
demands. In contrast, leatherback turtles
exploiting geographically closer, yearround prey resources in more tropical
habitats (e.g., Sulu Sulawesi and South
China Seas) may remigrate annually
(Lontoh 2014).
The DPS also exhibits genetic
population structure. While mtDNA
analyses of 106 samples from Indonesia,
Papua New Guinea, and Solomon
Islands did not detect genetic
differentiation among nesting
aggregations (Dutton et al. 2007),
microsatellite DNA analyses indicate
fine-scale genetic structure (Dutton
2019; NMFS SWFSC unpublished data).
The wide distribution and variance in
foraging strategies likely buffers the DPS
to some degree against local
catastrophes or environmental changes
that would limit prey availability. The
distribution of nesting beaches
throughout four countries, although
primarily concentrated in three, helps to
buffer the entire DPS from major
environmental catastrophes, because
disturbances are not likely to similarly
affect all countries during the same
seasons. Additionally, the fine-scale
genetic structure among nesting
aggregations is indicative of
metapopulation dynamics, which may
also provide the DPS with some
resilience.
Diversity
The West Pacific DPS exhibits genetic
diversity, with six haplotypes identified
in 106 samples from Solomon Islands,
Papua Barat Indonesia, and Papua New
Guinea (Dutton 2006; Dutton et al. 2007;
Dutton and Squires 2008). This may
provide the DPS with the raw material
necessary for adapting to long-term
environmental changes, such as cyclic
or directional changes in ocean
environments due to natural and human
causes (McElhany et al. 2000; NMFS
2017). The population also exhibits
temporal nesting diversity, with various
proportions of the population nesting
during different times of the year
(summer versus winter) which helps to
increase resilience to environmental
impacts. The foraging strategies are also
diverse, with turtles using seven
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ecoregions of the Pacific Ocean. Diverse
foraging strategies likely provide some
resilience against local reductions in
prey availability or catastrophic events,
such as oil spills or typhoons, by
limiting exposure from a single event to
only a portion of the DPS. We conclude
that diversity within the DPS provides
it with some level of resilience to
threats.
Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
The destruction or modification of
habitat is a threat to this DPS. Primary
impacts to nesting beaches include
erosion and ocean inundation, which
may be caused by natural processes.
Nesting beaches of the West Pacific
DPS are dynamic, high profile beaches
that are subject to erosion, such as
during King Tides (naturally occurring,
predictable highest tides), which are
common seasonal occurrences. In
Indonesia, the Bird’s Head Peninsula
beaches are also subject to seasonal
patterns of erosion and accretion.
Changes in the currents brought on by
monsoons beginning in September
cause major erosion at Jamursba-Medi
that often removes the entire beach,
making the habitat unsuitable for
nesting until accretion begins again in
March (Hitipieuw et al. 2007). This
natural erosion has been documented to
impact many nests at Jamursba-Medi
(Hitipeuw et al. 2007). Arguably,
western Pacific leatherbacks have been
dealing with such changes in beach
habitats over time, and a turtle’s long
reproductive lifespan in general is
designed to sustain nest loss during a
few bad years or seasons. For example,
during the 2003/2004 nesting season, 80
percent of marked nests at JamursbaMedi (Warmamedi beach) washed away
before they hatched (Hitipeuw et al.
2007). However, given the low
abundance of the population, the loss
(or continued loss over time) of nests is
a concern.
At Wermon, the inundation of nests
from high tides is a threat during the
winter months. During the 2008/2009
winter nesting season, 26 percent of
nests laid at Wermon were inundated by
tidal activity (Wurlianty and Hitipeuw
2009). During the 2004/2005 nesting
season, 23 percent of nests were lost to
inundation (Wurlianty and Hitipeuw
2005). During the 2003/2004 nesting
season, 10.7 percent of all nests at
Wermon were below the high water
mark and were subsequently washed
away by high tides (Hitipeuw et al.
2007). Tapilatu and Tiwari (2007)
stressed that any management plan
developed for Papua will need to
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address the impact of inundation and
beach erosion.
Beach erosion is also a threat to nests
in Papua New Guinea, where strong
storms and tidal surges result in
substantial erosion and changes to
beaches throughout the Huon Coast. For
example, much of the Labu Tale nesting
beach was lost to erosion during the
2012/2013 nesting season (Pilcher
2013). The differences in beach width
along the Huon Coast place some
beaches at more risk of inundation and
erosion, such as Kamiali Beach, which
is half the width and significantly
narrower than Busama Beach (Pilcher
2008). At Kamiali, the average distance
of nests to the sea was 3.2 m, compared
to 6.2 m at Busama; the distances to the
vegetation line were comparable across
sites (1.3 m and 1.7 m, respectively;
Pilcher 2013).
In Vanuatu, there has been low
hatching success in some years due to
storms, floods, and high water (Petro et
al. 2007; WSB 2016).
In recent years, management and
conservation practices have included
relocating erosion-prone nests to bolster
hatchling production. However, these
projects are funding-dependent
throughout the range of the West Pacific
DPS. At Jamursba-Medi, ‘‘doomed’’
nests (i.e., those that are likely to be lost
to erosion or inundation) are sometimes
relocated to a more stable section of
beach; 15 nests were relocated during
the 2017 summer nesting season (Tiwari
et al. in prep.). At Wermon, nests are
relocated to avoid erosion and tidal
inundation, and increasingly due to
Ipomea root invasion (Tiwari et al. in
prep), but beach management activities
are project-dependent. At Wermon
during the 2017/18 winter nesting
season, nests could not be relocated
because of the lack of permission from
the beach owners, and all but three
nests washed away (Tiwari et al. in
prep). In Papua New Guinea, 22 of 47
nests (47 percent) at Kamiali beach were
relocated to protect them from storm
surge and erosion during the 2011/2012
nesting season, and 41 percent of nests
were relocated during the 2009/2010
season (Pilcher 2012). In the Solomon
Islands, efforts to relocate ‘‘doomed’’
nests is an ongoing and necessary
management strategy to help bolster
hatchling production, given that a large
proportion of nests are inundated or
have very low hatching success (Goby et
al. 2010; TDA 2013; Jino et al. 2018).
A large, significant portion of nests
(i.e., 10.7 percent to nearly all) are
exposed to the reduction and
modification of nesting habitat, as a
result of erosion and inundation. This
threat impacts the DPS by reducing
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nesting and hatching success, which has
been documented throughout the
nesting range of the DPS (NMFS and
USFWS 2013; Bellagio Sea Turtle
Conservation Initiative 2008). While
West Pacific leatherback turtles have
undoubtedly evolved to sustain changes
in beach habitats given their proclivity
to select highly dynamic and typically
narrow beach habitats, and therefore at
the population level can sustain some
level (albeit unquantified level) of nest
loss. However, the increasing frequency
of storms and high water events perhaps
as a result of climate change can result
in increased and perhaps unnatural loss
of nests. Such impacts may lower the
productivity of the DPS. Based on the
information presented above, we
conclude that habitat loss and
modification is a threat to the DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
The primary threat to the West Pacific
DPS is the harvest (both legal and
illegal) of leatherback turtles and their
eggs. Leatherback turtles are protected
by regulatory mechanisms in all four
nations where the DPS nests, but laws
are largely ignored and not consistently
enforced. This is due to the extreme
remoteness of beaches, customary and
traditional community-based ownership
of natural resources (which includes sea
turtles), and overall lack of institutional
capacity and funding for enforcement.
Furthermore, the cultural and socioeconomic dynamics in these nations
confound community buy-in and
conservation efforts (Kinch 2006;
Gjersten and Pakiding 2012; von Essen
et al. 2014). Additionally, there are
nuances related to indigenous harvest
(and the definition thereof), some of
which is permitted in these nations.
Turtle poaching affects both nesting
females on beaches and turtles in their
foraging habitats (Bellagio Sea Turtle
Conservation Initiative 2008; Kinch
2009; Suarez and Starbird 1996; Tiwari
et al. 2013; WWF 2018). Turtle poaching
has been documented in all four
countries where this DPS nests. Egg
poaching is a well-documented threat
(past and current) and is widespread
throughout the range of the DPS
(Bellagio Sea Turtle Conservation
Initiative 2008; NMFS and USFWS
2013; Tiwari et al. 2013; Tapilatu et al.
2017).
In Indonesia, the poaching of turtles
and eggs continues to occur, though egg
harvest and exploitation of females has
been minimized at Jamursba-Medi and
Wermon beaches due to the presence of
monitoring programs and educational
outreach. Large-scale egg poaching
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occurred at Jamursba-Medi between
1980 and 1993, whereby approximately
4 to 5 boats per week (from May to
August) collected 10,000 to 15,000 eggs
per boat (Tapilatu et al. 2013).
Commercial egg harvest has been
effectively eliminated since beach
monitoring was established at that
beach in 1993 (Hitipeuw et al. 2007).
However, recent survey efforts indicate
that most, if not all, sea turtle eggs
(including leatherback turtles) are
poached at other Bird’s Head Peninsula
beaches and sold in local markets
(Tapilatu et al. 2017). At Buru Island,
Indonesia, between 2016 and 2017,
eight females were poached (WWF
2018), and over the past 20+ years, three
to five nesting females have likely been
taken annually (J. Wang, NMFS, pers.
comm., 2018). In 2017, 114 of 203
leatherback nests were harvested at
Buru Island (WWF 2018). In 2018, due
to education provided by the newly
established WWF program on Buru
Island, local community-based efforts in
four villages now prohibit female and
egg harvest. While protective laws exist
in Indonesia, enforcement is largely
lacking in areas where monitoring
programs do not exist.
In Indonesia, foraging leatherback
turtles are also harvested in the waters
of the Kei Islands, Maluku Province,
where a recognized indigenous
subsistence harvest of immature and
adult turtles (average size 145 to 170
cm; range 52 to 203 cm) occurs and has
likely been a key feature of the local
traditional culture for centuries
(Compost 1980; Hamman et al. 2006;
Hitipeuw and Lawalata 2006, 2008).
Within the Kei Islands, customary law
(‘‘hak adat’’) authorizes a ritual
leatherback turtle hunt in the nine
villages of the traditional kingdom of
the Nufit people. Starbird and Suarez
(1994) brought attention to this hunt
when they reported that approximately
200 turtles were harpooned in three
months (October to December) of 1994,
with as many as 13 taken in one day.
Over the past three decades, sporadic
monitoring efforts have estimated that
up to 100 individuals are harvested
annually (Suarez and Starbird 1996;
Hitipeuw and Lawalata 2008; WWF
2018). At one point, it was assumed that
harvest pressure had declined and was
no longer an issue (NMFS and USFWS
2013). However, recent surveys indicate
that harvest continues, with
conservative estimates of 431 turtles
killed over an 8-year period (an average
of 53.9 turtles annually), typically
between August to February (Hitipeuw
and Lawalata 2008), and at least 103
turtles harvested in 2017 (WWF 2018).
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Most concerning perhaps is that some of
the turtle meat harvested may be
commercially sold as dried meat (i.e.,
leatherback ‘‘jerky’’ locally known as
dendeng), which is illegal to sell and
inconsistent with indigenous traditional
practices. Of four genetic samples
acquired in 1995 from turtles harvested
in the Kei Islands, three were assigned
to Birds Head Indonesian region and the
fourth sample was not definitive (66
percent probability, with 34 percent
probability to Solomon Islands),
although it could also be from the
Indian Ocean or from an undetermined
location (NMFS SWFSC unpublished
data 2018).
In Papua New Guinea, turtle and egg
poaching is a major threat despite the
fact that leatherback turtles have been
protected since the 1976 Fauna
(Protection and Control) Act. The illegal
take of both eggs and turtles likely
continues throughout the country due to
lack of community-based awareness,
reliance on traditional communitybased practices, institutional capacity,
and law enforcement (Bellagio Sea
Turtle Conservation Initiative, 2008).
The killing of nesting females has also
been well documented throughout
Papua New Guinea (Bellagio Sea Turtle
Conservation Initiative 2008; Kinch
2009; Pilcher 2013). For example, at
Bougainville Island, surveys of
community members identified that 21
nesting females were poached during
the last decade (Kinch 2009). However,
the harvest of eggs is likely the most
prolific threat in Papua New Guinea. If
unprotected, egg harvest (compounded
by intense dog predation described
below) resulted in the loss of 70 to 100
percent of nests (Quinn and Kojis 1985;
Hirth 1993; Bellagio Sea Turtle
Conservation Initiative 2008; Pilcher
2013). For example, during a one week
survey in January 2009 at Bougainville
Island, almost 100 percent of the 46
documented nests were poached (Kinch
2009). It is likely that near total egg
collection occurred throughout the
Huon Coast between World War II and
the establishment of the Huon Coast
Leatherback Turtle Monitoring and
Conservation Program in 2003 (Bellagio
Sea Turtle Conservation Initiative 2008;
Pilcher and Chaloupka 2013; Pilcher
2013). The Huon Coast Project, which
operated between 2003 and 2013,
helped to reduce egg and turtle harvest
due to program involvement and
community incentive funds received in
exchange for non-harvest agreements
(Pilcher 2013). As a result of the
program, hatchling production (i.e.,
percent of eggs yielding hatchlings)
increased from zero to approximately 60
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percent (Pilcher 2009, 2011, 2013;
WPRFMC 2015). The Project ended in
2013, and unfortunately egg harvest
resumed since there was no incentive
for communities to maintain their noharvest agreements (John Ben, Huon
Coast Leatherback Turtle Project, pers.
comm., 2020).
In Vanuatu and the Solomon Islands,
the poaching of females and collection
of eggs is also well documented
(Bellagio Sea Turtle Conservation
Initiative 2008; NMFS and USFWS
2013). In Vanuatu, MacKay et al. (2014)
reported the harvest of five nesting
females between 1999 and 2008.
However there is a general
understanding that nesting females were
typically harvested (Petro et al. 2007).
Of the 315 nests documented on Isabel
Island, Solomon Islands during a
January 2011 site visit at Sasokolo and
Litogahira beaches, the majority of nests
had been poached (Tiwari 2011
unpublished data). Historically, nearly
all nesting females and eggs were
poached on Redova for consumption
(Tiwari 2011 unpublished data). In
response, financial incentive programs
have been established to protect nests
and females whereby villagers are paid
a financial reward for each nest that
hatches successfully (TDA 2013). On
Vangunu Island, 10 to 20 nesting
females were poached annually, in
addition to near-total egg collection
(Jino et al. 2018). In response to
declining population trends, the
community declared a moratorium on
the harvest of leatherback turtles in
1999 (Jino et al. 2018), and a community
incentive program providing financial
awards has helped to reduce harvest
pressure (TDA 2013). Despite these
efforts and protective legislation, the
poaching of females and eggs likely
persists throughout the Solomon Islands
(TDA 2013: Tiwari 2011 unpublished;
MacKay et al. 2014).
Within the West Pacific DPS, many
nesting females, foraging turtles, and
eggs are exposed to both illegal
poaching and legal harvest. The taking
of turtles reduces abundance. The taking
of nesting females reduces both
abundance and productivity. Such
impacts are high because they directly
remove the most productive individuals
from the DPS, reducing current and/or
future reproductive potential. Egg
harvest reduces productivity; the
persistent, and near-total (at some
locations) collection of eggs guarantees
that future population recruitment (i.e.,
nesting female abundance) will be
reduced or eliminated. Given the
declining nesting trend and current
nesting female abundance of this DPS,
the continued and unregulated poaching
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or harvest of leatherback turtles and
eggs is unsustainable. Further, the
harvest of approximately 100 foraging
leatherback turtles annually at the Kei
Islands, Indonesia is likely an
unsustainable practice given the current
low abundance of the population. We
conclude that overutilization is a major,
and the primary, threat to the West
Pacific DPS, accelerating its risk of
extinction.
Disease or Predation
While we could not find any
information on disease for this DPS,
predation of eggs is a major and welldocumented threat to the West Pacific
DPS, likely second to poaching (i.e.,
nests not taken by humans are typically
predated; Bellagio Sea Turtle
Conservation Initiative 2008).
In Indonesia, predation of eggs by
feral pigs, feral dogs, and monitor
lizards has been documented, with feral
pig predation being the most
detrimental (Hitipeuw and Maturbongs
2002; Tapilatu and Tiwari 2007;
Bellagio Sea Turtle Conservation
Initiative 2008). Nest predation by
domestic and/or feral dogs has been
recorded in both Jamursba-Medi and
Wermon. Predation of nesting females
by crocodiles has also been documented
at Wermon beach (Bellagio Sea Turtle
Conservation Initiative 2008; UNIPA,
pers. comm., 2018). At Jamursba-Medi,
between June and July of 2005, 29.3
percent of nests were destroyed by pigs
(Tapilatu and Tiwari 2007). Intensive
management effort at Jamursba-Medi
reduced feral pig predation of nests to
five percent during the 2016 and 2017
nesting seasons (Tiwari et al. in prep).
Feral pigs and dogs depredated 17.5
percent of all nests at Wermon during
the 2003 and 2004 winter nesting season
(Hitipeuw et al. 2007). At Wermon, 21
percent of nests were lost to predation
during the 2004/2005 nesting season
(Wurlianty and Hitipeuw 2005). At Buru
Island in 2017, 16 nests were lost to
predation by dogs, wild boar, lizards, or
saltwater crocodiles (WWF 2018).
In Papua New Guinea, predators of
eggs include feral dogs, monitor lizards,
and ghost crabs (Kinch 2009).
Depredation of nests by village dogs was
determined to be an intense threat to
nests, with dogs consuming all nests
laid during the 2003/2004 and 2004/
2005 nesting seasons at Kamiali beach
(Pilcher 2006; I. Kelly, NMFS, pers.
comm., 2018). Predation of nesting
females by crocodiles has also been
documented in a number of locations in
Papua New Guinea (Bellagio Sea Turtle
Conservation Initiative 2008; Kinch
2009). To protect nests, Huon Coast
communities developed and placed
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bamboo grids over nests to prevent dogs
from preying on the eggs (Pilcher 2006;
2009). This, along with efforts to reduce
egg harvest by humans, resulted in
increased hatching production from
zero to approximately 60 percent
between 2006 and 2013, with over 2,300
nests saved producing approximately
100,000 hatchlings (Pilcher 2009; 2011;
2013; WRFMC 2015). However, this
project ended in 2013, and it is
unknown if egg protection continues, or
if nest predation has resumed.
In this DPS, a large proportion of eggs
are exposed to predation, especially by
dogs and pigs. Predation primarily
results in the loss of eggs, and the
impact of this threat is a reduction of
productivity. Though leatherback turtles
generally produce a large number of
eggs and hatchlings, predation is
widespread throughout the range of the
DPS, and in some areas, predation rates
are as high as 100 percent. We conclude
that predation poses a threat to the West
Pacific DPS.
Inadequacy of Existing Regulatory
Mechanisms
The West Pacific DPS is protected by
several regulatory mechanisms. For
each, we review the objectives of the
regulation and to what extent it
adequately addresses the targeted threat.
Leatherback turtles are protected by
legislation in all four of the nations
where the West Pacific DPS nests
(Indonesia, Papua New Guinea,
Solomon Islands, and Vanuatu). It is
generally illegal to harvest leatherback
turtles and their eggs. However, laws are
not typically enforced or followed given
customary marine tenure systems that
dictate near-shore rights. Lack of
enforcement or implementation of
protective laws may be due to: Overall
lack of in-country institutional capacity
and funding for enforcement; the
extreme remoteness of beaches;
customary marine tenure or traditional
community-based ownership of natural
resources in these nations (which
includes sea turtles; Kinch 2006;
McDonald 2006) and regulatory
government-led legislation, which may
be incompatible with traditional
practices (von Essen et al. 2014). There
are also nuances related to indigenous
harvest (and the definition thereof),
which is not prohibited in these nations.
As a result, most leatherback nesting
beaches with the exception of JamursbaMedi and Wermon (i.e., beaches with
established long-term monitoring
programs) are not currently protected
(or only minimally protected) from
harvest or poaching of eggs, nesting
females, or other anthropogenic threats.
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In Indonesia, all sea turtles are
protected by law, but there are
allowances for indigenous peoples
(although indigenous provisions are not
clearly defined). The 1990 Government
Regulation Act number 5 concerning the
Conservation of the Natural Resources
and the Ecosystem, makes the trade of
protected wildlife illegal, and those
found liable can be punished to a
maximum of 5-year prison term and
fined 100 million Indonesia Rupiah
(approximately 6,500 USD). The
protection of all sea turtle species
(Government Regulation No. 7 on
Preserving Flora and Fauna Species)
came into effect in 1999 (Zainudin et al.
2007). The use of protected wildlife is
allowed for the purposes of research,
science, and rescue of the wildlife itself.
While the trade and exploitation of
turtles is illegal in Indonesia, there still
exists a documented harvest of green
turtles in Bali, which contributes to
public confusion regarding sea turtle
protections (Westerlaken 2016).
In Papua New Guinea, the leatherback
turtle is the only species protected
under the 1976 Fauna (Protection and
Control) Act, which makes killing of
leatherback turtles or taking of
leatherback turtle eggs illegal, with fines
of 500 to 1000 kina (approximately 100
to 300 USD). Any person who buys or
sells or offers for sale, or has in
possession leatherback turtle eggs or
meat can also be fined. The Act makes
provisions for persons with customary
rights to take turtles, but states that sea
turtles cannot be taken, killed, or sold
from May through July (Kinch 2006).
This is typically the nesting season for
hard-shelled sea turtle species, but
leatherback turtles nest primarily during
the winter months (November to
February). As with most Melanesian
countries, lands are locally-owned and
managed, and the national government
has little influence outside major cities
(Kinch 2006).
The Solomon Islands Fisheries Act
(1993) regulations protect nesting turtles
and eggs during the breeding season
(June to August and November to
January); prohibit the sale, purchase, or
export of sea turtle species or their
parts; and contain specific protections
for leatherback turtles. In the Solomon
Islands, more than 85 percent of the
land is held under customary (locallymanaged) marine tenure, and the vast
majority of the population still lives in
rural areas making a living from the
natural resources on those lands. For
centuries, communities have practiced
traditional models of resource
stewardship, making implementation
and enforcement of national regulations
nearly impossible. Instead, natural
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resource governance must originate
from chiefs and village leaders, which
requires extensive educational outreach
to encourage traditional approaches that
may be supported by legal or ‘modern’
enforcement measures (McDonald
2006).
Fisheries Regulations under the
Vanuatu Fisheries Act (2009) prohibit
the take, harm, capture, disturbance,
possession, sale, purchase of or
interference with any turtle nest (or any
turtle in the process of nesting) and the
import, or export of green, hawksbill,
and leatherback turtles or their products
(shell, eggs, or hatchlings). The Act also
prohibits the possession of turtles in
captivity. A person may apply in
writing to the Director of Fisheries for
an exemption from all or any of these
provisions for the purposes of carrying
out customary practices, education,
and/or research. Similar to Papua New
Guinea and the Solomon Islands,
natural resource governance in Vanuatu
is best directed, realized, and
implemented at the community level
and not via national legislation.
Fortunately, traditional practices are
experiencing a renaissance in Vanuatu
and may complement current regulatory
marine resource management efforts
(Hickey et al. 2006).
Throughout the foraging range of the
DPS, there are numerous regulatory
mechanisms that protect turtles within
the DPS. These include: RFMOs such as
the Western and Central Pacific
Fisheries Commission (WCPFC) and the
IATTC and fisheries management
regulations in 32 nations where this
DPS may occur (Harrison et al. 2018).
The WCPFC adopted a Sea Turtle
Conservation and Management Measure
(CMM 2008–03) to mitigate the impacts
on turtles from commercial shallow-set
fisheries operating in the Western and
Central Pacific Ocean. The measure
included the adoption of FAO (2009)
guidelines to reduce sea turtle mortality
through safe handling practices and to
reduce bycatch by implementing one of
three methods by January 2010. The
three methods to choose from are: (1)
Use only large circle hooks with offsets
of ≤10°; (2) use whole finfish bait; or (3)
use any other mitigation plan or activity
that has been approved by the
Commission. This sea turtle
conservation measure is specific to selfidentified shallow-setting, swordfishtargeting fleets. It does not apply to the
international Pacific longline deep-set
tuna-targeting fisheries, which comprise
the majority of the longline fisheries and
are also known to interact with
leatherback turtles (Lewison et al. 2004;
Beverly and Chapman 2007; Roe et al.
2014; Wallace et al. 2013). Technical
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analysis of the sea turtle conservation
measure found a very small percentage
of shallow-set fisheries to be in
compliance, with less than one percent
of Western and Central Pacific Ocean
longline effort implementing mitigation
measures, even though approximately
20 percent of longline effort consists of
shallow sets (Clarke 2017). Further,
many RFMO members are not meeting
the five percent observer coverage
requirement resulting in limited bycatch
reporting (Clarke 2017).
In summary, regulatory mechanisms
exist to protect leatherback turtles and
their eggs throughout the range of this
DPS. However, most are inadequate to
reduce the threat that they were
designed to address due to a lack of
implementation or enforcement or
inclusion of provisions for indigenous
harvest. Regulations are also misaligned
with established traditional practice and
management systems. As a result,
poaching and bycatch remain major
threats to the DPS. In summary, we
consider the inadequacy of the
regulatory mechanisms to be a threat to
the DPS.
Fisheries Bycatch
Fishery bycatch in coastal and pelagic
fisheries is a major threat to the West
Pacific DPS, which is exposed to
domestic and international fisheries
throughout its extensive foraging range.
At-sea bycatch of leatherback turtles has
been documented for a variety of gillnet
and longline fisheries in the Pacific
Ocean, but little is known about the
total magnitude or full geographic
extent of mortality. Satellite telemetry
studies have identified movements and
revealed fidelity to foraging regions of
the DPS, specifically in habitats of the
North Pacific Ocean, southwestern
Pacific Ocean, and Indo-Pacific tropical
seas (Bailey et al. 2012; Benson et al.
2011, Seminoff et al. 2012; Roe et al.
2014). The summer nesting component
of the population exhibits strong site
fidelity to the central California foraging
area (Benson et al. 2011) which puts
them at risk during migrations of
interacting with U.S. and international
pelagic longline fleets operating
throughout the Central and North
Pacific Oceans. For example, several of
the turtles tagged in Papua Barat,
Indonesia were known or suspected to
have been killed in fisheries operating
off Japan, Philippines, and Malaysia
(Benson et al. 2011).
Historically, significant leatherback
bycatch was documented in the North
Pacific high seas driftnet fishery, which
expanded rapidly during the late 1970s
but was banned in 1992 by a UN
resolution (summarized in Benson et al.
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2015). Wetherall et al. (1993) estimated
that over 750 leatherback turtles were
killed in Japanese, Korean, and
Taiwanese driftnet fisheries during the
1990 to 1991 season, with potentially
5,000 to 10,000 leatherback turtles
bycaught between the late 1970s and
1992. Based on current knowledge of
movement patterns (Benson et al. 2011),
the majority of these bycaught turtles
would have originated from western
Pacific nesting beaches after their boreal
summer nesting period. Thus, high seas
driftnet fishery bycatch was likely a
significant contributor to the population
declines observed at nesting beaches
during the 1980s and 1990s (Benson et
al. 2015).
Many nations are involved in longline
fishing in the Pacific Ocean, where two
types of vessels are used: (1) Large
distant-water freezer vessels that
undertake long (months) voyages and
operate over large areas of the region;
and (2) smaller offshore vessels with ice
or chill capacity that typically
undertake trips of about one month.
Target species are yellowfin, bigeye,
albacore tuna, and swordfish. The total
annual number of longline vessels in the
western and central Pacific region has
fluctuated between 3,000 and 6,000 for
the last 30 years, including the 100 to
140 vessels in the Hawaii longline
fisheries (NMFS 2018).
Pelagic Fisheries
International longline fisheries are
characterized by inconsistent reporting
and traditional gear configurations,
including J-style hooks with squid bait,
which result in higher interaction and
mortality rates than for modified gear
(Beverly and Chapman 2007; Lewison et
al. 2004; Swimmer et al. 2017). For
example, the Taiwan and China tuna
longline fisheries are estimated to have
bycatch rates several times higher than
Hawaii longline fisheries (Bartram and
Kaneko 2008; Chan and Pan 2012).
Analyzing multi-national turtle bycatch
data from 1990 to 2004, Molony (2005)
found that the purse seine fishery and
the deep, shallow, and albacore longline
fisheries (operating between 15° N and
31° S) take an average of about 100
leatherback turtles annually. Lewison et
al. (2004) collected fish catch data from
40 nations and turtle bycatch data from
13 international observer programs to
estimate global longline bycatch of
loggerhead and leatherback turtles in
2000. In the Pacific Ocean, they
estimated 1,000 to 3,200 leatherback
turtle (juvenile and adult) mortalities
from pelagic longlining in 2000
(Lewison et al. 2004). Using effort data
from Lewison et al. (2004) and bycatch
data from Molony (2005), Beverly and
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Chapman (2007) estimated sea turtle
longline bycatch to be approximately 20
percent of that estimated by Lewison et
al. (2004), approximately 200 to 640
leatherback turtles annually. These
estimates include turtles from the East
and West Pacific DPS. While the results
of each of these studies may be feasible,
the Lewison et al. 2004 estimates were
based on available data at that time (i.e.,
less than 30 percent of longline fishing
effort) that was skewed toward fishing
fleets with relatively better management
and data reporting systems, and hence
extrapolations may have overestimated
interaction rates (Clarke et. al. 2014).
However, Beverly and Chapman (2007)
applied different catch per unit effort
(CPUE) estimates in calculations
differentiated between deep-set and
shallow-set fisheries which have
different interaction rates and, hence,
their estimates may be more realistic.
Despite scientific evidence showing
that use of circle hooks and finfish bait
significantly reduces leatherback turtle
bycatch rates in longline fisheries
(Gilman et al. 2007; Swimmer et al.
2017), nations are not required to use
this hook/bait combination. The WCPFC
Sea Turtle Conservation and
Management Measure (CMM 2008–03)
only applies to fleets using shallow-set
gear targeting swordfish. Additionally,
observer program coverage levels in
WCPFC longline fisheries have not
reached the required five percent
coverage rate, resulting in limited
bycatch reporting and likely
underreporting (Clarke 2017). Further,
existing sea turtle mitigation measures
are currently only being applied to
approximately one percent of shallowset longline fisheries in the Convention
Area, even though approximately 20
percent of the longline effort consists of
shallow-sets (Clarke 2017).
A workshop convened to assess the
effectiveness of WCPFC’s Sea Turtle
Conservation and Management Measure
found limited reductions in interactions
and mortalities (Clarke 2017). Fishery
observer data collected between 1989
and 2015 of 34 purse seine and longline
fleets across the Pacific documented a
total of 2,323 sea turtle interactions, of
which 331 were leatherback turtles
(Clarke 2017). Two bycatch hotspot
areas were identified: One in central
North Pacific (which likely reflects the
100 percent observer coverage in the
Hawaii shallow-set longline fishery) and
a second hotspot in eastern Australia
(Clarke 2017). However, analysis of the
data also found that overall
conservation benefits would have been
greater had mitigation measures also
been applied to deep-set gear and not
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only to shallow-set swordfish fisheries
(Clarke 2017).
While bycatch in pelagic shallow-set
swordfish-targeting longline fisheries
has received the most attention to date,
comparable studies for deep-set tunatargeting fisheries are not available due
to the more complex nature of these
fisheries. There may be fewer
interactions because deep-set fisheries
(operating at depths more than 60 m)
generally have lower bycatch rates, but
they also have higher mortality rates
than shallow-set gear (Lewison et al.
2004; Kaplan 2005; Gilman et al. 2007).
Pelagic deep-set tuna-targeting fisheries
cannot be ignored because they also
have the potential to interact with
leatherback turtles and constitute four
times greater effort than shallow-set
fisheries yet do not have RFMO gear
mitigation requirements (Clarke 2017).
Wallace et al. (2013), and a global
review based on that study (FAO 2014),
categorized longline and gillnet fisheries
interactions with West Pacific
leatherback turtles as high risk but low
impact for longline and gillnet gear,
likely due to insufficient data from this
data-poor region. Bycatch in small-scale
coastal fisheries has been a significant
contributor to population declines in
many regions (Kaplan 2005; Peckham et
al. 2007; Alfaro-Shigueto et al. 2011),
yet there is a significant lack of
information from coastal and smallscale fisheries, especially from the
Indian Ocean and Southeast Asian
region (Lewison et al. 2014).
Southeast Asian Fisheries
Waters of Southeast Asia are heavily
fished by a variety of gillnets, trawls,
fish traps, and a range of different hook
and line gears, involving hundreds of
thousands of fishers (FAO 2011). The
West Pacific DPS nests, migrates, and
forages throughout this densely
populated and heavily exploited coastal
region (Bellagio Sea Turtle Conservation
Initiative 2008; Benson et al. 2011;
Lewison et al. 2014; Roe et al. 2014;
Harrison et al. 2018).
There are few quantitative estimates
of fisheries interactions near nesting
beaches of this DPS, and existing reports
provide only brief snapshots of impacts
or are outdated. In Indonesia, between
1980 and 1993, shark gillnets off the
nesting beaches of Jamursba-Medi killed
two to three nesting females weekly
(Tapilatu et al. 2013). As a member of
the WCPFC and the IOTC, Indonesia
must comply with reporting
requirements and conservation
measures as required by these RFMOs.
In 2006, of the 85 sea turtle interactions
observed in 539 sets on 10 tuna longline
vessels, 3 were adult leatherback turtles
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(Zainudin et al. 2007). Leatherback
turtles are known to migrate through
and forage within Philippine waters
(Benson et al. 2011), and in 2014, aerial
surveys observed leatherback turtles
foraging in high density fishing areas
(130 to 381 boats; MRF 2010, 2014).
Leatherback turtles have also stranded
dead or injured on Philippine beaches
as a result of fishery interactions,
typically with gillnet gear (Bagarinao
2011; Cruz 2006; MRF 2010; MWWP
2018 unpublished). In Malaysia, bycatch
studies using an interview-based
approach revealed that four leatherback
turtles were caught in gillnets the prior
year (Pilcher et al. 2008).
Fisheries operating out of Australia
and New Zealand may result in high
bycatch and mortality rates for the
winter nesting component of the DPS
that migrates into the Southern
Hemisphere (MacKay et al. 2014;
Harrison et al. 2018). In Australia, some
bycatch records exist for pelagic
longline fisheries (Robins et al. 2002;
Stobutzki et al. 2006), prawn trawls off
Queensland and Northern Territory,
gillnet fisheries off Queensland and
Tasmania, and pot gear off Tasmania
(Limpus 2009). Gillnet sea turtle
bycatch is reported as widespread and
includes anecdotal reports of
leatherback turtles taken in Tasmanian
tuna gillnet fisheries (Limpus 2009).
Between 2004 and 2014, the
Australian shallow-set fishery had an
estimated 29 to 178 leatherback
interactions, based on two to 10
observations (average = 4.6 interactions)
and four to 10 percent observer coverage
(MacKay et al. 2014). These data are
similar to bycatch information
extrapolated from interviews with
Australian fishers (Robins et al. 2002)
which identified 162 leatherback turtles
interactions in 2001 (MacKay et al.
2014). Australia has a sea turtle
mitigation plan for its Eastern Tuna and
Billfish Fishery which sets ‘‘trigger
level’’ interaction rates of ≤0.0048
turtles per 1,000 hooks for each turtle
species or 0.0172 turtles per 1,000 hooks
overall (DAFF 2009 in Clarke et al.
2014). In 2013, Australia reported that
the trigger levels had been exceeded for
the third year in a row and as a
consequence the Australian Fisheries
Management Authority required that
shallow-set vessels in these fisheries use
large circle hooks consistent with the
WCPFC sea turtle measure (CMM 2008–
03; Clarke et al. 2014).
In New Zealand, records document
288 instances of stranding or
commercial and recreational bycatch of
leatherback turtles from 1892 to 2015
(Godoy et al. 2016). New Zealand’s
surface longline fishery captured 90
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leatherback turtles between 2008 and
2015 (Godoy et al. 2016). This is likely
an underestimate because data were
based on low observer coverage (5.8
percent overall), with limited observer
overage during the peak time of
leatherback abundance in New Zealand
waters (January to March). Strandings
can also provide opportunities for
researchers to identify fisheries
interactions. MacKay et al. (2014)
identified 19 mortalities in New
Zealand and 29 mortalities in Australia.
Although the cause of most strandings
was often unknown, leatherback turtles
have been found entangled in crab pot
gear and monofilament fishing nets and
ropes. Longline fishing is concentrated
off southern Queensland and New
South Wales, Australia and is the
suspected cause of 41 percent of
strandings (n = 12). In Victoria,
Tasmania and South Australia, 61
percent of strandings (n = 17) involved
suspected entanglement in inshore
fishing gear and crab pots (MacKay et al.
2014).
U.S. Pacific Pelagic Fisheries
Detailed bycatch data are available for
U.S.-managed pelagic fisheries
operating in the central and eastern
Pacific Ocean due to regulatory
mandates and high levels of observer
coverage. Longline fisheries, based out
of Hawaii and American Samoa, may
interact with foraging turtles of the West
Pacific DPS. However, only two
interactions involved individuals of the
East Pacific DPS in 1995 and 2011 (P.
Dutton, NMFS, pers. comm., 2018).
Prior to 2001, the Hawaii longline
fishery was estimated to capture about
110 leatherback turtles annually,
resulting in approximately 9 annual
mortalities (McCracken 2000). Since
2005, the fishery has reduced its
estimated mortality to seven leatherback
turtles annually, and data confidence
increased significantly due to increased
observer coverage (NMFS 2018). The
fishery was closed in 2001 under court
order and re-opened in 2004 as two
separate fisheries: A shallow-set
swordfish-targeting fishery and a deepset tuna-targeting fishery. Management
requirements include: Gear modification
(e.g., circle hooks and fin-fish bait) and
handling measures designed to reduce
sea turtle bycatch rates and posthooking mortality in both fisheries; an
annual hard-cap limit on the number of
allowable interactions in the shallow-set
fishery; 100 percent observer coverage
in the shallow-set fishery; and 20
percent observer coverage in the deepset fishery (50 CFR 665 (Subparts A–C);
NMFS 2012, 2014, 2015). The shallowset fishery has been closed three
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additional times since reopening in
2004: In 2006, after reaching the hard
cap for loggerhead turtle interactions (n
= 17); in 2011, after reaching the hard
cap for leatherback turtle interactions (n
= 16); and in 2018 under a stipulated
settlement after the Ninth Circuit Court
of Appeals held that NMFS’ no jeopardy
determination for loggerheads in the
2012 biological opinion (9th Circuit
2017) was arbitrary and capricious. See
Turtle Island Restoration Network v.
U.S. Dep’t. of Commerce, 878 F.3d 725
(9th Cir. 2017). Since 2004, leatherback
turtle interactions in the shallow-set
component of the fishery have been
reduced by 84 percent from 0.03 to 0.01
BPUE as a result fisheries regulations
(Swimmer et al. 2017). Between 2004
and 2017, there have been 99 total
leatherback turtle interactions in the
shallow-set fishery (or approximately 8
turtles annually), based on 100 percent
observer coverage (WPRFMC 2018).
Between 2002 and 2016, an estimated
168 interactions may have occurred in
the Hawaii deep-set fishery (or
approximately 12 annually), based on
an extrapolation of data collected at a
level of 20 percent observer coverage
(WPRFMC 2018). Observer coverage of
the American Samoa longline fishery
has varied over time from 5 to 40
percent and has had an estimated 59
interactions between 2010 and 2017
(WPRFMC 2018).
The U.S. tuna purse seine fishery
operating in the Western and Central
Pacific Ocean anticipates up to 11
leatherback turtle interactions annually
(NMFS 2006). However, the fishery had
fewer interactions, with approximately
16 leatherback turtle interactions
between 2008 and 2015 based on
observer coverage ranging from 20 to
100 percent (NMFS unpublished data).
From 1990 to 2009, there were 24
observed leatherback turtle interactions
in the California drift gillnet fishery
based on 15.6 percent per year observer
coverage (Martin et al. 2015). Genetic
analyses indicated that almost all
originated from the West Pacific DPS
(Dutton et al. 1999; NMFS SWFSC
unpublished). In 2001, NMFS
implemented regulations (i.e., a large
time/area closure in Central California)
that reduced interactions by
approximately 80 to 90 percent, with
only two leatherback turtle interactions
(both alive) observed based on 20 to 30
percent observer coverage since
regulations were implemented (NMFS
West Coast Region unpublished). Drift
gillnet fishing is prohibited annually
from August 15 to November 15 within
the California leatherback turtle
conservation area. Currently, NMFS
anticipates up to 10 interactions (or 7
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mortalities) over a 5-year period (NMFS
2013).
In addition, nine fixed gear fisheries
operate off the U.S. West Coast,
including the Federally-managed
sablefish pot fishery and the statemanaged California Dungeness crab
fishery. Since 2008, only one
leatherback interaction has been
documented in the sablefish fishery
(NMFS 2013). The state-managed
Dungeness crab fishery may be a newly
emerging threat: Two documented
leatherback entanglements in pot gear
(mainline or surface buoy) occurred in
2015 and 2016. Fishing effort was high,
and the fishery had shifted into the
Central California region, which
overlaps somewhat with leatherback
foraging habitat (S. Benson, NMFS, pers.
comm., 2018). In 2019, the State of
California settled with a non-profit
organization in response to a complaint
that the commercial Dungeness crab
fishery was taking leatherback sea
turtles (and other large whales) without
authorization under section 10 of the
ESA. The California Dungeness crab
fishery closed in mid-April 2019 as part
of the settlement agreement and again
on May 15, 2020 (just the Central
Management Area), due to significant
risk of marine life entanglement. The
northern part of California remains open
until mid-July unless CDFW decides to
take further management action (i.e., if
risks to large whales and/or leatherbacks
is elevated in that area).
East Pacific Pelagic Fisheries
The West Pacific DPS has a vast transPacific range. Some individuals forage
in the East Pacific Ocean, where
leatherback turtles are caught in
fisheries of Peru and Chile (Donoso and
Dutton 2010; Alfaro-Shigueto et al.
2007, 2011, 2018). Of 59 leatherback
turtles caught in East Pacific fisheries,
an estimated 15 percent of individuals
sampled originated from the West
Pacific DPS (Dutton et al. 2000; Donoso
and Dutton 2010). Information compiled
by IATTC on sea turtle interactions with
pelagic longline fisheries operating in
the East Pacific is limited, given that
requirements for longline observer
coverage of five percent was only
implemented in January 2013 (Clarke et
al. 2014). Additional information on
East Pacific fisheries are presented in
the bycatch section for the East Pacific
DPS.
Summary of Fisheries Bycatch
We conclude that individuals of this
DPS are exposed to high fishing effort
throughout their foraging range, in
coastal waters near nesting beaches, and
when migrating to and from nesting
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beaches, though very little fisheries data
are available for coastal areas. Bycatch
rates in international pelagic and coastal
fisheries are high, and these fisheries
have limited management regulations
despite hotspots of high interactions in
Southeast Asia (Lewison et al. 2004,
2014; Alfaro-Shigueto et al. 2011;
Wallace et al. 2013; Clarke 2017).
Annual interaction and mortality
estimates are only available for U.S.managed pelagic fisheries, which
operate under extensive fisheries
regulations that are designed to
minimize the capture and mortality of
endangered and threatened sea turtles
(NMFS 2013; Swimmer et al. 2017;
NMFS 2018). Mortality reduces
abundance, by removing individuals
from the population; it also reduces
productivity, when nesting females are
killed. We conclude that fisheries
bycatch is a major threat to the West
Pacific DPS.
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Vessel Strikes
Vessel strikes are a threat to the West
Pacific DPS. Between 1981 and 2016,
there were 11 documented vessel strikes
in central California (NMFS West Coast
Region, unpublished data 2018). Many
vessel strikes are not reported, and
turtles are not recovered.
The range of the DPS overlaps with
many high-density vessel traffic areas.
Though the potential for exposure is
high, we are only aware of 11 vessel
strikes in recent decades. Vessel strikes
resulting in mortality would lower the
abundance of the DPS. However,
available data does not support
characterizing this as a high or moderate
impact. We conclude that vessel strikes
pose a threat to the DPS, albeit of less
concern than other impacts such as
overutilization and fisheries
interactions.
Pollution
Pollution includes contaminants,
marine debris, and ghost fishing gear.
Leatherback turtles can ingest small
debris, causing internal damage and
blockage. Larger debris can entangle
animals, leading to reduced mobility,
starvation, and death. Given the amount
of floating debris in the Pacific Ocean
(Lebreton et al. 2018), marine debris has
the potential to be a significant threat to
the DPS. Presently available data do not
allow for quantifying the precise extent
of the threat.
Leatherback turtles feed exclusively
on jellyfish and other gelatinous
organisms and as a result may be prone
to ingesting plastics resembling their
food source (Schuyler et al. 2013).
Lebreton et al. (2018) estimated plastic
debris accumulation to be at least
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79,000 (45,000 to 129,000) tonnes in the
Great Pacific Garbage Patch, a 1.6
million km2 of subtropical waters
between California and Hawaii. This
figure is four to 16 times greater than
previously reported. Entanglement in
ghost fishing gear is also a concern
(Gilman et al. 2016), and derelict nets
made up approximately 46 percent by
piece, and 86 percent by weight, of
debris floating in this area (Lebreton et
al. 2018). The highest risk areas within
the range of the West Pacific DPS where
animals may encounter significant
amounts of debris includes the north
Pacific gyre, the South China Sea, and
off of the east coast of Australia (Schuler
et al. 2015). However, WedemeyerStrombel et al. (2015) found no plastics
in the gastrointestinal tracts of two
leatherback carcasses from American
Samoan and Hawaiian longline fisheries
from 1993 to 2011. Clukey et al. (2017)
found no plastics in the gastrointestinal
tracts of three leatherback carcasses
from Pacific longline fisheries captured
between 2012 and 2016. However, it is
very difficult to obtain dead leatherback
turtles to study these effects, and given
the great amount of plastics within
environment, such results may
underestimate ingestion impacts.
Few studies of pollutants and their
effect on leatherback turtles were
available within the range of this DPS.
Harris et al. (2011) found the heavy
metal exposure in leatherback turtles
foraging off the coast of California to be
nine times higher than the St. Croix
nesting population, although levels
were not expected to be lethal. We do
not know if there were sub-lethal
effects. Stewart et al. (2011) found that
PCBs are more likely to be transferred
from females to their eggs than from the
environment to eggs.
Given the large amount of marine
debris within the range of the DPS, we
expect exposure to be high for all life
stages despite low sample sizes of
leatherback turtles with ingested marine
debris. Potential impacts include death
and injury. However, quantitative
estimates of such impacts are not
available. We conclude that pollution
may be a threat to the DPS.
Natural Disasters
The best available scientific and
commercial data indicate that natural
disasters are a threat to the DPS but do
not allow the impact to be quantified.
Natural disasters within the range of
this DPS include: Tsunamis, typhoons,
earthquakes, and flash floods. Such
environmental events are periodic, with
localized impacts that do not persist
over time. These events may reduce nest
incubation and hatching success in one
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season or at few locations. While
leatherback turtles have undoubtedly
evolved to sustain such natural impacts,
the increasing frequency of
environmental events as a result of a
changing climate, which can affect the
frequency and intensity of high tides
and large storms, may hamper
productivity and conservation activities
(Goby et al. 2010; S. Benson, NMFS,
pers. comm., 2018). Such events may
pose additional threats by depositing
marine debris on nesting beaches and in
occupied waters. The 2011 Japan
tsunami and the 2006 Indonesian
earthquake and resulting tsunami likely
deposited large amounts of debris (i.e.,
millions of tons) into the foraging and
migrating habitats of the DPS (Hafner et
al. 2014; NOAA 2015). We conclude
that natural disasters pose a potential
threat to the West Pacific DPS.
Climate Change
Climate change is a threat to the West
Pacific DPS. A warming climate and
rising sea levels can impact leatherback
turtles through changes in beach
morphology, increased sand
temperatures leading to a greater
incidence of lethal incubation
temperatures, changes in hatchling sex
ratios, and the loss of nests or nesting
habitat due to beach erosion (Benson et
al. 2015).
Elevated egg incubation temperatures
can lead to mortality. During the 2009/
2010 nesting season at the Huon Coast
(Papua New Guinea), Pilcher (2010)
found higher incubation temperatures
(32 to 33 °C) in exposed nests compared
to shaded nests (29 to 30 °C). Sea turtles
exhibit temperature-dependent sex
determination. The incubation
temperature determines sex ratios and
the duration of incubation (i.e.,
thermosensitive period). Along the
Huon Coast, incubation duration
decreased during the nesting season as
beach temperatures warmed. During the
2006/2007 nesting season, nests laid in
November hatched in 61.8 ± 4.2 days,
and nests laid in February hatched in
55.8 ± 3.4 days (n = 171 nests;
Steckenreuter et al. 2010). Assuming
that hatchlings were male at
temperatures less than 29.2 °C and
female at temperatures greater than 30.5
°C, Steckenreuter et al. (2010) estimated
that only 7.7 percent of the hatchlings
were female, indicating a highly maleskewed sex ratio. However, given the
Pilcher (2010) results, sex ratios are
likely variable over time and space.
Climatic change may also alter rainfall
levels, which may cool beaches and
offset increases in sand temperature. At
Wermon, the sand is black, yet beach
temperatures are lower, perhaps because
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peak nesting coincides with the
monsoon season (Tapilatu and Tiwari
2007). Sand temperatures fluctuate
between 28.6 and 34.9 °C at JamursbaMedi and between 27.0 and 32.7 °C at
Wermon (Tapilatu and Tiwari 2007).
Hatching success of nests undisturbed
by feral pig predation was significantly
lower in Jamursba-Medi (25.5 percent)
than Wermon (47.1 percent). Although
there was significant variation between
beaches, Tapilatu and Tiwari (2007)
concluded that high sand temperatures
may exceed the thermal tolerance of
leatherback embryos, resulting in high
embryo mortality and low hatching
success at Jamursba-Medi. Further,
Tapilatu and Tiwari (2007) concluded
that high average sand temperatures
may suggest a female-biased population
at Jamursba-Medi. However, the mean
incubation period of 61.5 ± 4.7 days
(Tapilatu and Tiwari 2007) was similar
to the length of incubation recorded in
Papua New Guinea during the cooler
November period, which Steckenreuter
et al. (2010) suggested produced a malebiased sex ratio.
Tapilatu et al. (2013b) found that the
daily average sand temperatures during
the boreal summer (from 2005 to 2012)
ranged from 26.5 to 34.9 °C, suggesting
the production of female-biased sex
ratios and potentially lower hatching
success. Further, histological
examination of dead hatchlings from
both summer and winter nesting
seasons from 2009 to 2019 produced a
female-biased sex ratio, which is
consistent with the relatively warm
thermal profiles of the nesting beaches
(Tapilatu et al. 2013b). Additional
impacts of climate change include
increased sea level rise and storm
frequency, resulting in greater nest
inundation and beach erosion. As sea
level rises, King Tides are likely to have
a greater effect on nests. Climate change
may also affect prey availability. Saba et
al. (2007, 2012) identified a correlation
between the reproductive frequency of
the East Pacific DPS and ENSO events.
Because the West DPS also forages in
the East Pacific Ocean, it too may be
exposed to variability in productivity.
The threat of climate change is likely
to modify the nesting and foraging
conditions for turtles of the DPS.
Impacts are likely to affect productivity.
Negative impacts and low hatching
success due to high beach temperatures
and coastal erosion have already been
documented and are likely to become
worse, and thus we conclude that
climate change is a threat to the West
Pacific DPS.
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Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply to
turtles of the West Pacific DPS (for a
description of each effort, please see the
section on conservation efforts for the
overall species): Convention on the
Conservation of Migratory Species of
Wild Animals, Convention on Biological
Diversity, Convention on International
Trade in Endangered Species of Wild
Fauna and Flora, Convention for the
Protection of the Marine Environment
and Coastal Area of the South-East
Pacific (Lima Convention), Convention
for the Conservation and Management of
Highly Migratory Fish Stocks in the
Western and Central Pacific Ocean
(WCPF Convention), Convention for the
Protection of the Natural Resources and
Environment of the South Pacific
Region, Convention Concerning the
Protection of the World Cultural and
Natural Heritage (World Heritage
Convention), Eastern Pacific
Leatherback Network, Eastern Tropical
Pacific Marine Corridor Initiative, FAO
Technical Consultation on Sea TurtleFishery Interactions, IAC, MARPOL,
IUCN, The Memorandum of
Understanding of a Tri-National
Partnership between the Government of
the Republic of Indonesia, the
Independent State of Papua New Guinea
and the Government of Solomon
Islands, Ramsar Convention on
Wetlands, RFMOs, Secretariat of the
Pacific Regional Environment
Programme, UNCLOS, and UN
Resolution 44/225 on Large-Scale
Pelagic Driftnet Fishing. Although
numerous conservation efforts apply to
the turtles of this DPS, they do not
adequately reduce its risk of this DPS,
they do not adequately reduce its risk of
extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the West Pacific DPS is at high risk of
extinction. The DPS exhibits a total
index of nesting female abundance of
1,277 females at two currently
monitored beaches over the most recent
remigration interval. These beaches may
represent 75 percent of total DPS
nesting activity. This abundance makes
the DPS vulnerable to stochastic or
catastrophic events that increase its
extinction risk. This DPS exhibits low
hatching success and decreasing nest
and population trends due to past and
current threats, which are likely to
further lower abundance and increase
the risk of extinction. The DPS exhibits
genetic diversity and metapopulation
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48399
structure, with nesting aggregations
distributed throughout four nations.
Nesting occurs during two seasons
(winter and summer), with year-round
nesting at some locations and uses
multiple foraging areas, throughout the
Pacific Ocean. Thus, the DPS has some
resilience to stochastic events and
environmental perturbations at nesting
beaches and foraging areas. However, its
abundance and declining trends place
the DPS at risk of extinction as a result
of past threats.
Current threats also contribute to the
risk of extinction of this DPS. The
overutilization of turtles and eggs, as a
result of legal and illegal harvest, is the
primary threat to this DPS, reducing
abundance and productivity.
Abundance and productivity are further
reduced by fisheries bycatch. Juvenile
and adult turtles are taken by numerous,
international, coastal, and pelagic
fisheries throughout the extensive, panPacific foraging range of the DPS.
Predation (especially by dogs and pigs)
reduces productivity at high rates.
Erosion and inundation result in habitat
loss and modification that reduces
productivity and contributes to low
hatching success. Additional threats
include: Pollution, vessel strikes, and
natural disasters. Climate change is an
increasing threat that results in reduced
productivity. Though many regulatory
mechanisms exist, they do not
adequately reduce threats.
We conclude, consistent with the
team’s findings, that the West Pacific
DPS is at risk of extinction. Its nesting
female abundance makes the DPS highly
vulnerable to threats. The declining
nesting trend further contributes to its
risk of extinction. While the DPS has
spatial structure and diversity, the
resilience provided by those factors is
likely to be eroded by the reduced and
declining abundance. Past egg and turtle
harvest reduced the abundance and
productivity of this DPS and remains a
primary threat. Fisheries bycatch is also
a primary threat that reduces abundance
by removing mature and immature
individuals from the population.
Predation is also a major threat to
productivity. Though numerous
conservation efforts apply to this DPS,
they do not adequately reduce the risk
of extinction. We conclude that the
West Pacific DPS is in danger of
extinction throughout its range and
therefore meets the definition of an
endangered species. The threatened
species definition does not apply
because the DPS is currently in danger
of extinction (i.e., at present), rather
than on a trajectory to become so within
the foreseeable future.
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East Pacific DPS
The Team defined the East Pacific
DPS as leatherback turtles originating
from the East Pacific Ocean, north of 47°
S, south of 32.531° N, east of 117.124°
W, and west of the Americas. In the
south, the cold waters of the Antarctic
Circumpolar Current likely restrict the
nesting range of this DPS. We placed the
northern and western boundaries at the
border between the United States and
Mexico because this DPS forages
primarily in the East Pacific Ocean, off
the coasts of Central and South
America.
The range of the DPS (i.e., all
documented areas of occurrence) is
centered in the eastern Pacific Ocean
but may include distant waters for
foraging, as demonstrated by a turtle
satellite-tracked to waters off the Tonga
Trench and a turtle captured by the
Hawaii longline fishery, genetically
assigned to the population we refer to in
this finding as the East Pacific DPS (P.
Dutton, NMFS, pers. comm., 2018).
Records indicate that the DPS occurs in
the waters of the following nations:
Chile; Colombia; Costa Rica; Ecuador; El
Salvador; France (Clipperton Island);
Guatemala, Honduras; Mexico;
Nicaragua; Panama; Peru; and the
United States (Hawaiian Islands)
(Wallace et al. 2013).
Leatherback turtles of the East Pacific
DPS nest primarily on beaches in
Mexico, Costa Rica, and Nicaragua. In
Mexico, where the largest nesting
aggregations occur, nesting beaches are
found in 11 states, over 7,828 kilometers
as far north as Baja California Sur (Sarti
2002). The following beaches in Mexico
host approximately 40 to 50 percent of
total nesting for the nation: Mexiquillo
(Michoaca´n), Tierra Colorada
(Guerrero), and Cahuita´n, Chacahua,
and Barra de la Cruz (Oaxaca; Gaona
Pineda and Barraga´n Rocha 2016). In
Costa Rica, approximately 75 percent of
nesting occurs within the Parque
Nacional Marino Las Baulas
(Guanacaste Province) at three nesting
beaches: Playa Ventanas; Playa Grande;
and Playa Langosta (based on recent
abundance estimates from 2011–2015;
Santidria´n Tomillo et al. 2017). In
Nicaragua, small numbers of leatherback
turtles nest on Playa Salamina-Costa
Grande and Veracruz de Acayo
(Chacocente Wildlife Refuge) (FFI 2018).
Rare nesting events have been
documented in Guatemala (n = 6), El
Salvador (n = 4), and Panama (n = 4),
with none in Honduras (Sarti et al.
1999).
Generally, the nesting season starts in
October and ends in March (Santidria´n
Tomillo et al. 2007; Eckert et al. 2012).
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Nesting is generally bound between 10°
N and 20° N, falling within the
northeast corner of the Intertropical
Convergence Zone. The nesting beaches
share similarly warm temperatures,
moderate annual rainfall, and seasonal
dynamics (Saba et al. 2012). In general,
nesting beach habitat for leatherback
turtles is associated with deep water
and strong waves and oceanic currents,
but shallow water with mud banks are
also used by leatherback turtles.
Beaches with coarse-grained sand and
free of rocks, coral, or other abrasive
substrates also appear to be selected by
leatherback turtles (reviewed by Eckert
et al. 2012).
Foraging areas of the East Pacific DPS
include coastal and pelagic waters of the
southeastern Pacific Ocean. Leatherback
turtles are widely dispersed on the high
seas throughout the eastern Pacific
Ocean (Shillinger et al. 2008). They also
forage in coastal areas off the coast of
Peru and Chile (Alfaro-Shigueto et al.
2007; Eckert 1997; Donoso and Dutton
2010). Using satellite telemetry,
Morreale et al. (1996) tracked the
movements of eight post-nesting females
and identified a persistent southbound
migration corridor from Las Baulas
National Park toward the Galapagos
Islands. Eckert (1997) found a similar
pattern, tracking seven post-nesting
females from Mexiquillo in a similar
direction; while three continued to the
same foraging habitat as the Costa Rican
nesting females, four shifted their
movements away from the South
American coast, when a strong El Nin˜o
caused a warm water anomaly.
Additional tracking of 46 post-nesting
females from Las Baulas National Park
over a 3-year period (2004/2005 to 2006/
2007) confirmed the persistent
migratory corridor (Shillinger et al.
2008). The turtles navigated the
equatorial current system, south to
around 5° S latitude and negotiated the
strong alternating eastward-westward
flows of the equatorial current,
swimming predominantly in a
southward direction and moving rapidly
through the productive equatorial
region. They then dispersed throughout
the South Pacific Gyre ecosystem,
which is characterized by low
phytoplanktonic biomass. The South
Pacific Gyre contains ample
mesoplankton forage base, as
demonstrated by tuna longline fisheries
effort in the eastern tropical Pacific
Ocean (Shillinger et al. 2008). Of the 46
turtles, only one leatherback moved into
coastal foraging areas, which had been
documented earlier by Eckert (1997).
During the course of the tracking
duration, this female occupied
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nearshore foraging habitats along the
coast of Central America, which
represents highly productive areas when
compared with oceanic areas.
Researchers have hypothesized that
high bycatch along the coastal areas of
Central and South America could have
extirpated a coastal migratory
phenotype in this population (Saba et
al. 2007). Recently, Harrison et al.
(2018) determined that post-nesting
females from Las Baulas National Park
spent 78.2 percent of their time on the
high seas, 17.8 percent of their time in
Costa Rica’s EEZ, and 3.7 percent of
their time around the Galapagos Islands.
In summary, preferred foraging areas
for the East Pacific DPS are
characterized by low sea surface
temperatures and high mesoscale
variability. Post-nesting females migrate
relatively quickly through areas that
contain the strong equatorial currents as
well as high chlorophyll-a
concentrations, likely because of the
strong currents. While swimming speed
was significantly higher in areas of high
chlorophyll levels, the association
between these two variables was weak
(Shillinger et al. 2008). Once past this
area, they appear to forage in the
southern part of their range in the South
Pacific Subtropical Convergence, where
there is a sharp gradient in primary
production. In this area, Ekman
upwelling may accelerate the transport
of nutrients and consequently increase
prey availability. Seasonally,
leatherback turtles from the East Pacific
DPS foraged at higher southerly
latitudes during the austral summer
(November to February), which may
reflect seasonal patterns in prey
abundance during higher latitudes
(Bailey et al. 2012).
Abundance
The total index of nesting female
abundance for the East Pacific DPS is
755 females. We based this total index
on 13 nesting aggregations in: Mexico
(Mexican Commission for Natural
Protected Areas; L. Sarti, CONANP,
pers. comm. 2018); Costa Rica
(Santidria´n Tomillo et al. 2017;
Leatherback Trust 2018); and Nicaragua
(FFI 2018). Our total index does not
include several unquantified nesting
aggregations in Mexico, Costa Rica, and
Nicaragua. To calculate the index of
nesting female abundance for nesting
beaches in Mexico (i.e., 572 females), we
added the total number of nesting
females between the 2013/2014 and
2016/2017 nesting seasons (i.e., a 4-year
remigration interval; L. Sarti, CONANP,
pers. comm., 2018) at each beach. We
performed a similar calculation for
Costa Rica (n = 165 females). To
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calculate the index of nesting female
abundance in Nicaragua (i.e., 20
females), we divided the total number of
nests between the 2014/2015 and 2017/
2018 nesting seasons (i.e., a 4-year
remigration interval; Santradia´n Tomillo
et al. 2007) by the clutch frequency (7.2
clutches/season; Santradia´n Tomillo et
al. 2007).
This number represents an index of
nesting females for this DPS because it
only includes available data from
recently and consistently monitored
nesting beaches. While rare or sporadic
nesting may occur on other beaches,
consistent and standardized monitoring
only occurs at these beaches, which are
for the most part protected.
Our total index of nesting female
abundance is similar to published
abundance estimates for this DPS. The
IUCN Red List assessment estimated the
total number of mature individuals
(males and females) at 633 turtles, based
first on dividing the average annual
number of nests (n = 926) by the
estimated clutch frequency (n = 7.2,
Reina et al. 2002) to obtain an average
annual number of nesting females. This
value was then multiplied by the
average remigration interval (n = 3.7
years, Reina et al. 2002; Santidria´n
Tomillo et al. 2007) to obtain a total
number of adult females that included
nesting as well as non-nesting turtles. In
order to account for adult males, the
authors assumed that the sex ratio of
hatchlings produced on nesting beaches
in the East Pacific (approximately 75
percent female, or 3:1 female:male ratio)
reflected the natural adult sex ratio
(Wallace et al. 2013). A more recent
analysis of primary sex ratios that
included multiple years of data and
considered hatching success (i.e., lower
in hot nests) estimated primary sex
ratios at Playa Grande, Costa Rica as
approximately 85 percent female
(Santidria´n Tomillo et al. 2014). In
Mexico, the female to male ratio is
closer to 1.1:1 (A. Barragan, Kutzari,
pers. comm., 2019).
In Mexico, the beaches included in
our total index represent approximately
70 to 75 percent of total nesting in that
nation (Gaona Pineda and Barragan
Rocha 2016). However, our total index
does not include nesting females from
Agua Blanca (40 km in Baja California);
Playa Ventura (6 km), Playa San
Valentı´n (21 km), Piedra de
Tlacoyunque (44 km in Guerrero), and
La Tuza (16 km in Oaxaca) (Sarti et al.
2007). These beaches are not regularly
monitored for nesting, which is thought
to be rare or of low abundance (L. Sarti,
CONANP, pers. comm., 2018).
In Costa Rica, 75 percent of nesting
occurred at Las Baulas National Park
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(summarized in Santidria´n Tomillo et
al. 2017), although the recent nesting at
other beaches may lower this
percentage. These beaches include:
Naranjo, Cabuyal, Nombre de Jesu´s,
Ostional, and Caletas. The longest data
set was provided for Naranjo, which has
been intermittently covered from 1971
to 2015. Limited nesting has been
documented at Playa Coyote and at
Playa Caletas, which is a high energy
eight kilometer beach located on the
Nicoya Peninsula (Squires 1999). Given
the lack of nesting events for Caletas in
recent years, it may no longer host
leatherback nesting, despite the fact that
the Playa Caletas/Ario National Wildlife
Refuge was created in 2004 to protect
leatherback turtles (Gaos et al. 2008).
In Nicaragua, leatherback turtles nest
at three beaches. Salamina Costa Grande
and Veracruz de Acayo (in the Rio
Escalante Chacocente Wildlife Refuge)
host the most nesting and have been
subject to the most consistent
monitoring. Small numbers of females
also nest at Juan Venado National
Reserve, which is not consistently
monitored (V. Gadea, FFI, personal
communication, 2018).
Nesting is rare in other nations (Sarti
et al. 1999). Nesting is very uncommon
in Ecuador with one record of a female
attempting to nest (according to local
reports) in Atacames, a province of
Esmeraldas (Salas 1981). Sarti et al.
(1999) reported six nests at Playa
Puntilla, El Salvador, but overall nesting
is low and/or unknown throughout the
nation. In Guatemala, nesting is rare,
with reports by Sarti et al. (1999)
recording only eight nests during an
entire season, and more recently, zero to
six nests per year along the Pacific coast
of Guatemala (Muccio and Flores 2015).
Past nesting sites included Hawai beach,
La Candelaria, Taxico, Santa Rosa, and
the zone adjacent to the border with El
Salvador, as reported by Chaco´nChaverri (2004). Although nesting has
been documented at Barqueta National
Refuge, little is known about nesting in
Panama (Chaco´n-Chaverri 2004).
Our total index of nesting female
abundance (755 females) places the DPS
at risk for environmental variation,
genetic complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). These
processes, working alone or in concert,
place small populations at a greater
extinction risk than large populations,
which are better able to absorb losses in
individuals. Due to its small size, the
DPS has relatively little capacity to
buffer such losses. Historical abundance
estimates were much greater (e.g.,
75,000 leatherback nesting females
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48401
estimated in Pacific Mexico from a 1980
aerial survey ((Pritchard 1982).
However, this estimate was derived
from a brief aerial survey and may have
been an overestimate (Pritchard 1996)),
indicating that this population at one
time had the capacity for a much larger
nesting population. Therefore, the
current nesting female abundance is
likely an indicator of past and current
threats, and given the intrinsic problems
of small population size, elevates the
extinction risk of this DPS.
Productivity
The East Pacific DPS exhibits a
decreasing nest trend since monitoring
began, with a 97.4 percent decline since
the 1980s or 1990s, depending on the
nesting beach (Wallace et al. 2013).
Despite intense conservation efforts, the
decline in nesting had not been reversed
as of 2011 (Benson et al. 2015). We
found a declining nest trend at some of
the remaining, small nesting
aggregations. Abundance at Las Baulas,
Costa Rica (previously the single largest
nesting aggregation) at its peak was
seven times the current abundance at
Playa Barra de la Cruz/Playa Grande,
Mexico (currently the largest nesting
aggregation). From 1988/1989 to 2015/
2016, the number of nesting females at
Las Baulas declined ¥15.5 percent
annually (sd = 3.8 percent; 95 percent
CI = ¥23.1 to ¥7.8 percent; f = 0.998;
mean annual nests = 315).
In recent decades (after a historical
decline), nest counts have increased at
some beaches in Mexico. The Playa
Tierra Colorada nest trend has increased
by 0.6 percent annually (sd = 8.9
percent; 95 percent CI = ¥17.1 to 18.9
percent; f = 0.536; mean annual nests =
153) between the 1996/1997 and 2016/
2017 nesting seasons. Over the same
time period, nesting at Playa Barra de la
Cruz/Playa Grande increased by 9.5
percent annually (sd = 8.0 percent; 95
percent CI = ¥6.5 to 25.8 percent; f =
0.918; mean annual nests = 122). In
contrast, nest counts at Cahuita´n
decreased from 1997/1998 through
2016/2017, with a median trend of ¥4.3
percent annually (sd = 9.7 percent; 95
percent CI = ¥22.1 to 17.6 percent; f =
0.716; mean annual nests = 123).
We lack adequate data on nesting in
Nicaragua to estimate trends.
Our trend analysis yields similar
results to other published findings. The
IUCN Red List assessment concluded
that this subpopulation is decreasing
and has declined by ¥97.4 percent over
the past three generations (Wallace et al.
2013). The number of nests at Mexico
nesting beaches has declined
precipitously in recent decades (Benson
et al. 2013). Historically, Mexico hosted
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the largest leatherback turtle nesting
aggregation in the world, with 75,000
nesting females estimated during an
aerial survey in 1980 ((Pritchard 1982).
However, this estimate was derived
from a brief aerial survey and may have
been an overestimate (Pritchard 1996)).
Prior to that aerial survey, Marquez et
al. (1981) reported that the nesting
beach of San Juan Chacahua (Oaxaca)
was the most important nesting site in
Mexico, with approximately 2,000
females nesting each season.
Researchers also identified Tierra
Colorada and Mexiquillo as important
nesting sites, with approximately 3,000
to 5,000 nests per season. Monitoring of
the nesting assemblage at Mexiquillo
has been continuous since 1982. During
the mid-1980s, more than 5,000 nests
per season were documented along 4 km
of this nesting beach. By 1993, less than
100 nests were counted along the entire
18 km beach (Sarti 2002). According to
Sarti et al. (1996), nesting declined at
this location at an annual rate of over 22
percent from 1984 to 1995. Researchers
from the National University of Mexico
recorded 3,000 to 5,000 nests annually
from 1982 to 1989 at primary nesting
beaches, with sharp declines observed
in 1993 to 1994 at the nesting sites at
Mexiquillo, Tierra Colorada, Chacahua
and Barra de la Cruz. These early
reports were generally snapshots (e.g.,
local unpublished data) of leatherback
nesting activity in Mexico, until 1995,
when a more coordinated conservation
effort took shape in the form of
complete nesting surveys for the entire
Pacific coast of Mexico (Eckert 1997). In
1995, ‘‘Proyecto Laud’’ (Leatherback
Project) was formed to estimate the
population size using comprehensive
surveys. In 1995 and 1996, Proyecto
Laud estimated approximately 1,100
females nesting throughout Mexico; the
next two seasons, they estimated
between 236 and 250 nesting females,
and declines continued. Currently,
based on data from 2014 through 2018
(preliminary) between 100 and 250
females nest at all the protected beaches
in Mexico.
In Costa Rica, the number of nesting
females per season declined from 1,367
females in 1988 to 117 females in 1998
(Spotila 2000). While there were
increases in the number of nesting
females during the 1999/2000 season
(224 females) and 2000/2001 season
(397 females), the population has shown
a steady decline, with less than 30
nesting females in recent years (i.e.,
through 2016; The Leatherback Trust
2018).
In Nicaragua, 108 leatherback turtles
nested on Playa Chacocente from
October to December, 1980; in January
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1981, 100 turtles nested in a single night
on Playa El Mogote (Arauz 2002). An
aerial survey of Playa El Mogote during
the 1998/1999 nesting season revealed a
nesting density of 0.72 turtles per
kilometer (Sarti et al. 1999 in Arauz
2002). During the 2000/2001 nesting
season, community members near Playa
El Mogote reported that 210 leatherback
nests had been deposited. That number
decreased to 29 nests during the 2001/
2002 nesting season (Arauz 2002). At
Playa Veracruz 48 nesting females were
identified between 2002 and 2010
(Urteaga et al. 2012). Between 2002 and
2014, Salazar et al. (2019) recorded 340
nests, indicating a downward trend.
Considering the best available data,
nesting has declined in Nicaragua.
Nesting females of the East Pacific
DPS are generally smaller and produce
fewer eggs per clutch than turtles from
other leatherback populations (Sarti et
al. 2007; Piedra et al. 2007; Santidria´n
Tomillo et al. 2007). For example in
Mexico, nesting females have a mean
size of 144 cm CCL and 62 eggs per
clutch; the average total fecundity per
females was estimated to be 341 eggs
per season, with a maximum of 744 eggs
deposited in a season (Sarti et al. 2007).
The low productivity parameters,
drastic reductions in overall nesting
female abundance, and current declines
in nesting place the DPS at risk of
extinction, especially given the limited
nesting female abundance.
Spatial Distribution
The DPS is characterized by
somewhat continuous and low density
nesting across long stretches of beaches
along the coast of Mexico and Central
America. Santidria´n Tomillo et al.
(2017) found a contraction of the Costa
Rica’s overall nesting distribution since
the 1990s.
The best available genetic data
indicate a high degree of connectivity
among nesting aggregations. Dutton et
al. (1999) did not find any genetic
differentiation between nesting
populations in Mexico (Playa
Mexiquillo) and Costa Rica (Playa
Grande) based on analysis of mtDNA
control region sequences. Additional
analyses of mtDNA sequences and
nuclear DNA (microsatellites) from
three index nesting beaches in Mexico
also failed to find genetic differentiation
(Barragan and Dutton 2000; Dutton et al.
unpublished).
Based on monitoring of tagged nesting
females, researchers documented female
interchange between nesting beaches
within Mexico and within Costa Rica.
However, only one interchange has been
documented between Mexico and Costa
Rica (Sarti et al. 2007). Interchange
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between nesting beaches may occur
during or between nesting seasons and
may depend on the distance between
nesting sites, which can be fairly large,
especially in Mexico. For example, the
distance between Tierra Colorada and
Cahuita´n is 25 kilometers, and up to
18.7 percent of nesting females visit
both beaches within a season (average of
nine percent). Mexiquillo is located
approximately 475 kilometers from the
closest other nesting beach (Tierra
Colorada), and researchers found no
interchange of females within seasons.
However, a few females were found to
nest in either Mexiquillo and/or Tierra
Colorado between seasons (Sarti et al.
2007).
In Costa Rica, nesting females move
among the three nesting beaches of Las
Baulas National Park, within and
between seasons, particularly between
Playa Grande and Playa Langosta,
although researchers study both Playa
Grande and Playa Ventanas in
combination. According to data
gathered over 10 years of research (mid
1990s through the mid-2000s), an
average of 71 percent of females nested
only on Playa Grande, 10 percent nested
only on Playa Langosta, and 18 percent
nested on both beaches in a given
season. In other seasons, females have
been shown to shift and nest primarily
on a different beach. Within two
seasons, 82 percent of nesting females at
Playa Langosta also nested at Playa
Grande and 100 percent of nesting
females at Playa Langosta within three
seasons occasionally also nested at
Playa Grande (Santidria´n Tomillo et al.
2007). At the less abundant nesting
beaches in Costa Rica, the exchange rate
between females ranged between 7 and
28 percent. For example, at Ostional, 12
out of the 43 identified females were
observed at least once at other sites (28
percent), while at Naranjo, 4 out of 21
identified females were also observed at
other beaches (19 percent). At Cabuyal,
2 out of 15 turtles were observed at
other beaches (13 percent), while 1 out
of 15 females at Caletas were observed
elsewhere (7 percent) (Santidria´n
Tomillo et al. 2017).
The foraging range of the DPS extends
into coastal and pelagic waters of the
southeastern Pacific Ocean. Individuals
forage in the Pacific Gyre ecosystem and
along the coasts of Peru and Chile, with
variation resulting from the location of
upwelling and ENSO effects.
Researchers have hypothesized that
high bycatch along the coastal foraging
phenotype in this population (Saba et
al. 2007). Recently, Harrison et al.
(2018) determined that post-nesting
females from Las Baulas National Park
spent 78.2 percent of their time on the
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high seas, 17.8 percent of their time in
Costa Rica’s EEZ, and 3.7 percent of
their time around the Galapagos Islands.
Multiple nesting and foraging
distributions likely help to buffer the
DPS against local catastrophes or
environmental changes that would
otherwise modify nesting habitat or
limit prey availability. Nesting
aggregations are largely connected.
However, there is less exchange among
distant nesting beaches. Foraging turtles
are vulnerable to perturbations in ocean
conditions due to climate change,
ENSO, and the Pacific Decadal
Oscillation.
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Diversity
The East Pacific DPS exhibits genetic
diversity, as demonstrated by moderate
to high mtDNA haplotypic diversity (h
= 0.66–0.71; Dutton et al. 1999). Such
diversity likely provides the DPS with
some capacity for adapting to long-term
environmental changes, such as cyclic
or directional changes in ocean
environments due to natural and human
causes (McElhany et al. 2000; NMFS
2017). Nesting habitat is mainly
restricted to mainland beaches along the
same coast. The DPS does not exhibit
temporal or seasonal nesting diversity,
with most nesting occurring between
October and March. This limits
resilience. For example, short-term
spatial and temporal changes in the
environment are likely to affect all
nesting females in a particular year. The
foraging strategies are somewhat
diverse, with turtles foraging in coastal
and oceanic waters. However, most
turtles forage in the East Pacific Ocean,
where they are similarly exposed to the
effects of climate change, ENSO, or the
Pacific Decadal Oscillation. Thus, the
DPS has limited resilience.
Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
The destruction or modification of
habitat is a threat at many nesting
beaches used by turtles of the East
Pacific DPS. Foraging habitat has also
been characterized as marginal,
particularly in the eastern tropical
Pacific Ocean (pelagic environment) due
to relatively low productivity. Coastal
habitat, which is normally associated
with high productivity, may have been
marginalized due to high levels of
interactions with coastal artisanal
fisheries.
Development threatens the DPS by
modifying the preferred beach habitat
for nesting. Sustained and substantial
development along the northern and
southern ends of the nesting beach at
Playa Grande in Las Baulas National
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Park, and in adjacent areas, has resulted
in the loss of nesting beach habitat in
addition to the removal of much of the
natural beach vegetation. As a result,
erosion has increased and led to other
environmental damages to sand that are
associated with human development,
including significant changes to
elevation, water content, particle size,
pH, salinity, organic content and
calcium carbonate content (Clune and
Paladino 2008). Within the past two
decades, beachfront development in the
town of Tamarindo (across Tamarindo
Bay from Playa Grande) has resulted in
the degradation of nesting beach habitat,
including: Pollution from artificial light,
solid and chemical wastes, beach
erosion, unsustainable water
consumption, and deforestation. Hotels
in this area have replaced a significant
leatherback nesting area at Playa
Tamarindo, which hosted significant
nesting in the 1970s and 1980s (Wallace
and Piedra 2012). Playa Langosta, which
is just across from Tamarindo, is
inundated with lights and noise from
the town (Wallace and Piedra 2012).
Currently, development has been
curtailed due mainly to water issues
(i.e., drought). Any additional
development would damage the current
hydrology. The Leatherback Trust, a
local nonprofit working at Las Baulas
National Park, has acquired some
properties to prevent development, but
property costs have increased over time.
At Las Baulas National Park, 10 percent
of nests were being inundated by tidal
flows. To mitigate this threat, nests at
risk of tidal inundation were relocated
to another site on the same beach or into
a hatchery. Hatchling production
slightly increased due to the
establishment of the hatchery, where
approximately two percent of hatchlings
were produced from 1998 to 2004
(Santidria´n Tomillo et al. 2007). We
conclude that coastal development in
Costa Rica is a threat to this DPS.
In Mexico, the extent of development
near nesting beaches is generally low,
given the remoteness of the beaches in
Baja California and on the mainland.
Reviewing the location of these nesting
beaches, we found very few roads or
development nearby. The main nesting
beaches remain somewhat isolated, with
very few roads or development adjacent
to the nesting beaches. Thus, there is
limited threat due to artificial lighting
and generally little to no beach driving
except perhaps that associated with
monitoring efforts (L. Sarti, CONANP,
pers. comm., 2018). In 2002, the
Commission for Natural Protected Areas
designated two of the index beaches
(Mexiquillo and Tierra Colorada) as
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48403
natural protected areas (turtle
sanctuaries), which helped protect
nesting habitat. Subsequently, in 2003,
three of the index beaches (Mexiquillo,
Tierra Colorada, and Cahuita´n) were
listed as Ramsar Sites, which are
wetland sites designated to be of
international importance under the
Ramsar Convention.
At Veracruz de Acayo beach in
Nicaragua, Salazar et al. (2019) note that
while conservation efforts has reduced
the threat of poaching, the
establishment of tourism-focused
coastal development that do not comply
with the existence of management plans
could threaten the nesting habitat.
While nesting beaches within this
DPS are generally remote and/or
protected due to monitoring and
existence of national parks and wildlife
refuges, nesting females, hatchlings, and
eggs at Las Baulas National Park (Costa
Rica) nesting beaches are exposed to the
modification of nesting habitat, as a
result of development. This threat
impacts the DPS by reducing nesting
and hatching success, thus lowering the
productivity of the DPS. We conclude
that habitat loss and modification is a
threat to the East Pacific DPS.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
The harvest of nesting females and
eggs was the primary cause of the
historical decline in abundance of the
East Pacific DPS. Since then, laws have
been passed to protect eggs and turtles.
However, poaching still occurs.
In Mexico, Sarti et al. (2007)
attributed the decline of nesting females
to the killing of adult females and
intensive egg harvest. Adult females
were historically killed at nesting
beaches and in open waters (Sarti et al.
1994; Sarti et al. 1998). Since 1990, the
harvest of turtles and eggs has been
prohibited by national legislation.
However, poaching pressure remains
high wherever beach patrols do not
occur (Santidria´n Tomillo et al. 2017).
For example, Mexiquillo produced
hatchlings every season in the 1980s.
However, even with efforts to protect
the nests in place, 60 to 70 percent of
the total number of clutches were
poached. Nichols (2003) notes that
leatherback turtles were once harvested
off Baja California, but their meat is now
considered inferior for human
consumption. At present, leatherback
turtles are not generally captured for
their meat or skin, but the poaching of
nesting females has been known to
occur on beaches such as Piedra de
Tlacoyunque, Guerrero (Sarti et al.
2000).
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Although poaching of turtles and eggs
has been consistently reduced over the
years, it still occurs at high levels.
Effective conservation and protection
depends on human presence at the
nesting beaches (Santidria´n Tomillo et
al. 2017). Without such protection,
poaching is likely to escalate. This may
have occurred at one of the primary
nesting beaches (Mexiquillo), where
monitoring and conservation has not
taken place in recent years due to safety
concerns (L. Sarti, CONANP, pers.
comm., 2018). Since the mid-1990s,
Proyecto Lau´d has been relocating
clutches (usually within 1–2 hours of
deposition) to protected fenced areas
and releasing hatchlings in different
areas of the beach. These efforts are
intended to protect the eggs from
poachers/predators and the hatchlings
from predators (Sarti et al. 2007).
In Costa Rica, the population decline
was predominantly caused by egg
harvest. Ninety percent of eggs were
collected on one of the major nesting
beaches, Playa Grande, a decade or more
prior to the reduction of nesting females
(Santidria´n Tomillo et al. 2007). In the
1950s, there were few nesting females at
Playa Grande (Wallace and Piedra
2012). In the late 1960s and early 1970s,
the number of nesting turtles increased
to more than 100 nesting females
nightly (Wallace and Piedra 2012). In
the early 1970s, newly constructed
roads provided access to people from
distant villages and cities, and egg
harvest increased to more than 90
percent by the late 1970s (Wallace and
Piedra 2012). Such high levels of egg
harvest persisted for nearly two decades
(Wallace and Saba 2009). Despite
protection of nesting beaches at Las
Baulas National Park, illegal poaching of
eggs still occurs, though rarely. The
black market for eggs remains strong;
local bars throughout Guanacaste and
elsewhere continue to offer shots of raw
sea turtle egg yolks accompanying beer
or liquor (Wallace and Piedra 2012).
In 1991, the Parque Nacional Marino
Las Baulas was created and
subsequently ratified by law in 1995.
The Park consists of three leatherback
nesting beaches: Playa Grande, Playa
Ventanas, and Playa Langosta. The
establishment of the park ensured
increased protection at all three nesting
beaches, greatly reducing egg poaching
in the area. Poaching of eggs was
reduced from 90 percent prior to 1990/
1991, to 50 percent in 1990/1991, 25
percent in 1991 through 1993, and near
0 percent in 1993/1994 (Santridia´n
Tomillo et al. 2007). To mitigate
poaching, nests are often relocated.
However, relocation may reduce
hatching success (reviewed in
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Herna´ndez et al. 2007; Eckert et al.
2012). In Playa Grande, Costa Rica,
fewer females were produced in
translocated nests; cooler nests due to a
lower number of metabolizing embryos
may have reduced hatchling success
(Sieg et al. 2011).
In Nicaragua, prior to protection in
the early 2000s, poachers took nearly
100 percent of the nests at the three
nesting beaches. Nesting beach
protection has occurred at Veracruz
since 2002, Juan Venado since 2004,
and Salamina since 2008. An average of
ten community team members (mostly
ex-poachers) monitor beaches
seasonally. From 2002 to 2010, up to
420 nests were recorded and an
estimated 94 were protected (Urteaga et
al. 2012). While Veracruz de Acayo and
Salamina are protected at 100 percent,
Isla Juan Venado is not permanently
monitored. Therefore, poaching is likely
to occur. Poaching occurs at high levels
at other beaches, such as Playa El
Mogote. During the 2001/2002 nesting
season, 23 of 29 nests were poached (79
percent), and the remaining six nests
were protected in a hatchery (Arauz
2002). Due to the high level of poaching
in this area, when possible, researchers
from Flora & Fauna International
relocated 98 nests between 2002 and
2004. However, these nests had a low
emergence rate (22 percent; Urteaga and
Chaco´n 2008).
Extensive and prolonged effects of
comprehensive egg harvest have
depleted the leatherback population in
Costa Rica and Mexico, with egg harvest
levels of nearly 90 percent for about two
decades (Sarti et al. 2007; Santidria´n
Tomillo et al. 2008; Wallace and Saba
2009). Currently, nesting females and
eggs of the East Pacific DPS are exposed
to poaching. Though efforts have
reduced the levels of poaching of both
eggs and nesting turtles, egg poaching
remains high and affects a large
proportion of the DPS. Poaching of
nesting females reduces both abundance
(through loss of nesting females) and
productivity (through loss of
reproductive potential). Such impacts
are high because they directly remove
the most productive individuals from
DPS, reducing current and/or future
reproductive potential. Egg harvest
reduces productivity only, but over a
long period of time, this also reduces
recruitment and thus abundance. Given
the high exposure and impacts, we
conclude that overutilization, as a result
of poaching, poses a major threat to the
DPS.
Disease or Predation
Little is known about diseases and
parasites in leatherback turtles, although
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fibropapillomatosis has been described
as a major epizootic disease in hard
shelled turtles. A fibropapilloma tumor
(in regression) was found on one nesting
female at Mexiquillo, Mexico in 1997
(Huerta et al. 2002). Various bacteria
have also been documented in
leatherback eggs. Soslau et al. (2011)
sampled eggs laid on a Costa Rican
beach to determine if bacteria were
contributing to the low hatching rate (50
percent). The bacteria identified (i.e.,
species of the Bacillus, Pseudomonas,
and Aeromonas genera) are known
pathogens to humans and may account
for developmental arrest of the turtle
embryo (Soslau et al. 2011).
Numerous predators prey on East
Pacific leatherback turtles throughout
their life stages. Eggs and hatchlings are
eaten by crabs, ants, birds, reptiles,
mammals, and fish (Eckert et al. 2012).
In Costa Rica, during the 1993/1994
nesting season, several nests were lost to
predation and infestation by maggots
(Schwandt et al. 1996). In the Nicoya
Peninsula, on the Pacific coast of Costa
Rica, Squires (1999) documented
evidence of potential nest predation by
dogs, coyote, and raccoon. Predation of
hatchlings by dogs and raccoons has
increased in Playa Grande due to an
increase in development in the area (P.
Santridia´n Tomillo, The Leatherback
Trust, pers. comm., 2019).
For adult turtles, principal predators
at sea include killer whales, crocodiles
(Pritchard 1981), and sharks, while
nesting females are taken by crocodiles
(Bedding and Lockhart 1989), tigers, and
jaguars (Pritchard 1971). Sarti et al.
(1994) observed a lone male killer whale
feeding on a single gravid female near
Michoaca´n, Mexico, apparently
consuming only certain parts of the
turtle and discarding others (e.g., female
reproductive organs). In summary, eggs,
hatchlings, and some adults are exposed
to predation. For this DPS, the primary
impact is to productivity (i.e., reduced
egg and hatching success). Predation on
nesting females, while rare, reduces
abundance and productivity. Nest
predation is mitigated through screening
of nests, relocation of nests to hatcheries
and releasing hatchlings in safer areas of
the beach, and protecting nesting
females from large predators such as
dogs and jaguars (Sarti et al. 2007); some
of these efforts are funded through the
MTCA. We conclude that predation is a
threat to the East Pacific DPS.
Inadequacy of Existing Regulatory
Mechanisms
Several international regulatory
mechanisms apply to turtles in this
DPS. The IAC, in particular, prohibits
the harvest of turtles and eggs. CITES
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limits all international trade of the
species. There are also international
efforts to reduce fisheries bycatch.
In 2015, at the 7th Conference of the
Parties, the IAC resolved to prioritize
conservation actions in their work
programs that would help ‘‘reverse the
critical situation of the leatherback sea
turtle in the Eastern Pacific.’’
Specifically, parties were urged to: (1)
Submit leatherback bycatch information
annually to the IAC Secretariat; (2)
improve leatherback turtle fishery
monitoring efforts through the use of onboard observers; (3) report annually on
the measures they have taken to reduce
leatherback bycatch in their fisheries;
(4) enhance leatherback nest monitoring
and protection to increase hatchling
survival and protect nesting beach
habitat; (5) foster safe handling and
release of incidentally bycaught
leatherback turtles in fisheries; and (6)
agree to a five-year strategic plan
containing key activities related to the
resolution (CIT–COP7–2015–R2). The
strategic plan was patterned after the
Regional Action Plan for Reversing the
Decline of the Eastern Pacific
Leatherback (https://
savepacificleatherbackturtles.org) and
included measures to reduce fisheries
bycatch of adult and subadult
leatherback turtles, the identification of
high risk areas with fisheries and
leatherback turtles, the identification
and protection of important areas for
leatherback turtle survival in different
life stages, the elimination of any
consumption and illegal use of
leatherback turtles, and nesting site
protection.
As mandated by the 1994 North
American Agreement for Environmental
Cooperation, the Commission for
Environmental Cooperation (CEC)
encourages Canada, the United States,
and Mexico to adopt a continental
approach to the conservation of flora
and fauna. In 2003, this mandate was
strengthened as the three North
American nations launched the
Strategic Plan for North American
Cooperation in the Conservation of
Biodiversity. The North American
Conservation Action Plan (NACAP)
initiative began as an effort promoted by
the three nations, through the CEC, to
facilitate the conservation of marine and
terrestrial species of common concern.
In 2005, the CEC supported the
development of a NACAP for Pacific
leatherback turtles by Canada, the
United States, and Mexico. Identified
actions in the plan addressed three main
objectives: (1) Protection and
management of nesting beaches and
females; (2) reducing mortalities from
bycatch throughout the Pacific Basin;
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and (3) waste management, control of
pollution, and disposal of debris at sea.
In 2015, the Eastern Pacific
Leatherback Network (also known as La
Red de la Tortuga Lau´d del Oce´ano
Pacifico (Red Lau´d OPO)
(www.savepacificleatherbacks.org)) was
formed to address the critical need for
regional coordination of East Pacific
leatherback conservation actions to
track conservation priorities and
progress at the population level. This
network has brought together
conservationists, researchers,
practitioners and government
representatives from 22 institutions
across nine East Pacific nations with
varying priorities, capacities and
historical experiences in leatherback
research and conservation to contribute
to shared activities, projects, and goals.
Through these efforts, Red Lau´d OPO
now has mutually-agreed upon
mechanisms for sharing information and
data, as well as standardized protocols
for nesting beach monitoring and
bycatch assessments/fishing practices.
The Convention for the Protection of
Natural Resources and Environment of
the South Pacific, also known as the
Noumea Convention, has been in force
since 1990 and includes 26 Parties (as
of 2013). The purpose of the Convention
is to protect the marine environment
and coastal zones of the South-East
Pacific, and beyond that area, the high
seas up to a distance within which
pollution of the high seas may affect
that area.
In 2015, the IATTC passed a
resolution that requires large longline
vessels fishing in the eastern tropical
Pacific Ocean to carry observers.
Cooperating parties that have
documented interactions with sea
turtles in their longline fleet are
required to maintain at least five percent
observer coverage and provide an
annual report to the IATTC.
Unfortunately, the forms used by
observers to report incidents are not
standardized, so in some cases, the
reports did not include species
identification, condition of the released
turtles, and location of the interactions,
and the five percent minimum coverage
is often not met. Nations without
reported bycatch of sea turtles simply
provided a statement to that effect. In
the few reports we reviewed,
leatherback turtles comprised some of
the bycatch in the eastern tropical
Pacific Ocean, but there were few
details on the events (C. Fahy, NMFS,
pers. comm., 2018). In 2007, the IATTC
passed a resolution requiring nations to
conduct research on sea turtle bycatch
reduction measures in their longline
fleets (e.g., use of circle hooks and fish
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bait). Despite results in both the Atlantic
and Pacific longline fleets showing that
use of circle hooks/fish bait significantly
reduced leatherback bycatch rates
(Swimmer et al. 2017), nations are not
required to use this hook/bait
combination. In 2017, at an IATTC sea
turtle bycatch reduction workshop, the
United States presented findings on
longline bycatch reduction and
proposed a stronger resolution that
would require use of this methodology.
However, some nations resisted, and the
resolution did not move forward for
consideration at the annual IATTC
meeting.
Throughout the world, illegal,
unreported, and unregulated (IUU)
fishing leads to underestimates of
bycatch. In Mexico, there is a lack of
effective fisheries governance, resulting
in highly uncertain fishery statistics. For
example, from 1950 to 2010, total
fisheries catch, including estimated IUU
catch and discarded bycatch, was nearly
twice as high as the official statistics
(Cisneros-Montemayor et al. 2013).
Thus, the bycatch threat of commercial
fisheries in Mexico may be higher than
currently estimated.
In addition, several international
treaties and/or regulatory mechanisms
protect East Pacific leatherback turtles.
While no single law or treaty can be 100
percent effective at minimizing
anthropogenic impacts to sea turtles in
these areas, there are several
international conservation agreements
and laws in the region that, when taken
together, provide a framework within
which sea turtle conservation advances
can be made (Frazier 2012). In addition
to protection provided by local marine
reserves throughout the region, sea
turtles may benefit from the following
broader regional effort: (1) The Eastern
Tropical Pacific (ETP) Marine Corridor
(CMAR) Initiative supported by the
governments of Costa Rica, Panama,
Colombia, and Ecuador, which is a
voluntary agreement to work towards
sustainable use and conservation of
marine resources in these nations’
waters; (2) the ETP Seascape Program
managed by Conservation International
that supports cooperative marine
management in the ETP, including
implementation of the CMAR; (3) the
IATTC and its bycatch reduction efforts
through resolutions on sea turtles,
observer coverage, etc.; (4) the IAC,
which is designed to lessen impacts on
sea turtles from fisheries and other
human impacts; and (5) the Permanent
Commission of the South Pacific (Lima
Convention), which has developed an
Action Plan for Sea Turtles in the
Southeast Pacific.
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Most nations within the range of the
East Pacific DPS have laws prohibiting
the harvest of turtles and eggs. This
applies to nesting turtles and those
captured at sea. National laws in Mexico
(1990 Presidential Decree), Costa Rica
(2002 Presidential Decree N°8325: The
Law of Protection, Conservation, and
Recuperation of Marine Turtles), and
Nicaragua (Law No. 651 and Ministrial
Resolution No. 043–2005) protect
nesting females and eggs and nesting
beaches. However, poaching remains a
major threat. Although laws prohibit the
harvest of turtles in Peru, fishermen
consume leatherback turtles bycaught in
small-scale fisheries (Alfaro-Shigueto et
al. 2011), indicating inadequate
enforcement of existing laws. In other
nations where leatherback turtles of this
DPS are bycaught, the turtles are
released and not retained (e.g., Chile;
Donoso and Dutton 2010).
Several protected areas have been
established throughout the range of the
DPS. Most of the nesting beaches in
Mexico and Costa Rica are protected
from egg and turtle poaching, with
effective monitoring to ensure low
levels of poaching. Poaching likely
continues at unprotected and remote
beaches, and at those that contain an
extensive coastline that is difficult to
monitor and protect. Protected nesting
beaches in Mexico include: Mexiquillo
(until 2013); Playa de Tierra Colorada,
Playa Cahuita´n, Playa San Juan, Bahia
de Chacahua, and Playa Barra de la
Cruz. Protected nesting beaches in Costa
Rica include: Las Baulas National Park
(Playa Grande, Playa Langosta, and
Playa Ventanas), Naranjo (National
Park), Cabuyal (under no official
management category), Nombre De Jesu´s
(under no official management
category), Ostional (wildlife refuge), and
Caletas (wildlife refuge). Protected
nesting beaches in Nicaragua include:
Salamina-Costa Grande, Veracruz de
Acayo (Chacocente Wildlife Refuge).
Marine protected areas also exist. The
waters of the Las Baulas National Park,
which represents a hotspot for internesting females and breeding males, are
protected out to 22.2 km as a no-take
zone for all fishing activity. However,
satellite telemetry data for nesting
females at these beaches over three
seasons revealed that the turtles move
well outside these boundaries during
their inter-nesting period, which makes
them vulnerable to fisheries outside the
park (Shillinger et al. 2010). Data from
44 females that were tagged off Las
Baulas National Park revealed a high
use habitat within 6 nm from the
nesting beaches, but overall revealed a
generally large range, covering over
33,000 km2, from the Nicoya Peninsula,
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east into the Gulf of Nicoya in Costa
Rica, and north to coastal habitats
within 30 kilometers offshore from
southern Nicaragua. The marine areas
adjacent to this protected boundary are
not managed under any type of status
(Shillinger et al. 2010). Fisheries within
Costa Rica and Nicaragua’s EEZ include
trawl, gillnet and longline that continue
to operate.
In summary, numerous regulatory
mechanisms exist to protect leatherback
turtles, eggs, and nesting habitat
throughout the range of this DPS.
Although the regulatory mechanisms
provide some protection to the species,
many do not adequately reduce the
threat that they were designed to
address, generally as a result of limited
implementation or enforcement. As a
result, bycatch, incomplete nesting
habitat protection, and poaching remain
threats to the DPS. We conclude that the
inadequacy of existing regulatory
mechanisms is a threat to the East
Pacific DPS.
Fisheries Bycatch
Bycatch in commercial and
recreational fisheries, both on the high
seas and off the coasts, is the primary
threat to the East Pacific DPS. This
threat affects the DPS by reducing the
abundance of all life stages of the DPS
(with the likely exception of hatchlings).
Integrating catch data from over 40
nations and bycatch data from 13
international observer programs,
Lewison et al. (2004) estimated the
numbers of leatherback turtles taken
globally by pelagic longliners to be more
than 50,000 leatherback turtles in just
one year (2000). With over half of the
total fishing effort (targeting tuna and
swordfish) occurring in the Pacific
Ocean, an estimated 20,000 to 40,000
leatherback turtles interacted with
longline fishing during the year studied.
Fishing effort was highest in the central
South Pacific Ocean (south of Hawaii),
which overlaps with the foraging range
of this DPS. Because observers are in
place on only a fraction of longline
vessels in the eastern tropical Pacific
Ocean, and a requirement came into
effect only recently through an IATTC
resolution, these estimates are likely a
minimum. More recently, Molony
(2005) and Beverly and Chapman (2007)
estimated sea turtle longline bycatch to
be approximately 20 percent of that
estimated by Lewison et al. (2004), or
approximately 200 to 640 leatherback
turtles annually. Where tuna species are
targeted, bycatch of turtles in the deepset longline gear often results in
mortality due to drowning. Additional
studies indicate the high impact of
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industrial longline fleets on leatherback
turtles (e.g., Spotila et al. 1996, 2000).
In their global study of sea turtle
bycatch, where available, Wallace et al.
(2013) found that longline bycatch had
a low impact, but that net bycatch had
a high impact on the East Pacific RMU.
The impact of local artisanal fleets
(using gillnets and longlines) that fish
closer to shore is less documented.
In Mexico, leatherback turtles wash to
shore entangled in longlines and
driftnet, indicating interaction and
mortality (Sarti et al. 2007). OrtizAlvarez et al. (2019) conducted a
bycatch survey across 48 different ports
(933 fishers) in Mexico, Nicaragua and
Costa Rica between October 2016 and
July 2017 in an effort to improve the
understanding of leatherback bycatch in
artisanal fisheries, particularly where
data are lacking. The surveys
represented on average over 30 percent
of the fishing fleet per port for both
Nicaragua and Costa Rica and 6 percent
per port for Mexico. In Mexico, where
gillnets were the most frequently
reported gear, fishers (n = 709) reported
an estimated bycatch of 300 leatherback
turtles in the previous year, with 65
percent in ‘‘good condition;’’ 76 percent
of fishers released turtles alive (three
percent consumed or sold the turtles).
Estimated average bycatch rates per
vessel were 1.0 for Costa Rica and
Nicaragua and 2.3 for Mexico. In Costa
Rica, leatherback turtles were primarily
caught in longlines and released alive;
75 percent of the Costa Rican fishermen
reported that bycaught leatherback
turtles were in ‘‘good condition.’’ In
Nicaragua, where gillnets were the most
frequently reported gear, 18 percent of
fishers reported that leatherback turtles
were in ‘‘good condition;’’ 76 percent of
fishers released turtles alive (six percent
consumed or sold the turtles; OrtizAlvarez et al. (2019).
Recent surveys of 765 Ecuadorian,
Peruvian, and Chilean fishermen (at 43
ports, representing 28 to 63 percent of
ports) reported the following
leatherback interaction rates (as a
percentage of total interactions with sea
turtles): 2.81 percent of 40,480
interactions (32.5 percent mortality) in
Ecuador, 14.87 of 5,828 interactions
(50.8 percent mortality) in Peru, and
27.83 percent of 170 interactions (3.2
percent mortality) in Chile (AlfaroShigueto et al. 2018). Mortality rates
reported for all sea turtles were 3.2
percent in Chile, 32.5 percent in
Ecuador, and 50.8 percent in Peru
(Alfaro-Shigueto et al 2018).
The swordfish gillnet fisheries in Peru
and Chile may have contributed to the
decline of the DPS. The decline in the
nesting population at Mexiquillo
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occurred at the same time that effort
doubled in the Chilean driftnet fishery
(Eckert 1997). Using data collected from
Frazier and Montero (1990) regarding
leatherback takes in a swordfish gillnet
fishery from one port in Chile (San
Antonio), and extrapolating to other
ports in Chile and Peru, with an
increased level of effort observed
through the mid-1990s, Eckert (2007)
estimated that a minimum of 2,000
leatherback turtles were killed annually
by the combined swordfish fishing
operations (only gillnet) off Peru and
Chile. After some fleets switched from
large mesh gillnet to longline to target
swordfish, this estimate has declined by
at least an order in magnitude. Research
conducted in the Chilean large-mesh
gillnet fishery to reduce bycatch of
marine mammals and sea turtles
indicates that less than five leatherback
turtles have interacted with the fishery
(on observed vessels) since 2014, and all
were released alive (C. Fahy, NMFS,
pers. comm., 2018).
In Peru, the capture of leatherback
turtles has been prohibited since 1976,
although retention of bycaught
leatherback turtles continues (FAO
2004). From 1985 to 1999, based on
field books, diaries, specimen data
sheets, fishery statistics files and
unpublished reports, 30 leatherback
turtles were captured in fisheries (in
Alfaro-Shigueto et al. 2007). From July
2000 to November 2003, observers at 8
ports, from Mancora in northern Peru to
Morro Sama in the south, reported 133
leatherback turtles caught by artisanal
fishing gear, with 76 percent caught in
gillnets and 24 percent caught in
longlines targeting fish, sharks, and rays
(Alfaro-Shigueto et al. 2007). Of the total
caught, 41.4 percent (n = 55) were
released alive and 58.6 percent (n = 78)
were retained for human consumption.
Of the leatherback turtles retained and
measured (n = 6), the size ranged from
98 to 123 cm curved carapace length
(CCL), indicating that both subadults
and adults are encountered by artisanal
fisheries off Peru. Researchers recently
assessed and quantified sea turtle
mortality levels in one fishing village in
central-southern Peru (San Andre´s)
through sampling dump sites (97.3
percent) and strandings (2.7 percent)
over a 5-year period (2009 to 2014). Of
953 carapaces recorded, leatherbacks
comprised only 1.4 percent of sea turtles
(n = 13). However, this study still
confirmed that they were consumed or
sold for human consumption. With a
mean CCL of 113.0 cm (range: 80 to 135,
n = 10), 70 percent of the leatherbacks
were juveniles and 30 percent were subadults. There were no adults.
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Researchers noted that the meat was
used to support separate demands:
Fishermen families’ consumption, local
trade, and ‘‘special’’ orders from Lima
(Quispe et al. 2019). Using data from
shore-based and on-board observers,
Alfaro-Shigueto et al. (2011) estimated
the mean annual leatherback bycatch as
follows: 40 turtles (with a range of 37 to
44) in the driftnet fishery, with 80
percent released alive; six turtles (with
a range of 3 to 9) in the dolphinfish
longline fishery, all released alive; and
26 turtles (with a range of 24 to 27) in
the shark longline fishery, all released
alive. Alfaro-Shigueto et al. (2015)
assessed the bycatch of leatherback
turtles in driftnet vessels in northern
Peru (through at-sea monitoring) and
central Peru (shore-based monitoring).
From December 2013 to November
2014, 31 leatherback turtles were
captured, of which 13 died. Interactions
occurred primarily with juveniles and
subadults (mean CCL was 125.1 ± 14.8).
Nearshore driftnets from San Jose
(northern Peru) captured 20 leatherback
turtles (five dead). At least one animal
was butchered, indicating that even
animals caught alive may be killed,
despite Peruvian laws restricting such
practices. Approximately 3,000 net
vessels fish along the coast of Peru, but
only a fraction were included in this
study (Alfaro-Shigueto et al. 2015).
Efforts are being made to patrol nets to
reduce bycatch, conduct extensive
education and outreach, and increase
regulation and enforcement (AlfaroShigueto et al. 2015). A review of
information collected from official
statistics, literature, and surveys of
beaches and dumpsites revealed that the
size of captured leatherback turtles
declined over the years. In 1987, the
mean CCL of captured leatherback
turtles was 117 ± 10.65 cm, while in
2005, the mean CCL was 109.27 ± 14.4,
possibly indicating overexploitation due
to systematic and sustained harvests,
particularly during El Nin˜o years
(Campos et al. 2009). Greater captures of
all sea turtles, including leatherback
turtles, occurred during periods of El
Nin˜o, when turtles are more likely to be
found in more coastal waters (where
there is increased artisanal fishery
activity) due to environmental
variability and availability of jellyfish in
those areas (Campos et al. 2009).
In Chile, a commercial fishery was
established in 2001 that permitted
longlining for swordfish (shallow-set)
with the condition that all vessels were
required to take an observer on board to
collect information on bycatch. Between
2001 and 2005, over 10 million hooks
were observed, and leatherback turtles
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were the most common species caught
(n = 284), with the majority (n = 282)
released alive. Leatherback turtles were
caught primarily between 24° S and
38° S (furthest south was 38°39′ S and
84°15′ W) in less than 4 percent of the
sets with an overall mean of 0.0268
turtles per one thousand hooks. Size
estimates revealed both juveniles and
adults. Fishermen were trained to use
the best practices for de-hooking,
disentangling, and releasing sea turtles,
which likely increased the survival rate
of leatherback turtles (Donoso and
Dutton 2010). Researchers recently
presented information on the incidental
capture of sea turtles in industrial and
artisanal longlines, gillnets and artisanal
espinel (i.e., small-scale handline or
longline) fisheries all targeting
swordfish off Chile (Za´rate et al. 2019).
Over an 8-year period (2006–2014), 182
leatherbacks were documented as
bycatch (mortality of bycaught turtles
was not reported). Over this study
period, 44 percent of turtles were caught
in industrial longline, 28 percent in
artisanal espinel, 17 percent in gillnets
and 11 percent in artisanal longline
(with sea turtle species undefined).
Researchers noted that while observer
coverage in the industrial longline fleet
has been generally high (>70 percent of
total fishing trips), the monitoring
coverage of artisanal espinel and gillnets
is very low (<3 percent). Thus, these
estimates of bycatch can be considered
minimal. While the number of
industrial and artisanal vessels has
declined (from 12 vessels in 2001 to 3
vessels in 2014, the number of artisanal
espinel and gillnet vessels has not
declined, remaining around 90 vessels
(Za´rate et al. 2019).
We conclude that juvenile and adult
life stages of the East Pacific DPS are
exposed to high fishing effort
throughout their foraging range and in
coastal waters near nesting beaches.
Mortality is also high in some fisheries,
with reported mortality rates of up to 58
percent due in part to the use of gillnets
and as well as consumption of bycaught
turtles in Peru. As noted above, there
have been efforts by individual nations
and regional fishery management
organizations to mitigate and reduce the
threat of bycatch, but those efforts have
not been successful at ameliorating the
risks. We conclude that fisheries
bycatch remains a major threat to the
East Pacific DPS.
Pollution
Pollution is a threat to the East Pacific
DPS. Pollution includes contaminants,
marine debris, and ghost fishing gear.
The South Pacific Garbage Patch,
discovered in 2011 and confirmed in
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2017, contains an area of elevated levels
of marine debris and plastic particle
pollution, most of which is concentrated
within the ocean’s pelagic zone and in
area where leatherback turtles forage for
many years of their life. The area is
located within the South Pacific Gyre,
which spans from waters east of
Australia to the South American
continent and as far north as the
Equator.
Given the amount of floating debris in
the Pacific Ocean (Lebreton et al. 2018),
marine debris has the potential to be a
significant threat to the East Pacific
leatherback population. The precise
impact cannot be quantified using the
best available data. Leatherback turtles
subsist primarily on jellyfish and other
gelatinous zooplankton and may be
prone to ingesting plastics resembling
their food source (Mrosovsky 1981;
Schuler et al. 2013, 2015). Dead
leatherback turtles have been found
choked on plastic bags, and phthalates
derived from plastics have been found
in leatherback egg yolk (Lebreton et al.
2018).
Prior to the early 1990s, high seas
driftnet fisheries freely operated in the
Pacific Ocean and interacted with
thousands of sea turtles. Researchers
estimated that over 1,000 leatherback
turtles were taken by the combined
fleets of Japan, Korea, and Taiwan
during a one-year period (Wetherall
1997). However, because genetic
analyses of Pacific leatherback turtles
were relatively new at that time, the
data does not indicate the nesting beach
origin of those bycaught leatherback
turtles. In 1992, a UN moratorium
banned high seas driftnet fisheries, so
that active large scale driftnets no longer
pose a threat to leatherback turtles.
However, numerous discarded driftnets
continue to entangle and drown
leatherback turtles in a phenomenon
known as ‘‘ghost fishing’’ (Gilman et al.
2016),
In 2007, the IATTC passed a
resolution pertaining to sea turtle
bycatch in purse seine and longline
fisheries which primarily target tuna. In
order to address the marine debris and
potential interactions with sea turtles in
the eastern tropical Pacific Ocean,
fishermen are required to disentangle
sea turtles entangled in fish aggregating
devices, even if the device does not
belong to the vessel.
Only a few studies of levels or effects
of toxins on leatherback turtles have
examined effects to their health and
fitness, as well as any effects to eggs and
hatchlings. Sill et al. (2008) sampled
non-viable leatherback eggs and
hatchlings that died in the egg chamber
at Las Baulas National Park. Researchers
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analyzed the samples for metals and
other toxicants to explore the
relationship between pollution and
hatching success for 30 females. Metal
levels were highly variable, but there
were no significant differences within
and between groups of females, and
none of the pesticides tested were
present in the samples (Sill et al. 2008).
Overall, the study found no relationship
between metal concentrations and
hatching success. The researchers
postulated that eggs may take up some
metals from the nest environment and
deposit other metals in the egg shell, as
unhatched eggs contained more nickel,
copper, and cadmium and contained
significantly less iron, manganese and
zinc than dead hatchlings (Sill and
Paladino 2008).
As with all leatherback turtles,
entanglement in and ingestion of marine
debris and plastics is a threat that likely
kills several individuals a year.
However, data are not available because
most affected turtles are not observed.
Given the amount of pollution turtles
are exposed to throughout their lifetime,
this has the potential to be a significant
threat to the East Pacific leatherback
population, although the impact cannot
be quantified using the best available
data. We conclude that pollution is a
threat to this DPS.
Oceanographic Regime Shifts
The East Pacific DPS is affected by
oceanographic regime shifts. In the
eastern equatorial Pacific Ocean,
reductions in productivity parameters
are primarily associated with ENSO,
during which sex ratios become biased
up to 100 percent female (Santidria´n
Tomillo et al. 2014). There is also an
effect on hatching and emergence
success in North Pacific Costa Rica
(Santidria´n Tomillo et al. 2012): During
El Nin˜o years, hatching success is very
low due to dry and hot conditions on
the nesting beaches and is high during
La Nin˜a events due to increased
precipitation in this area. La Nin˜a
events are characterized by high
phytoplankton productivity, cooler sea
surface temperatures, enhanced
precipitation in northwestern Costa
Rica, and cooler air temperatures. These
factors lead to increases in the biomass
and distribution of gelatinous
zooplankton, the primary food of
leatherback turtles. Foraging success
and the frequency of reproduction are
enhanced following such periods of
high primary productivity (Saba et al.
2007). Nesting seasons that follow the
La Nin˜a events, result in peaks in the
number of nesting females, higher than
average hatching success and emergence
rates, and a larger proportion of male
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hatchlings (Saba et al. 2012). Saba et al.
(2008) found that a shift from 1 °C to
¥1 °C in the El Nin˜o sea surface
temperature anomaly resulted in a fivefold increase in leatherback remigration
probabilities at Playa Grande. Such
large-scale regime shifts are likely to
affect the entire DPS. Productivity is
positively (La Nin˜a) or negatively (El
Nin˜o) impacted. Wallace et al (2006)
hypothesize that prey availability
related to ENSO exacerbates the effects
of fisheries bycatch mortality, resulting
in declining trends. Because of the small
abundance of the DPS, extended El Nin˜o
events are likely to pose a threat to the
East Pacific DPS.
Climate Change
Climate change is a threat to the East
Pacific DPS. The impacts of climate
change include: Increases in
temperatures (air, sand, and sea
surface); sea level rise; increased coastal
erosion; more frequent and intense
storm events; and changes in
oceanographic regimes and currents.
Climate projections assessed by the
IPCC indicate that Central America is
very likely (defined as 90 to 99 percent
probability; IPCC 2007) to become
warmer and likely (defined as 66 to 90
percent probability; IPCC 2007) to
become drier by 2100 (Saba et al. 2012).
In addition, climate variability is likely
to change the strength and frequency of
El Nin˜o events, although there is less
scientific consensus on the frequency
and magnitude of changes to these
events. A climate-forced population
dynamics model developed by Saba et
al. (2012) showed sea surface
temperatures to be highly correlated
with large phytoplankton productivity
throughout a 100-year projection to the
year 2100. Relative to a stable nesting
population given mean surface air
temperatures and precipitation from
1975 to 1999, Saba et al. (2012)
estimated that the nesting population at
Playa Grande would decline at a rate of
7 (±1) percent per decade over the next
century of climate change under a
scenario which considered increasing
emissions from 2000 to 2100 (A2
scenario). Similar declines occurred for
other scenarios (Special Report on
Emissions Scenarios 2007). The nesting
population was projected to remain
stable up until around 2030 but reduced
75 percent by the year 2100. Hatching
success and emergence rates, which
would decrease associated with 2.5 °C
warming of the nesting beaches, served
as a primary driver of the decline.
Santidria´n Tomillo et al. (2012)
developed a similar climate forcing
model, which considered projected
changes associated with El Nin˜o events
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and demonstrated that hatching success
would decline from approximately 42 to
18 percent by 2100, while emergence
rates would decline between
approximately 76 to 29 percent. The
authors concluded that even with
protection at the primary nesting
beaches in Costa Rica, with the general
warming of Central America in the near
future, the chances of a new nesting area
emerging with more ideal conditions
(i.e., cooler and wetter) is unlikely
(Santidria´n Tomillo et al. 2012).
Increasing sand temperature is an
existing threat to the DPS. The longterm data set on leatherback turtles
nesting at Playa Grande, Costa Rica
indicates reduced emergence success,
skewed sex ratios, and increased
hatchling mortality as a result of
increased sand temperature (Santidria´n
Tomillo et al. 2015). From 2004 to 2013,
primary sex ratios fluctuated between a
minimum sex ratio of 41 percent
females (and the only year with a malebiased hatchling production) to 100
percent females produced during two
seasons (Santidria´n Tomillo et al. 2014).
Low emergence success and low
hatchling output (i.e., higher mortality
as a result of high sand temperatures)
were associated with a strongly biased
female ratio, because these resulted
from female-producing high
temperatures. Variability in these results
occur during and between nesting
seasons, largely due to highly variable
climatic conditions in northwestern
Costa Rica, resulting in ‘‘boom-bust’’
cycles in leatherback hatchling
production and primary sex ratios (in
Santidria´n Tomillo et al. 2014). Sand
temperatures are projected to continue
to increase, which will likely result in
a further decline in the number of
hatchlings produced (Santidria´n
Tomillo et al. 2014). An increase in the
percentage of females could potentially
benefit the productivity of the DPS in
the short-term. However, any such
benefits would be tempered by the
associated lower emergence and
hatchling success rates. Relocation of
sea turtle clutches that may be
‘‘doomed’’ due to high sand
temperatures and inundation is a
common conservation practice,
particularly at areas with warming
beaches. However, relocation is not
always possible and is also associated
with lower emergence and hatchling
success rates.
In addition to climate change
influencing the nesting beach habitat of
eastern Pacific leatherback turtles, the
impacts of a warming ocean may also
affect the environmental variables of
their pelagic migratory and foraging
habitat, which may further increase
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population declines. As mentioned
previously, the preferred foraging
habitat of eastern Pacific is
characterized by relatively low sea
surface temperatures and low levels of
chlorophyll-a. Using information
derived from satellite tracked
leatherback turtles, which established
migratory pathways and core foraging
habitat (as summarized in Shillinger et
al. 2008), in combination with
generalized additive mixed models,
researchers were able to project that
between 2001 and 2100, there would be
a net loss of the core foraging habitat of
the DPS. The loss was predicted to be
a 15 percent decline over the next
century (Willis-Norton et al. 2014).
Depending on whether this population
is able to shift their preferred migratory
routes and foraging habitat over time
(which is unclear), remigration intervals
may shorten or lengthen, which could
influence reproductive productivity.
Climate change is a threat to the East
Pacific DPS that affects nesting females
(e.g., remigration interval and fitness),
their progeny (e.g., hatching success,
embryonic development, and
feminization of hatchlings), and foraging
subadult and adult leatherback turtles.
Detrimental impacts of increased sand
temperatures have already occurred and
are likely to continue or worsen.
Foraging areas will also be impacted via
changes in ocean productivity, sea
surface temperatures, and availability of
prey.
Conservation Efforts
There are numerous efforts to
conserve the leatherback turtle. The
following conservation efforts apply to
turtles of the East Pacific DPS (for a
description of each effort, please see the
section on conservation efforts for the
overall species): Convention on the
Conservation of Migratory Species of
Wild Animals, Convention on Biological
Diversity, Convention on International
Trade in Endangered Species of Wild
Fauna and Flora, Convention for the
Protection of the Marine Environment
and Coastal Area of the South-East
Pacific (Lima Convention), Convention
for the Conservation and Management of
Highly Migratory Fish Stocks in the
Western and Central Pacific Ocean
(WCPF Convention), Convention
Concerning the Protection of the World
Cultural and Natural Heritage (World
Heritage Convention), Eastern Pacific
Leatherback Network, Eastern Tropical
Pacific Marine Corridor Initiative, FAO
Technical Consultation on Sea TurtleFishery Interactions, IAC, MARPOL,
IUCN, Ramsar Convention on Wetlands,
RFMOs, Secretariat of the Pacific
Regional Environment Programme,
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UNCLOS, and UN Resolution 44/225 on
Large-Scale Pelagic Driftnet Fishing.
Although numerous conservation efforts
apply to the turtles of this DPS, they do
not adequately reduce its risk of
extinction.
Extinction Risk Analysis
After reviewing the best available
information, the Team concluded that
the East Pacific DPS is at high risk of
extinction. The DPS exhibits a total
index of nesting female abundance of
755 females at monitored beaches. Such
a limited nesting population size makes
this DPS vulnerable to stochastic or
catastrophic events that increase its
extinction risk. This DPS exhibits a
decreasing nest trend, which along with
lower than-average productivity metrics,
has the potential to further reduce
abundance and increase the risk of
extinction. The nesting range is
somewhat limited to the Pacific Central
American coast, with little diversity
among sites. Thus, stochastic events
could have catastrophic effects on
nesting for the entire DPS, with no
distant subpopulations to buffer losses
or provide additional diversity. Most
foraging occurs in the eastern Pacific
Ocean, which is subject to
oceanographic regimes shifts that
expose the DPS to low-productivity
events. Based on these demographic
factors, we find the DPS to be at risk of
extinction as a result of past threats.
Current threats also contribute to the
risk of extinction of this DPS. Fisheries
bycatch is the major threat, capturing,
and often killing, turtles throughout
their foraging areas, thus reducing
abundance. There are few mechanisms
in place, including internationally
through the IATTC or other bilateral or
international instruments and through
monitoring and enforcement of coastal
fisheries laws, to mitigate or reduce
bycatch. Overutilization is also a major
threat. Historically, harvest of turtles
and eggs reduced the once high
abundance of turtles to current low
levels. The poaching of eggs continues,
reducing productivity, especially at
unprotected beaches, where egg
collection may reach 100 percent and
nesting females may also be at risk of
poaching. The effects of climate change,
including the observed and predicted
increase in frequency and strength of
ENSO events (i.e., oceanographic regime
shifts), are threats to this DPS, given its
restricted foraging range and the
vulnerability of nesting beaches to high
sand temperatures and low levels of
rainfall, which affect sex ratios and
emergence and hatching success (i.e.,
productivity). Additional threats
include: Habitat loss and modification;
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predation; and pollution. Development
modifies nesting habitat. However, most
beaches are protected throughout the
nesting range. Though many regulatory
mechanisms are in place, they do not
adequately reduce the impact of these
threats. Further, it is important to note
that efforts (e.g., relocation) to protect
and mitigate threats from the harvest of
turtles and eggs, predation, and
environmental impacts related to
erosion and lethal temperatures are
dependent upon the presence of
monitoring or management programs.
Some of these are dependent on funding
from the MTCA. Even when undertaken,
these efforts may not be successful.
We determine, consistent with the
Team’s findings, that the East Pacific
DPS is currently in danger of extinction.
Its nesting female abundance and
declining trend make the DPS highly
vulnerable to threats. Though numerous
conservation efforts apply to this DPS,
they do not adequately reduce the risk
of extinction. We conclude that the East
Pacific DPS is currently in danger of
extinction throughout its range and
therefore meets the definition of an
endangered species. The threatened
species definition does not apply
because the DPS is currently at risk of
extinction (i.e., at present), rather than
on a trajectory to become so within the
foreseeable future.
Leatherback Turtle, Overall Species
The petition under review sought
specifically to identify the NW Atlantic
population of leatherback sea turtles as
a separate DPS and assign it a different
status from the global listing. As
explained throughout this finding, we
have determined that seven leatherback
populations would satisfy the tests for
recognition under our DPS Policy (i.e.,
that they are discrete from one another
and significant to the overall species),
and we have referred to these
hypothetically, for purposes of our
analysis only, as DPSs. This includes
the NW Atlantic DPS. However, we
have also determined that, even if these
populations were formally recognized as
DPSs through a listing process under the
Act, each of the DPSs would have the
same status as the overall species,
which is currently listed throughout its
range (globally) as endangered. Nothing
in the petition or in the best available
information we have reviewed has led
us to conclude that there is any basis to
disturb the long-standing global listing,
which remains in effect and is
unaffected by this finding. For
completeness, here we present an
overview of current information
pertaining to the status of the overall
species, including a summary of some of
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the key information from the DPSspecific sections as well as an
evaluation of the demographic factors
affecting the overall species.
As explained in the Background
section, the leatherback turtle was
originally listed as endangered in 1970
under the precursor to the ESA and was
carried forward as an ‘‘endangered
species’’ when the ESA became
effective. The Services designated the
nesting beaches at Sandy Point, St.
Croix (43 FR 43688; September 26,
1978) and surrounding marine waters
(44 FR 17710; March 23, 1979) as
critical habitat. NMFS designated
additional marine habitat along 41,914
square miles (108,558 square km) of the
U.S. West Coast as critical habitat (77
FR 4170; January 26, 2012). The
Services issued the recovery plans for
leatherback turtles in the U.S.
Caribbean, Atlantic, and Gulf of Mexico
(1991) and U.S. Pacific (1998; https://
www.fisheries.noaa.gov/action/
recovery-plans-leatherback-sea-turtle).
The species has the widest
distribution of any reptile, with a global
range extending from 71° N, based on an
at-sea capture off Norway (Carriol and
Vader 2002) to 47° S, based on an at-sea
sighting off New Zealand (Eggleston
1971; Eckert et al. 2012). The species
has several thermoregulatory
adaptations to allow such a large
latitudinal range, maintain its core
temperature while foraging, and avoid
overheating during nesting. These
include its large size, low metabolic
rates, countercurrent heat exchange at
the base of its limbs, and peripheral
insulation (Frair et al. 1972; Greer et al.
1973; Paladino et al. 1990; Fossette et al.
2009; Bostrom et al. 2010; Eckert et al.
2012; Casey et al. 2014; reviewed in
Wallace and Jones 2015).
Nesting is restricted to mainly tropical
or subtropical beaches. However,
nesting also occurs on temperate
beaches of the SW Indian Ocean
(Pritchard and Mortimer 1999). Nesting
usually occurs on high-energy beaches
(Pritchard 1976), resulting in high rates
of natural erosion. The primary factors
influencing shoreline suitability for
nesting appear to be a lack of abrasive
substrate material, a deep-water
approach to minimize energy
expenditure needed to reach nesting
sites, and proximity to oceanic currents
that can facilitate hatchling dispersal
(Eckert et al. 2012). Leatherback turtles
appear to prefer wide, long beaches with
a steep slope, deep rock-free sand, and
an unobstructed deep water or softbottom approach (Pritchard and
Mortimer 1999; Eckert et al. 2015). As
a result, it has been proposed that the
choice of nesting location is based on
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site characteristics within a geographic
location (MacKay et al. 2014).
Foraging areas are generally
characterized by zones of upwelling,
including off the edges of continents,
where major currents converge, and in
deep-water eddies (Saba 2013).
Important foraging areas include but are
not limited to: upwelling off the west
coasts of North and South America
(Benson et al. 2011; Roe et al. 2014);
Benguela Current Marine Ecosystem
(Honig et al. 2007); and Canadian waters
on the Scotian Shelf (James et al. 2005a,
2006b, 2007b).
Abundance
Adding together the total indices of
nesting female abundance for all DPSs,
the total index of nesting female
abundance for the species is 32,174
females. This number, however, should
be considered as a compilation of seven
populations ranging in size from 27 to
20,659 nesting females because nesting
female exchange does not occur
between DPSs.
Comparisons with historical accounts
of nesting female abundance are
complicated by the discovery of new
nesting beaches over time, changes in
remigration intervals and/or clutch
frequency, and modified observational
effort. Abundance estimates for even
large nesting beaches were not available
prior to 1950 (Rivalan et al. 2006),
several large nesting beaches were not
discovered until the 1960s or later
(NMFS and USFWS 2013), and
monitoring efforts were variable over
time. Pritchard’s 1971 global estimate of
29,000 to 40,000 nesting females
included a maximum estimate (i.e.,
40,000 nesting females) based on the
assumption that large nesting
aggregations had yet to be discovered
(Pritchard 1971); this estimate did not
include large nesting female abundances
from the East Pacific and SE Atlantic
Oceans. At that time, the nesting
aggregation at Terengganu, Malaysia
nesting population was thought to be
one of the largest; however it has since
been extirpated (Chan and Liew 1996).
In 1982, Pritchard revised his initial
global estimate to 115,000 nesting
females, based largely on the nesting
beaches in Pacific Mexico (n = 75,000;
Pritchard 1982). However, the 1982
estimate was extrapolated from a brief
aerial survey and may have been an
overestimate (Pritchard 1996). When the
Mexico nesting population collapsed,
Spotila (1996) estimated the total global
estimate to be 34,500 nesting females,
with a range of 26,200 to 42,900 nesting
females. However, this estimate did not
include the nesting aggregation in
Gabon, which in 2002 was identified as
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the largest in the world at that time,
with tens of thousands of nesting
females (Witt et al. 2009). Recent data
indicate less than 9,000 nesting females
in Gabon (Formia in progress). Thus, we
find that leatherback nesting female
abundance has declined rapidly in
several populations. Our total index of
nesting female abundance for the
species, which does include the largest
nesting aggregations from all DPSs, is
lower than previous estimates by at least
10,000 females.
Species go extinct through the loss of
populations. Therefore, the loss of any
of these populations (which we refer to
in this finding hypothetically as DPSs)
would increase the extinction risk of the
species. Most of the DPSs exhibit total
indices of nesting female abundances
that place them at risk for
environmental variation, genetic
complications, demographic
stochasticity, negative ecological
feedback, and catastrophes (McElhany
et al. 2000; NMFS 2017). The current
total index of nesting female abundance
for the species reflects the impact of
threats that have affected the species to
this point. This reduced abundance
renders it particularly vulnerable to
threats and contributes to its extinction
risk.
Spatial Distribution
Productivity
Diversity
Nest trends are decreasing across the
species, except at the least abundant
nesting aggregation in Brazil (i.e., the SE
Atlantic DPS), with a total index of 27
nesting females, which is increasing by
4.8 percent annually. Current nest
trends are declining at rates ranging
from ¥0.3 percent (within the SW
Indian DPS) to ¥9.3 percent (the overall
decline for the NW Atlantic DPS).
Historical declines are even larger.
Aerial surveys of nesting beaches in
Mexico detected declines from over
70,000 nesting females in 1982 to fewer
than 250 in 1998, with an annual
mortality rate of 22.7 percent (Spotila
2000) and an overall decline of 97.4
percent in three generations (Wallace et
al. 2013). The Terengganu, Malaysia
nesting aggregation has declined by 17.9
percent annually from 1967 to 2010. It
was been reduced to less than one
percent of its original size between the
1950s and 1995 (Chan and Liew 1996)
and is now considered functionally
extirpated. Significant declines in
nesting have been documented for other
populations (Benson et al. 2015).
Declining nesting trends reflect the
impact of threats that have been
operating on the species, and these
trends increase the extinction risk of the
species.
Relative to other sea turtle species, the
leatherback turtle has low genetic
diversity and shallow mtDNA
coalescence (Dutton et al. 1999),
reflecting its recent global radiation, i.e.,
Post-Pleistocene expansion from a
refugium in the Indian Ocean (Dutton et
al. 1999). As a species, it uses diverse
and widely distributed nesting and
forage areas. Differences in size at
maturity, remigration rate, clutch
frequency, and clutch size likely reflect
environmental variability among DPSs
(Saba et al. 2008; Saba et al. 2015). The
age of the species and its flexible use of
multiple foraging and nesting areas
indicate that the species has some
resilience to stochastic and
environmental changes.
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The species occurs over a broad
spatial range, in tropical and temperate
waters worldwide, from 71° N to 47° S
(Goff and Lien 1988; Carriol and Vader
2002; McMahon and Hayes 2006;
Shillinger et al. 2008; Wallace et al.
2010; Benson et al. 2011; Eckert et al.
2012). It nests and forages across a wide
spatial range, which provides some
degree of resilience against local
impacts to nesting and foraging areas.
The DPSs are reproductively isolated
with little to no gene flow connecting
them. However, within some DPSs there
is fine-scale population structure
(Dutton et al. 1999; Dutton et al. 2003;
Dutton et al. 2013; Molfetti et al. 2013).
These subpopulations exhibit
metapopulation dynamics, which make
a DPS more resilient to stochastic and
environmental changes. It is likely that
all DPSs once exhibited such dynamics,
given the ephemeral, high-energy
beaches where they nest and their
regional, but not necessarily beachspecific, philopatry (Dutton et al. 1999;
Dutton et al. 2013). However, the
reduction of nesting aggregations within
a DPS has likely reduced or removed
this structure, and the associated
resilience, in some DPSs and in the
overall species.
Present or Threatened Destruction,
Modification, or Curtailment of Habitat
or Range
The destruction or modification of
nesting habitat is a threat to most
leatherback turtles, and in some areas,
this threat is major, as a result of
development, erosion, or obstruction
from logs. By the year 2025, the UN
Educational, Scientific and Cultural
Organization (2001) forecasts that
human population growth and
migration will result in 75 percent of
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people living within 60 km of the sea.
This will place significant additional
pressure on coastal habitats.
Coastal development and associated
activities cause accelerated erosion rates
and interruption of natural shoreline
migration (National Research Council
1990). Numerous beaches are eroding
due to both natural (e.g., storms, sea
level changes, waves, shoreline geology)
and anthropogenic (e.g., development
and expansion, construction of armoring
structures, groins, jetties, marinas,
coastal development, inlet dredging)
factors. Such shoreline erosion has led
and will continue to lead to a loss of
nesting habitat for leatherback turtles
and potential loss of nests from
inundation. Erosion or inundation and
accretion of sand above incubating nests
appear to be the principal abiotic factors
that negatively affect incubating egg
clutches in some areas (Dow et al. 2007;
USFWS 1999; NMFS and USFWS 2013).
Shoreline structuring can also
physically prevent females from
reaching suitable nesting habitat or
prevent them from returning to sea
(Witherington et al. 2011).
Low hatching success, relative to
other sea turtle species, is characteristic
of many leatherback populations despite
high fertility rates (reviewed by Bell et
al. 2003; Eckert et al. 2012). Nest
relocation is undertaken as a
conservation measure in some locations
when erosion (or poaching and
predation) threaten the viability of a
nest. However, studies have found that
hatching success of nests in hatcheries
or nests relocated to another area of a
beach is lower than in situ nests
(reviewed in Herna´ndez et al. 2007;
Eckert et al. 2012). In addition, nest
relocation results in altered sand
temperatures, which influences the sex
ratio of hatchlings produced (Sieg et al.
2011).
Coastal development and expansion
also contributes to habitat degradation
via artificial lighting (i.e., light
pollution). The presence of artificial
lighting on or adjacent to nesting
beaches alters the behavior of nesting
females (often deterring nesting) and is
often fatal to post-nesting females and
emerging hatchlings, when they are
attracted to terrestrial light sources and
drawn away from the water
(Witherington 1992; Sella et al. 2006;
Witherington et al. 2014). As hatchlings
head toward lights or meander along the
beach, their exposure to predators and
likelihood of desiccation are greatly
increased. Artificial lighting may also
affect hatchlings that successfully find
the water, causing them to be
misoriented after entering the surf zone
or while in nearshore waters.
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The modification of nesting habitat
generally results in loss of productivity
for the species, as a result of reductions
in nest and hatching success. In
addition, several DPSs experience
nesting beach habitat modifications
(e.g., artificial lighting, logs, and other
obstructions) that result in the death of
nesting females and hatchlings.
Therefore, abundance is also reduced,
posing an even greater threat to the
continued existence of the turtles of the
DPS. The loss and modification of
nesting habitat poses a major threat to
the species.
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Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
Historically, the harvest of turtles and
eggs was the primary threat to the
species, leading to the loss of severe
depletion of many nesting aggregations
worldwide (Spotila et al. 1996). At one
point in time, egg harvest was
ubiquitous with all nests taken at many
beaches (Chan and Liew 1996; Sarti et
al. 2007; reviewed by Eckert et al. 2012).
For the NW Atlantic, NE Indian, and
West Pacific DPSs, legal harvest of turtle
and/or eggs continues. Despite laws in
many countries, the poaching of eggs
continues at most nesting beaches,
ranging in severity from minor at
monitored or protected beaches to near
100 percent harvest at unmonitored
beaches. Nesting females, and turtles
caught at sea, continue to be poached
for their meat, eggs, and fat in many
locations (Eckert et al. 2012). As
described in detail in the prior sections
evaluating the status of each individual
DPS, the harvest of eggs and turtles is
a threat to each and to the species
overall, and for the NE Indian and West
Pacific DPSs, it is a primary threat. The
legal and illegal harvest of turtles and
eggs poses a threat to the species.
Disease or Predation
We do not have adequate information
on disease to assess its impact on the
species. However, we have enough
information to conclude that predation
is clearly a threat. Numerous species
prey on leatherback eggs and hatchlings.
Eckert et al. (2012) provide an
exhaustive list of the documented
predators for each life stage and area.
For eggs, common predators include
ants, ghost crabs, monitor lizards,
crows, mongoose, domestic and feral
dogs, and feral pigs (Eckert et al. 2012).
For hatchlings, common predators
include the terrestrial predators listed
above as well as numerous species of
carnivorous fish, including sharks.
Sharks and killer whales, and in some
areas jaguars and crocodiles, prey on
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subadult and adult turtles. Predation on
eggs and hatchlings is common and
reduces productivity of the species;
predation on subadults and adults is
less prevalent but reduces abundance
when it occurs. Predation is a threat to
the species, and for some DPSs, it is a
major threat.
Inadequacy of Existing Regulatory
Mechanisms
Numerous regulatory mechanisms
provide certain protections to sea turtles
at the international, regional, national,
and local levels. For example, the
harvest of sea turtles and their eggs is
prohibited by regional conventions and
national laws. Fisheries bycatch is also
addressed, although not
comprehensively, by several
international and national instruments
and/or governing bodies. However, as
we detail below and has been discussed
in prior sections reviewing each
individual DPS, these measures are
often poorly implemented or enforced,
resulting in inadequate protections
against the threats they are designed to
ameliorate.
In some nations (e.g., South Africa)
sea turtles were among the first species
to receive legal protections and have
been the focus of concentrated
conservation efforts. However, current
regulatory mechanisms often fall short
of preventing further population
declines and ensuring persistence
(Eckert et al. 2012). For many nations
the regulations in place are inadequate
(usually due to lack of enforcement and
implementation) to address the impacts
of a wide range of anthropogenic
activities that directly injure and kill
turtles, disturb eggs, disrupt necessary
behaviors, and alter terrestrial and
marine habitats used by the species. In
many areas, regulations for the harvest
of turtles and eggs are inadequate due to
a lack of enforcement. In some areas, the
regulation of fisheries bycatch do not
adequately reduce associated mortality.
Fishery observer coverage is often
inadequate to accurately estimate
leatherback bycatch.
Due in part to their worldwide
distribution and highly migratory
nature, combined with nesting site
fidelity, leatherback turtles require
international, national, regional, and
local protection. Hykle (2002) and
Tiwari (2002) reviewed the value of
some international instruments and
concluded that they vary in their
effectiveness. Often, international
treaties do not realize their full potential
because: They do not include all key
nations; do not specifically address sea
turtle conservation; are handicapped by
the lack of a sovereign authority to
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promote enforcement; and/or lack of
legally-binding requirements. Lack of
implementation or enforcement by some
nations may make them less effective
than if they were implemented in a
more consistent manner across the
target region. A thorough discussion of
this topic is available in the 2002
special issue of the Journal of
International Wildlife Law and Policy:
International Instruments and Marine
Turtle Conservation (Hykle 2002).
Additional information on national,
regional, and local protection is
provided in the prior sections of this
finding relating to each individual DPS.
In summary, numerous regulatory
mechanisms protect leatherback turtles,
eggs, and nesting habitat throughout the
range of the species. Although the
regulatory mechanisms provide some
protection, many do not adequately
reduce the threat that they were
designed to address, generally as a
result of limited implementation or
enforcement. As a result, bycatch,
incomplete nesting habitat protection,
and poaching remain threats to the
species. We conclude that the
inadequacy of the regulatory
mechanisms is a threat to the
leatherback turtle.
Fisheries Bycatch
Fisheries bycatch is the primary threat
to leatherback turtles (Crowder 2000;
Spotila et al. 2000; Lewison et al. 2004;
Wallace et al. 2011; Wallace et al. 2013;
Angel et al. 2014). It is a primary threat
to all DPSs. Leatherback turtles are
susceptible to bycatch in a wide range
of fisheries, from large scale commercial
to artisanal. Gear types that affect
leatherbacks include: longlines, purse
seines, driftnets, gillnets, trawls, pots/
traps, and pound nets (Gray and Diaz
2017). Turtles often drown after
becoming entangled in nets and other
gear or become injured and possibly die
as a result of hooking or interactions
with the gear. While bycatch in pelagic
shallow-set swordfish longline fisheries
has received the most attention to date,
small-scale coastal fisheries occur
worldwide, employing over 99 percent
of the world’s 51 million fishers (FAO
2011).
Bycatch data are most commonly
collected by trained observers on fishing
vessels or via surveys or interviews
(Lewison et al. 2015). Though often the
best available data on bycatch, observer
data generally cover less than five
percent of fisheries’ total effort
(Finkbeiner et al. 2011) and are rarely
available for small-scale fisheries
(Wallace et al. 2013; Lewison et al.
2015). The use of different metrics also
makes the data difficult to compare
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among fisheries, gear types, and regions
(Lewison et al. 2015). Therefore,
estimates of bycatch and resulting
mortality often underestimate the
magnitude of this threat.
Furthermore, IUU fishing is a
significant yet unquantified threat to sea
turtles worldwide. In addition to killing
and injuring turtles, it undermines
national and regional efforts to estimate
fisheries bycatch. IUU fishing represents
up to 26 million tonnes of fish caught
annually (https://www.fao.org/iuufishing/en/). We have no estimates of
the impacts to leatherback turtles from
IUU fishing, though interaction and
mortality rates are likely high because of
the magnitude of this additional fishing
pressure and because it is unregulated.
Generally, leatherback turtles do not
attempt to consume the bait associated
with fishing gear, as other sea turtles do,
but become entangled in fishing gear
(Lewison et al. 2015). Longline fisheries
involve the deployment of a horizontal
main line and vertical branchlines with
baited hooks, which may entangle
leatherback turtles. Bycatch reduction
measures include using circle hooks,
finfish bait, minimizing soak times, and
limiting mainline length (Angel et al.
2014; https://www.fisheries.noaa.gov/
national/bycatch/fishing-gear-pelagiclonglines#risks-to-sea-turtles). Purse
seines capture schools of fish in a
vertical wall of netting that can be
closed at the bottom (https://
www.fisheries.noaa.gov/national/
bycatch/fishing-gear-purse-seines);
bycatch rates are generally much lower
than longline bycatch rates (Angel et al.
2014). Leatherback turtles also become
entangled and drowned in drift or set
gillnets (https://www.fisheries.noaa.gov/
national/bycatch/fishing-gear-gillnets).
Gillnets can be devastating to
leatherback populations when set near
nesting beaches and represent the
primary threat to leatherback turtles in
some areas (e.g., Trinidad; Eckert and
Eckert 2005). Trawl fisheries drag nets
along the substrate or through the water
column and can capture and drown sea
turtles. Although TEDs may mitigate
this threat, they are not always required
or used in all areas. Vertical lines
extending and/or connecting pot and
trap gear with surface buoys commonly
entangle and can kill leatherback turtles.
Longline and net fisheries are often
the greatest threats to leatherback
turtles. In a global study of sea turtle
bycatch, Wallace et al. (2013) compiled
data (n = 239 records) published
between 1990 and 2011 to compare gear
types (longline, net, and trawl) and their
impacts to leatherback RMUs, which are
similar to the DPSs discussed in this
rule, though their exact boundaries
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differ. Wallace et al. (2013) defined high
bycatch impact as follows: A weighted
median bycatch per unit effort (BPUE)
greater than or equal to one; median
mortality rate greater than or equal to
0.5; and affecting adult or subadult
turtles. They found that longline
bycatch had a high impact on SW
Atlantic, SE Atlantic, and SW Indian
RMUs and that net bycatch had a high
impact on the NW Atlantic and East
Pacific RMUs (Wallace et al. 2013).
Integrating catch data from over 40
nations and bycatch data from 13
international observer programs,
Lewison et al. (2004) estimated the
numbers of leatherback turtles taken by
pelagic longliners to be more than
50,000 leatherback turtles in just one
year (2000). With over half of the total
fishing effort (targeting tuna and
swordfish) occurring in the Pacific
Ocean, an estimated 20,000 leatherback
turtles interacted with longline fishing
gear, with 1,000 to 3,200 mortalities in
2000 (Lewison et al. 2004). However,
Beverly and Chapman (2007) estimated
sea turtle longline bycatch mortality to
be approximately 20 percent of that
estimated by Lewison et al. (2004), or
approximately 200 to 640 leatherback
turtle mortalities annually. We consider
the estimate of Beverly and Chapman
(2007) to be more realistic, considering
the low nesting females abundance of
Pacific leatherback turtles, and because
Beverly and Chapman (2007) combined
the effort data from Lewison et al. (2004)
with bycatch data from Molony (2005)
that differentiated between deep-set and
shallow-set fisheries (which have
different interaction rates).
In the Pacific Ocean, Roe et al. (2014)
predicted leatherback turtle bycatch
hotspots by comparing the satellite
tracks of 135 adult turtles with longline
fishing effort. The greatest bycatch risk
occurred adjacent to primary nesting
beaches of the West Pacific DPS.
Bycatch risk was also high in the South
Pacific Gyre, where the East Pacific DPS
forages. Expanding on this study, a
study of observer data from 34
swordfish-targeting shallow-set longline
fleets found there were 331 leatherback
turtle interactions between 1989 and
2015 (Clarke 2017). Clarke (2017)
identified two bycatch hotspot areas:
Central North Pacific Ocean and eastern
Australia (Clarke 2017).
In the Atlantic Ocean, Fossette et al.
(2014) compared leatherback telemetry
data to longline fishing effort data from
ICCAT to identify nine areas in which
leatherback turtles are exposed to
bycatch associated with high longline
fishery pressure. The high pressure
fishing areas include foraging areas in
the North and South Atlantic Ocean and
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in waters off Brazil and western Africa.
These high pressure fishing areas are
not comparable to those identified by
Roe et al. (2014), who used a different
methodology, but both studies identify
high risk areas within each ocean basin.
Additional bycatch information that
we have set out in prior sections
specific to each DPS applies to our
consideration of the risk to the overall
species. In summary, fisheries bycatch
is a threat that is encountered by
numerous juvenile and adult
leatherback turtles. Mortality rates are
often high, and individuals that are
released may experience injuries or
sublethal effects associated with
entanglement, submergence, or
handling. Fisheries bycatch reduces
abundance, and when it prevents
nesting females from returning to
nesting beaches, reduces productivity as
well. Fisheries bycatch is the primary
threat to the leatherback species.
Vessel Strikes
Vessel strikes pose a threat to the
species throughout its range. As mature
individuals move from oceanic foraging
areas into coastal waters to reproduce,
they are exposed to a greater
concentration of vessels. Vessel strikes
off nesting beaches may injure or kill
these individuals, reducing the
abundance and productivity of the DPS.
Most vessel strikes likely go unnoticed
or unreported, making this threat
potentially much more significant that
documented occurrences would suggest.
Vessel strikes are a threat to the
leatherback species.
Pollution
We define pollution as including
contaminants, marine debris, and ghost
or derelict fishing gear. Such
interactions are likely to go unnoticed
and unreported and thus likely present
a more significant impact than
documented occurrences would suggest.
Leatherback turtles of all life stages are
vulnerable to oil spills, on land and at
sea, where exposure to oil and
dispersants occurs via contact (i.e.,
physical fouling), inhalation, or
ingestion (reviewed by Stacy et al. in
press).
Marine debris is ubiquitous
throughout the range of the species.
Marine debris includes plastics
(including plastic bags), microplastics,
derelict fishing gear (e.g., ghost nets and
other discarded or lost gear), and other
man-made materials. Leatherback turtles
may directly consume floating plastics,
mistaking it for their gelatinous prey or
accidentally ingest plastics while
foraging. In particular, plastic bags
appear similar to jellyfish in the marine
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environment, inappropriately triggering
the sensory cue to feed (Schuyler et al.
2014; Nelms et al. 2016). Plastic bags
have been found during necropsy of
stranded leatherback turtles, and
phthalates derived from plastics have
been found in leatherback egg yolk
(Lebreton et al. 2018). Mrosovsky et al.
(2009) reviewed 408 necropsy records
from 1885 to 2007 and found evidence
of plastic in the gastrointestinal tract of
34 percent of leatherback turtles,
including some cases in which the
plastic obstructed the passage of food
through the gut. The most commonly
identified items were plastic bags,
fishing lines, twine, and fragments of
mylar balloons. Ghost or derelict fishing
gear include discarded or lost nets, line,
and other gear. Ghost fishing gear can
drift in the ocean and fish unattended
for decades and kill numerous
individuals (Wilcox et al. 2013). The
main sources of ghost fishing gear are
gillnet, purse seine, and trawl fisheries
(Stelfox et al. 2016). Marine debris
affects leatherback turtles via ingestion
or entanglement and can reduce food
intake and digestive capacity, cause
distress and/or drowning, expose turtles
to contaminants, and in some cases
cause direct mortality (Mrosovsky et al.
2009; NMFS and USFWS 2013). In
terms of microplastics, all samples
analyzed from all species (including
leatherbacks) had microplastics evident
in their gastro-intestinal tracts (Duncan
et al. 2018). Given the increase of
pollution entering the marine
environment over the past 30 years or
approximately 5.2 to 19.3 million
tonnes per year (Lebreton et al. 2018),
we conclude that pollution is a threat to
the species.
Natural Disasters and Oceanographic
Regime Shifts
Leatherback turtles are susceptible to
the impacts of natural disasters and
oceanographic regime shifts as a result
of their nesting and foraging
preferences. Nesting usually occurs on
high-energy beaches that are inherently
unstable (Pritchard 1976) and which are
susceptible to natural erosion. The
primary factors influencing shoreline
suitability for nesting appear to be a lack
of abrasive substrate material, a deepwater approach to minimize energy
expenditure needed to reach nesting
sites, and proximity to oceanic currents
that can facilitate hatchling dispersal
(Eckert et al. 2012). Leatherback turtles
nest lower on the beach than other
species, exposing their nests to erosion
and inundation. Storm events, King
Tides, tsunamis, and hurricanes can
destroy or modify preferred nesting
beaches of some DPSs.
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Gelatinous prey have relatively low
energy content, requiring leatherback
turtles to consume large quantities to
meet metabolic demands (Heaslip et al.
2012; Jones et al. 2012). Leatherback
turtles likely maximize their caloric
intake by aligning their foraging
behavior to prey distribution
abundance. Foraging areas are generally
characterized by zones of upwelling,
including off the edges of continents,
where major currents converge, and in
deep-water eddies (Saba 2013). Some of
these areas experience oceanographic
regime shifts that alter water
temperature, downwelling, Ekman
upwelling, sea surface height,
chlorophyll-a concentration, and
mesoscale eddies (Bailey et al. 2013;
Benson et al. 2011). These shifts alter
prey availability, and thus productivity
parameters (e.g., remigration rates,
clutch size, and clutch frequency), for
leatherback turtles. Some DPSs are not
affected by such shifts because they
have access to diverse foraging areas,
such as: coastal and pelagic waters;
subtropical, temperate, and boreal
waters; and ephemeral eddies (Neeman
et al. 2015). Such flexibility allows the
leatherback turtle to consume large
amounts of prey at various locations
throughout the year.
We conclude that natural disasters
and oceanographic regime shifts are
threats to the species, affecting some but
not all populations, depending on the
location of nesting and foraging areas.
These threats reduce productivity by
reducing nesting, nesting habitat, and
nest and hatching success.
Climate Change
Climate change is a threat that affects
leatherback turtles of all life stages and
within all DPSs. A warming climate and
rising sea levels can impact leatherback
turtles through changes in beach
morphology, increased sand
temperatures leading to a greater
incidence of lethal incubation
temperatures, changes in hatchling sex
ratios, and the loss of nests or nesting
habitat due to beach erosion (Benson et
al. 2013).
Impacts from climate change,
especially due to global warming, are
already being observed and are likely to
become more apparent in future years
(IPCC 2007a). In its Fifth Assessment
Report, the IPCC (2014) stated that the
globally averaged combined land and
ocean surface temperature data has
shown a warming of 0.85 °C from 1880
to 2012. The mean rate of globally
averaged sea level rise was 1.7
millimeters annually between 1901 and
2010, 2.0 millimeters annually between
1971 and 2010, and 3.2 millimeters
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annually between 1993 and 2010.
Climate model projections exhibit a
wide range of plausible scenarios for
both temperature and precipitation over
the next several decades. The global
mean surface temperature change for the
period 2016 to 2035 relative to 1986 to
2005 will likely be in the range of
0.3 ° to 0.7 °C (medium confidence;
IPCC 2014). The global ocean
temperature will continue to warm, and
increases in seasonal and annual mean
surface temperatures are expected to be
larger in the tropics and Northern
Hemisphere subtropics (i.e., where
leatherback turtles nest; IPCC 2014).
Under Representative Concentration
Pathway 8.5, the change in global mean
sea level rise for the mid- and late 21st
century relative to the reference period
of 1986 to 2005 is projected to be 0.30
meters higher from 2046 to 2065 and
0.63 meters higher from 2081 to 2100,
with a rate of sea level rise during 2081
to 2100 of 8 to 16 millimeters annually
(medium confidence; IPCC 2014).
For all sea turtles, including
leatherback turtles, a warming climate
and rising sea levels are likely to result
in changes in beach morphology,
increased sand temperatures leading to
a greater incidence of lethal incubation
temperatures, changes in hatchling sex
ratios, and the loss of nests and nesting
habitat due to beach erosion (Benson et
al. 2015; Hamann et al. 2013).
Leatherback turtles are most likely to be
affected by climate change at nesting
beaches due to warming temperatures,
sea level rise, and storm events and due
to oceanic changes that are likely to
alter foraging and migration. Warming
temperatures and increased
precipitation at nesting beaches affect
reproductive output including hatching
success, hatchling emergence rate, and
hatchling sex ratios (e.g., Hawkes et al.
2009). Sea level rise results in a
reduction or shift in available nesting
beach habitat, an increased risk of
erosion and nest inundation (e.g., Boyes
et al. 2010), and reduced nest success
(Fish et al. 2005; Fuentes et al. 2010;
Fonseca et al. 2013). Increased
frequency and severity of storm events
impact nests and nesting habitat, thus
reducing nesting and hatching success
(e.g., Van Houtan and Bass 2007;
Fuentes and Abbs 2010). Changes in
productivity affect the abundance and
distribution of forage species, resulting
in changes in the foraging behavior and
distribution of leatherback turtles (e.g.,
Saba et al. 2008, 2012) as well as
changes in leatherback fitness and
growth. Changes in water temperature
lead to a shift in range and changes in
phenology (timing of nesting seasons,
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timing of migrations) and different
threat exposure (e.g., Saba et al. 2008,
2012).
Increasing sand temperatures will
alter the thermal regime of incubating
nests, resulting in altered sex ratios and
reduced hatching output (Hawkes et al.
2009). Leatherback turtles exhibit
temperature-dependent sex
determination (reviewed by Binckley
and Spotila 2015), whereby phenotypic
sex is determined by temperatures
experienced during the thermosensitive
period of egg incubation. A 1:1 sex ratio
is produced when this pivotal
temperature lies between 29.2 and 30.4
°C for leatherback turtles in Malaysia,
29.2 and 29.8 °C in French Guiana/
Suriname, and 29.2 and 29.5 °C in
Pacific Costa Rica (Binckley and Spotila
2015). Warmer temperatures produce
more female embryos (Mrosovsky et al.
1984; Hawkes et al. 2007), but
temperatures over 32 °C are likely to
result in death. As temperatures
continue to increase, emergence rates
decrease (Santidria´n Tomillo et al.
2015), removing any advantage of
increased female production. Santidria´n
Tomillo et al. (2015) conclude that
leatherback turtles may not survive if
temperatures rise as projected by
current climate change models.
Increases in precipitation might
temporarily reduce the temperatures at
some nesting beaches thereby mitigating
some impacts relative to increasing sand
temperatures.
Beach erosion and nest inundation
already threaten leatherback nesting
habitat globally. Sea level rise is likely
to increase the number of nests lost to
erosion and inundation. Such loss of
nests is especially problematic in areas
prone to storm events, which are likely
to increase in intensity and duration,
and in areas where coastal development
impedes natural shoreline migration.
Climate change is also likely to alter
the productivity in some marine
environments, which could affect
leatherback prey availability. With
reports on the increasing incidence of
jellyfish blooms in some locations, there
is the perception that jellyfish
abundance is increasing globally
(Condon et al. 2012), which could result
in more prey for leatherback turtles
(Hawkes et al. 2009). However, after
analyzing all available long-term
datasets on jellyfish abundance, Condon
et al. (2012) found that there is no
robust evidence for a global increase in
jellyfish. Rather, jellyfish populations
undergo larger, worldwide oscillations
with an approximate 20-year periodicity
(Condon et al. 2012). Additional
monitoring is needed to determine
whether the weak linear trend in
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jellyfish abundance since 1970
represents an actual increase or is a
phase of an oscillation (Condon et al.
2012). Therefore, the effects of climate
change on productivity are uncertain.
As described in prior sections with
respect to each individual population,
some impacts from climate change have
already been observed. At several
nesting beaches, increased erosion
occurs, and sex ratios are severely
skewed toward females. Beach erosion
reduces productivity. Although the
skew toward females could increase
productivity in the short-term, it is often
correlated with low hatching success.
For these reasons, climate change is a
threat to the species.
Conservation Efforts
The ESA requires the Services to
make their listing determinations solely
on the basis of the best scientific and
commercial data available, after
conducting a status review, and after
taking into account those efforts, if any,
being made by any State or foreign
nation to protect the species, whether by
predatory control, protection of habitat
and food supply, or other conservation
practices, within any area under its
jurisdiction, or on the high seas (16
U.S.C. 1533 (b)(1)(A)). In addition, the
Services published a policy for the
evaluation of domestic conservation
efforts which have yet to be
implemented or to show effectiveness
(68 FR 15100; March 28, 2003). We did
not identify any conservation efforts
that required such evaluation for
leatherbacks (i.e., the conservation
efforts reviewed are international in
nature or have already been
implemented to a sufficient degree that
they have a track record of being
effective or not being effective). Several
conservation efforts have been
previously discussed in prior sections
evaluating regulatory mechanisms with
respect to each DPS. Therefore, the list
below describes only those conservation
efforts that have not been previously
discussed and that apply generally to
the leatherback species rather than
being clearly associated with a
particular population. We considered
these efforts prior to making our listing
determination. After reviewing these
efforts, we concluded that they have
been somewhat effective, in that they
have prevented this endangered species
from going extinct. However, these
efforts have not reduced the threats to
a level at which protections under the
ESA are no longer necessary.
African Convention on the
Conservation of Nature and Natural
Resources (Algiers Convention):
Adopted in September 1968, the
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contracted states were ‘‘to undertake to
adopt the measures necessary to ensure
conservation, utilization and
development of soil, water, floral and
faunal resources in accordance with
scientific principles and with due
regard to the best interests of the
people.’’ The Algiers Convention
recently has undergone revision, and its
objectives are to enhance environmental
protection, foster conservation and
sustainable use of natural resources, and
harmonize and coordinate policies in
these fields with a view to achieving
ecologically rational, economically
sound, and socially acceptable
development policies and programs.
Additional information is available at
https://www.unep.ch/regionalseas/legal/
afr.htm.
Atlantic Sea Turtle Network (ASO):
Created in 2003 to foster greater
collaboration in southern Brazil,
Uruguay, and Argentina for the
protection of sea turtles and their
habitats. ASO represents dozens of local
and regional NGOs and government
agencies as well as hundreds of
community members. ASO and its
partners have significantly advanced
policies to protect sea turtles from
fisheries interactions, which is one of
the most severe threats in the region.
Brazil plays a major role in South
American (and global) sea turtle
conservation and research, and it serves
as an example to other countries. Projeto
TAMAR, a partnership of the Centro
TAMAR/ICMBio, government agencies,
and Fundaca˜o Pro´ TAMAR, has been
active since 1980. Today, the group
carries out sea turtle research and
conservation from 22 stations on the
coast and the offshore islands of Brazil.
Another NGO based in the southern
Brazilian state of Rio Grande do Sul,
called NEMA has been collecting
systematic sea turtle stranding data
since 1990. Those data have been
instrumental to conservation efforts in
Brazil and have shown that southern
Brazil has the highest stranding rates for
loggerheads in the western Atlantic
Ocean.
Association of Southeast Asian
Nations (The ASEAN) Ministers on
Agriculture and Forestry (AMAF): A
Memorandum of Understanding (MoU)
on ASEAN sea turtle conservation was
created in 1999. From this, a Sea Turtle
Conservation and Protection Program
and Work plan has developed; research
and monitoring activities have also been
produced regionally (Kadir 2000). The
objectives of this Memorandum of
Understanding, initiated by ASEAN, are
to promote the protection, conservation,
replenishing, and recovery of sea turtles
and their habitats based on the best
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available scientific evidence, taking into
account the environmental, socioeconomic and cultural characteristics of
the Parties. It currently has nine
signatory states in the South East Asian
Region (https://document.seafdec.or.th/
projects/2012/seaturtles.php).
Andaman and Nicobar Island
Environmental Team (ANET): A
division of the Centre for Herpetology/
Madras Crocodile Bank Trust has been
conducting surveys and monitoring
since 1991. Over the last few years,
conservation and monitoring of sea
turtles in these islands has been carried
by Dakshin Foundation and Indian
Institute of Science in collaboration
with ANET, centered around a
leatherback monitoring program on
Little Andaman Island. A multiinstitution stakeholder platform for
marine conservation, including
government and non- governmental
agencies, was established by these
groups to facilitate the conservation of
marine turtles and other endangered
species (Tripathy et al. 2012). The Trust,
along with the Wildlife Institute of India
and Ministry of Environment and
Forests, produced a series of manuals on
sea turtle conservation, management
and research to help forest officers,
conservationists, NGOs and wildlife
enthusiasts conduct sea turtle
conservation and research programs
(ANET, 2003 as cited in Shanker and
Andrews 2004). A consolidated manual
has been produced to achieve these
goals by Dakshin Foundation and the
Trust (Tripathy et al. 2012).
Central American Regional Network:
This collaborative effort created the
national sea turtle network in each
country of the region, as well as the
development of first hand tools, such as
a regional diagnosis, a 10-year strategic
plan, a manual of best practices, and
four regional training and information
workshops for people in the region (e.g.,
Chaco´n and Arauz, 2001). This initiative
is managed by stakeholders in various
sectors (private, non-governmental and
governmental) across the region.
Convention on the Conservation of
Migratory Species of Wild Animals
(CMS): This Convention, also known as
the Bonn Convention or CMS, is an
international treaty that focuses on the
conservation of migratory species and
their habitats. As of December 2018, the
Convention had 127 Parties, including
Parties from Africa, Central and South
America, Asia, Europe, and Oceania.
While the Convention has successfully
brought together about half the
countries of the world with a direct
interest in sea turtles, it has yet to
realize its full potential (Hykle 2002). Its
membership does not include a number
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of key countries, including Canada,
China, Indonesia, Japan, Mexico, Oman,
and the United States. Under the CMS,
two Memoranda of Understanding
(MOUs) apply to leatherback turtles:
The MOU concerning Conservation
Measures for Marine Turtles of the
Atlantic Coast of Africa and the MOU
on the Conservation and Management of
Marine Turtles and their Habitats of the
Indian Ocean and South-East Asia.
Additional information is available at
https://www.cms.int.
Convention on Biological Diversity
(CBD): The primary objectives of this
international treaty are: (1) The
conservation of biological diversity, (2)
the sustainable use of its components,
and (3) the fair and equitable sharing of
the benefits arising out of the utilization
of genetic resources. This Convention
has been in force since 1993 and had
193 Parties as of March 2013. While the
Convention provides a framework
within which are broad conservation
objectives, it does not specifically
address sea turtle conservation (Hykle
2002). Additional information is
available at https://www.cbd.int.
Convention on International Trade in
Endangered Species of Wild Fauna and
Flora (CITES): Known as CITES, this
Convention was designed to regulate
international trade in a wide range of
wild animals and plants. CITES was
implemented in 1975 and currently has
183 Parties. Although CITES has been
effective at minimizing the international
trade of sea turtle products, it does not
limit legal harvest within countries, nor
does it regulate intra-country commerce
of sea turtle products (Hykle, 2002). The
leatherback turtle is included (since
1977) in CITES Appendix I, which bans
trade, including individuals and
products, except as permitted for
exceptional circumstances, not to
include commercial purposes (Lyster
1985). Additional information is
available at https://www.cites.org.
Convention on the Conservation of
European Wildlife and Natural Habitats:
Also known as the Bern Convention, the
goals of this instrument are to conserve
wild flora and fauna and their natural
habitats, especially those species and
habitats whose conservation requires
the cooperation of several States, and to
promote such cooperation. The
Convention was enacted in 1982 and
currently includes 51 European and
African States and the European Union.
Additional information is available at
https://www.coe.int/t/dg4/
cultureheritage/nature/bern/default_
en.asp.
Convention for the Co-operation in
the Protection and Development of the
Marine and Coastal Environment of the
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West and Central African Region
(Abidjan Convention): The Abidjan
Convention covers the marine
environment, coastal zones, and related
inland waters from Mauritania to
Namibia. The Abidjan Convention
countries are Angola, Benin, Cameroon,
Cape Verde, Congo, Cote d’Ivoire,
Democratic Republic of Congo,
Equatorial Guinea, Gabon, Gambia,
Ghana, Guinea, Guinea-Bissau, Liberia,
Mauritania, Namibia, Nigeria, Sao Tome
and Principe, Senegal, Sierra Leone, and
Togo. The Abidjan Convention is an
agreement for the protection and
management of the marine and coastal
areas that highlights sources of
pollution, including pollution from
ships, dumping, land-based sources,
exploration and exploitation of the seabed, and pollution from or through the
atmosphere. The Convention also
identifies where co-operative
environmental management efforts are
needed. These areas of concern include
coastal erosion, specially protected
areas, combating pollution in cases of
emergency and environmental impact
assessment.
Convention for the Protection
Management and Development of the
Marine and Coastal Environment of the
Eastern African Region (Nairobi
Convention): The Nairobi Convention
was signed in 1985 and came into force
in 1996. This instrument ‘‘provides a
mechanism for regional cooperation,
coordination and collaborative actions,
and enables the Contracting Parties to
harness resources and expertise from a
wide range of stakeholders and interest
groups towards solving interlinked
problems of the coastal and marine
environment.’’ Parties are responsible
for ‘‘the conservation and wise
management of the sea turtle
populations frequenting their waters
and shores [and] agree to work closely
together to improve the conservation
status of the sea turtles and the habitats
upon which they depend.’’ The Western
Indian Ocean-Marine Turtle Task Force,
which was created under the Nairobi
Convention and the IOSEA, plays a role
in sea turtle conservation. This is a
technical, non-political working group
comprised of specialists from eleven
countries: Comoros, France (La
Re´union), Kenya, Madagascar,
Mauritius, Mozambique, Seychelles,
Somalia, South Africa, United Kingdom
and Tanzania, as well as representatives
from inter-governmental organizations,
academic, and non-governmental
organizations within the region.
Additional information is available at
https://www.unep.org/
NairobiConvention.
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Convention for the Protection of the
Marine Environment of the North-East
Atlantic: Also called the OSPAR
Convention, this 1992 instrument
combines and updates the 1972 Oslo
Convention against dumping waste in
the marine environment and the 1974
Paris Convention addressing marine
pollution stemming from land-based
sources. The convention is managed by
the OSPAR Commission, which is
comprised of representatives from 15
signatory nations (Belgium, Denmark,
Finland, France, Germany, Iceland,
Ireland, Luxembourg, The Netherlands,
Norway, Portugal, Spain, Sweden,
Switzerland, and United Kingdom), as
well as the European Commission,
representing the European Community.
The mission of the OSPAR Convention
‘‘. . . is to conserve marine ecosystems
and safeguard human health in the
North-East Atlantic by preventing and
eliminating pollution; by protecting the
marine environment from the adverse
effects of human activities; and by
contributing to the sustainable use of
the seas.’’ Leatherback turtles are
included on the OSPAR List of
Threatened and/or Declining Species
and Habitats, used by the OSPAR
Commission for setting priorities for
work on the conservation and protection
of marine biodiversity. Additional
information is available at https://
www.ospar.org.
Convention for the Protection and
Development of the Marine
Environment of the Wider Caribbean
Region: Also called the Cartagena
Convention, this instrument that
benefits turtles of the Northwest
Atlantic leatherback DPS, has been in
place since 1986 and currently has 38
member states and territories. Under
this Convention, the component that
relates to leatherback turtles is the
Protocol Concerning Specially Protected
Areas and Wildlife (SPAW) that has
been in place since 2000. The goals are
to encourage Parties ‘‘to take all
appropriate measures to protect and
preserve rare or fragile ecosystems, as
well as the habitat of depleted,
threatened or endangered species, in the
Convention area.’’ The SPAW protocol
has partnered with WIDECAST to
develop a program of work on sea turtle
conservation, which has helped many of
the Caribbean nations to identify and
prioritize their conservation actions
through Sea Turtle Recovery Action
Plans. Each recovery action plan
summarizes the known distribution of
sea turtles, discusses major causes of
mortality, evaluates the effectiveness of
existing conservation laws, and
prioritizes implementing measures for
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stock recovery. The objective of the
recovery action plan series is not only
to assist Caribbean governments in the
discharge of their obligations under the
SPAW Protocol, but also to promote a
regional capability to implement
science-based sea turtle management
and conservation programs. Additional
information is available at https://
www.cep.unep.org/about-cep/spaw.
Convention on Nature Protection and
Wildlife Preservation in the Western
Hemisphere (Washington or Western
Hemisphere Convention): Elements of
the Convention include the protection
of species from human-induced
extinction, the establishment of
protected areas, the regulation of
international trade in wildlife, special
measures for migratory birds and
stressing the need for co-operation in
scientific research and other fields are
all elements of wildlife conservation.
Additional information is available at
https://www.oas.org/juridico/english/
treaties/c-8.html.
Convention for the Protection of the
Marine Environment and Coastal Area
of the South-East Pacific (Lima
Convention): This Convention’s
signatories include all countries along
the Pacific Rim of South America from
Panama to Chile. Among other resource
management components, this
Convention established protocol for the
conservation and management of
protected marine resources. Stemming
from this Convention is the Commision
Permanente del Pacifico Sur (CPPS) that
has developed a Marine Turtle Action
Plan for the Southeast Pacific that
outlines a strategy for protecting and
recovering marine turtles in this region.
Convention for the Protection of the
Natural Resources and Environment of
the South Pacific Region (Noumea
Convention): This Convention has been
in force since 1990 and currently
includes 26 Parties. The purpose of the
Convention is to protect the marine
environment and coastal zones of the
South-East Pacific within the 200-mile
area of maritime sovereignty and
jurisdiction of the Parties and, beyond
that area, the high seas up to a distance
within which pollution of the high seas
may affect that area. Additional
information is available at https://
www.unep.org/regionalseas/
programmes/nonunep/pacific/
instruments/default.asp.
Convention Concerning the Protection
of the World Cultural and Natural
Heritage (World Heritage Convention):
The World Heritage Convention was
signed in 1972 and, as of November
2007, 185 states were parties to the
Convention. The instrument requires
parties to take effective and active
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measures to protect and conserve
habitat of threatened species of animals
and plants of scientific or aesthetic
value. The World Heritage Convention
currently includes 31 marine sites.
Additional information is available at
https://whc.unesco.org/en/
conventiontext.
Convention for the Conservation and
Management of Highly Migratory Fish
Stocks in the Western and Central
Pacific Ocean (WCPF Convention): The
convention entered into force on 19
June 2004. The WCPF Convention
draws on many of the provisions of the
UN Fish Stocks Agreement [UNFSA]
while, at the same time, reflecting the
special political, socio-economic,
geographical and environmental
characteristics of the western and
central Pacific Ocean (WCPO) region.
The WCPFC Convention seeks to
address problems in the management of
high seas fisheries resulting from
unregulated fishing, over-capitalization,
excessive fleet capacity, vessel reflagging to escape controls,
insufficiently selective gear, unreliable
databases and insufficient multilateral
cooperation in respect to conservation
and management of highly migratory
fish stocks.
Convention for the Prohibition of
Fishing with Long Driftnets in the South
Pacific: This regional convention, also
known as the Wellington Convention,
was adopted in 1989 in Wellington,
New Zealand, and entered into force in
1991. The objective of the Convention is
‘‘to restrict and prohibit the use of drift
nets in the South Pacific region in order
to conserve marine living resources.’’
Additional information is available at
https://www.mfat.govt.nz/Treaties-andInternational-Law/01-Treaties-forwhich-NZ-is-Depositary/0-Prohibitionof-Fishing.php.
Eastern Pacific Leatherback Network:
Also known as La Red de la Tortuga
Lau´d del Oce´ano Pacifico (Lau´d OPO)
(www.savepacificleatherbacks.org) was
formed to address the critical need for
regional coordination of East Pacific
leatherback conservation actions
necessary to track conservation
priorities and progress at the population
level. Led by Fauna & Flora
International, this network has brought
together conservationists, researchers,
practitioners and government
representatives from 22 institutions
across nine East Pacific countries with
varying priorities, capacities and
historical experiences in leatherback
research and conservation to contribute
to shared activities, projects, and goals.
Through these efforts, Lau´d now has
mutually-agreed upon mechanisms for
sharing information and data, as well as
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standardized protocols for nesting beach
monitoring and bycatch assessments/
fishing practices.
The Eastern Tropical Pacific Marine
Corridor (CMAR) is a regional and crossborder initiative for the conservation
and sustainable use of the region’s
marine and coastal resources. Its
objective is to sustainably manage
biodiversity through ecosystem based
management and the development of
regional intergovernmental strategies
with support of non-governmental
organizations and international
cooperation agencies.
United Nations’ Food and
Agricultural Organization (FAO)
Technical Consultation on Sea TurtleFishery Interactions: While not a true
international instrument for
conservation, the 2004 FAO of the UN’s
technical consultation on sea turtlefishery interactions was groundbreaking
in that it solidified the commitment of
the lead UN agency for fisheries to
reduce sea turtle bycatch in marine
fisheries operations. Recommendations
from the technical consultation were
endorsed by the FAO Committee on
Fisheries (COFI) and called for the
immediate implementation by member
nations and Regional Fishery
Management Organizations (RFMOs) of
guidelines to reduce sea turtle mortality
in fishing operations, developed as part
of the technical consultation. Currently,
all five of the tuna RFMOs call on their
members and cooperating non-members
to adhere to the 2009 FAO ‘‘Guidelines
to Reduce Sea Turtle Mortality in
Fishing Operations,’’ which describes
all the gear types sea turtles could
interact with and the latest mitigation
options. The Western and Central
Pacific Fisheries Commission (https://
www.wcpfc.int) has the most protective
measures (CMM 2008–03), which follow
the FAO guidelines and ensure safe
handling of all captured sea turtles.
Fisheries deploying purse seines, to the
extent practicable, must avoid
encircling sea turtles and release
entangled turtles from fish aggregating
devices. Longline fishermen must carry
line cutters and use dehookers to release
sea turtles caught on a line. Longliners
must either use large circle hooks,
whole finfish bait, or mitigation
measures approved by the Scientific
Committee and the Technical and
Compliance Committee.
Inter-American Tropical Tuna
Convention (IATTC) has enacted a
resolution to mitigate the impact of tuna
fishing vessels on sea turtles by
reducing bycatch, injury, and mortality
of sea turtles. The IATTC has also
developed a memorandum of
understanding with the IAC. For more
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information, see https://www.iattc.org/
PDFFiles/Resolutions/IATTC/_English/
C-07-03-Active_Sea%20turtles.pdf.
The International Commission for the
Conservation of Atlantic Tunas (ICCAT)
has adopted a resolution for the
reduction of sea turtle mortality
(Resolution 03–11), encouraging States
to submit data on sea turtle interactions,
release sea turtles alive wherever
possible, and conduct research on
mitigation measures. It calls for
implementing the FAO Guidelines for
sea turtles, avoiding encirclement of sea
turtles by purse seiners, safely handling
and releasing sea turtles, and reporting
on interactions. The Commission does
not have any specific gear requirements
applicable to longline fisheries. ICCAT
is currently undertaking an ecological
risk assessment to better understand the
impact of its fisheries on sea turtle
populations. For more information see
https://www.iattc.org/. Other
international fisheries organizations that
may influence leatherback turtle
recovery include the Southeast Atlantic
Fisheries Organization (https://
www.seafo.org) and the North Atlantic
Fisheries Organization (https://nafo.int).
These organizations regulate trawl
fisheries in their respective Convention
areas. Given that sea turtles can be
incidentally captured in these fisheries,
both organizations have sea turtle
resolutions calling on their Parties to
implement the FAO Guidelines on sea
turtles as well as to report data on sea
turtle interactions.
The Indian Ocean Tuna Commission
(IOTC) is playing an increased role in
turtle conservation. Resolution 05/08,
superseded by Resolution 09/06 on Sea
Turtles, sets out reporting requirements
related to interactions with sea turtles
and accordingly provides an executive
summary per species for adoption at the
Working Party on Ecosystem and Bycatch and then subsequently at the
Scientific Committee. In 2011, IOTC
developed a ‘‘Sea Turtle Identification
Card’’ to be distributed to all longliners
operating in the Indian Ocean
(www.iotc.com). In 2012, the Indian
Ocean Tuna Commission (IOTC) began
requiring its 31 contracting Parties to
report sea turtle bycatch and to use safe
handling and release techniques for sea
turtles on longline vessels.
Indian Ocean—South-East Asian
Marine Turtle Memorandum of
Understanding (IOSEA): Under the
auspices of the Convention of Migratory
Species, the IOSEA memorandum of
understanding provides a mechanism
for States of the Indian Ocean and
South-East Asian region, as well as
other concerned States, to work together
to conserve and replenish depleted
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marine turtle populations. This
collaboration is achieved through the
collective implementation of an
associated Conservation and
Management Plan. Currently, there are
33 Signatory States. The United States
became a signatory in 2001. The IOSEA
has an active sub-regional group for the
Western Indian Ocean, which has
improved collaboration amongst sea
turtle conservationists in the region.
Further, the IOSEA website provides
reference materials, satellite tracks, online reporting of compliance with the
Convention, and information on all
international mechanisms currently in
place for the conservation of sea turtles.
Finally, at the 2012 Sixth Signatory of
States meeting in Bangkok, Thailand,
the Signatory States agreed to
procedures to establish a network of
sites of importance for sea turtles in the
IOSEA region (https://
www.ioseaturtles.org).
Inter-American Convention for the
Protection and Conservation of Sea
Turtles (IAC): This Convention is the
only legally binding international treaty
dedicated exclusively to sea turtles and
sets standards for the conservation of
these endangered animals and their
habitats with a large emphasis on
bycatch reduction. The Convention area
is the Pacific and the Atlantic waters of
the Americas. Currently, there are 15
Parties. The United States became a
Party in 1999. The IAC has worked to
adopt fisheries bycatch resolutions,
carried out workshops on Caribbean sea
turtle conservation, and established
collaboration with other agreements
such as the Convention for the
Protection and Development of the
Marine Environment of the Wider
Caribbean Region and the International
Commission for the Conservation of
Atlantic Tunas. Additional information
is available at https://
www.iacseaturtle.org.
International Convention for the
Prevention of Pollution from Ships
(MARPOL): The MARPOL Convention
is a combination of two treaties adopted
in 1973 and 1978 to prevent pollution
of the marine environment by ships
from operational or accidental causes.
The 1973 treaty covered pollution by
oil, chemicals, and harmful substances
in packaged form, sewage and garbage.
The 1978 MARPOL Protocol was
adopted at a Conference on Tanker
Safety and Pollution Prevention which
included standards for tanker design
and operation. The 1978 Protocol
incorporated the 1973 Convention as it
had not yet been in force and is known
as the International Convention for the
Prevention of Marine Pollution from
Ships, 1973, as modified by the Protocol
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of 1978 relating thereto (MARPOL 73/
78). The 1978 Convention went into
force in 1983 (Annexes I and II). The
Convention includes regulations aimed
at preventing and minimizing accidental
and routine operations pollution from
ships. Amendments passed since have
updated the convention.
International Union for Conservation
of Nature (IUCN): The IUCN Species
Programme assesses the conservation
status of species on a global scale. This
assessment provides objective, scientific
information on the current status of
threatened species. The IUCN Red List
of Threatened Species provides
taxonomic, conservation status and
distribution information on plants and
animals that have been globally
evaluated using the IUCN Red List
Categories and Criteria. This system is
designed to determine the relative risk
of extinction, and the main purpose of
the IUCN Red List is to catalogue and
highlight those plants and animals that
are facing a higher risk of global
extinction (i.e., those listed as Critically
Endangered, Endangered and
Vulnerable). Additional information is
available at https://www.iucnRed
List.org/about.
Marine Turtle Conservation Act
(MTCA): The MTCA is a key element of
sea turtle protection in the United States
and internationally. This Act authorizes
a dedicated fund to support marine
turtle conservation projects in foreign
countries, with emphasis on protecting
nesting populations and nesting habitat.
Additional information is available at
https://www.fws.gov/international/
wildlife-without-borders/marine-turtleconservation-fund.html.
Memorandum of Agreement between
the Government of the Republic of the
Philippines and the Government of
Malaysia on the Establishment of the
Turtle Island Heritage Protected Area:
Through a bilateral agreement, the
Governments of the Philippines and
Malaysia established The Turtle Island
Heritage Protected Area (TIHPA), made
up of nine islands (6 in the Philippines
and 3 in Malaysia). The following
priority activities were identified:
management-oriented research, the
establishment of a centralized database
and information network, appropriate
information awareness programs, a
marine turtle resource management and
protection program, and an appropriate
ecotourism program (Bache and Frazier
2006).
Memorandum of Understanding of a
Tri-National Partnership between the
Government of the Republic of
Indonesia, the Independent State of
Papua New Guinea and the Government
of Solomon Islands: This agreement
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promotes the conservation and
management of Western Pacific
leatherback turtles at nesting sites,
feeding areas and migratory routes in
Indonesia, Papua New Guinea and
Solomon Islands. This is done through
the systematic exchange of information
and data on research, population and
migratory routes monitoring, nesting
sites and feeding areas management
activities for Western Pacific
leatherback turtles and by enhancing
public awareness of the importance of
conserving these turtles and their
critical habitats. https://
awsassets.wwf.or.id/downloads/mou_
trinationalpartneshipagreement_
clean.pdf.
Memorandum of Understanding
Concerning Conservation Measures for
Marine Turtles of the Atlantic Coast of
Africa (Abidjan Memorandum): This
MOU was concluded under the auspices
of the Convention on the Conservation
of Migratory Species of Wild Animals
(CMS) and became effective in 1999.
The MOU area covers 26 Range States
along the Atlantic coast of Africa
extending approximately 14,000 km
from Morocco to South Africa. The goal
of this MOU is to improve the
conservation status of marine turtles
along the Atlantic Coast of Africa. It
aims at safeguarding six marine turtle
species—including the leatherback
turtle—that are estimated to have
rapidly declined in numbers during
recent years due to excessive
exploitation (both direct and incidental)
and the degradation of essential
habitats. This includes the protection of
the life stages from hatchlings through
adults with particular attention paid to
the impacts of fishery bycatch and the
need to include local communities in
the development and implementation of
conservation activities. However,
despite this agreement, killing of adult
turtles and harvesting of eggs remains
rampant in many areas along the
Atlantic African coast. Additional
information is available at https://
www.cms.int/species/africa_turtle/
AFRICAturtle_bkgd.htm.
National Sea Turtle Conservation
Project in India: Launched in 1998 with
the aim of protecting Lepidochelys
olivacea, but it also has conservation
and protection strategies for all the other
turtle species nesting in the country.
This project was undertaken by the
Indian government to oversee: Surveys,
monitoring programs, fisheries
interactions, community and NGO
participation, awareness raising and
education, research support, and other
support for regional and international
co-operation and collaboration for sea
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turtles conservation (Choudhury et al.,
2001).
North American Agreement for
Environmental Cooperation: As
mandated by the 1994 North American
Agreement for Environmental
Cooperation, the Commission for
Environmental Cooperation (CEC)
encourages Canada, the United States,
and Mexico to adopt a continental
approach to the conservation of flora
and fauna. In 2003, this mandate was
strengthened as the three North
American countries launched the
Strategic Plan for North American
Cooperation in the Conservation of
Biodiversity. The North American
Conservation Action Plan (NACAP)
initiative began as an effort promoted by
the three countries, through the CEC, to
facilitate the conservation of marine and
terrestrial species of common concern.
In 2005, the CEC supported the
development of a NACAP for Pacific
leatherbacks by Canada, the United
States, and Mexico. Identified actions in
the plan addressed three main
objectives: (1) protection and
management of nesting beaches and
females; (2) mortality reduction from
bycatch throughout the Pacific Basin;
and (3) waste management, control of
pollution, and disposal of debris at sea.
Ramsar Convention on Wetlands: The
Convention on Wetlands, signed in
Ramsar, Iran, in 1971, is an
intergovernmental treaty, which
provides the framework for national
action and international cooperation for
the conservation and wise use of
wetlands and their resources. Currently,
there are 158 parties to the convention,
with 1,752 wetland sites, including
important marine turtle habitat.
Additional information is available at
https://www.ramsar.org.
Secretariat of the Pacific Regional
Environment Programme (SPREP):
SPREP’s turtle conservation program
seeks to improve knowledge about sea
turtles in the Pacific through an active
tagging program, as well as maintaining
a database to collate information about
sea turtle tags in the Pacific. SPREP
supports capacity building throughout
the central and southwest Pacific.
SPREP established an action plan for the
Pacific Islands (https://www.sprep.org/).
South-East Atlantic Fisheries
Organization (SEAFO): SEAFO manages
fisheries activities in the Southeast
Atlantic high seas area, excluding tunas
and billfish. SEAFO adopted Resolution
01/06, ‘‘to Reduce Sea Turtle Mortality
in Fishing Operations,’’ in 2006. The
Resolution requires Members to: (1)
Implement the FAO Guidelines; and (2)
establish on-board observer programs to
collect information on sea turtle
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interactions in SEAFO-managed
fisheries. This Resolution is not legally
binding. Additional information is
available at https://www.seafo.org.
South Atlantic Association: In the
southwest Atlantic, the South Atlantic
Association is a multinational group
that includes representatives from
Brazil, Uruguay, and Argentina and
meets bi-annually to share information
and develop regional action plans to
address threats including bycatch
(https://www.tortugasaso.org/). At the
national level, Brazil has developed a
national plan for sea turtle bycatch
reduction that was initiated in 2001
(Marcovaldi et al. 2002). This national
plan includes various activities to
mitigate bycatch, including time-area
restrictions of fisheries, use of bycatch
reduction devices, and working with
fishermen to successfully release livecaptured turtles. In Uruguay, all sea
turtles are protected from human
impacts, including fisheries bycatch, by
presidential decree (Decreto
Presidencial 144/98).
United Nations Convention on the
Law of the Sea (UNCLOS): To date, 155
countries, including most mainland
countries lining the western Pacific, and
the European Community have joined in
the convention. The United States has
signed the treaty and abides by some
provisions, but the Senate has not
ratified it. Aside from its provisions
defining ocean boundaries, the
convention establishes general
obligations for safeguarding the marine
environment through mandating
sustainable fishing practices and
protecting freedom of scientific research
on the high seas. Additional information
is available at https://www.un.org/Depts/
los/index.htm.
United Nations’ Food and
Agricultural Organization (FAO): The
FAO published guidelines for sea turtle
protection, entitled Technical
Consultation on Sea Turtle-Fishery
Interactions (FAO 2005). The UN 1995
Code of Conduct for Responsible
Fisheries (FAO 2004) provides
guidelines for the development and
implementation of national fisheries
policies, including gear modification
(e.g., circle hooks, fish bait, deeper sets,
and reduced soak time), new
technologies, and management of areas
where fishery and sea turtle interactions
are more severe. The guidelines stress
the need for mitigation measures, data
on all fisheries, fishing industry
involvement, and education for fishers,
observers, managers, and compliance
officers (FAO 2004).
United Nations Resolution 44/225 on
Large-Scale Pelagic Driftnet Fishing: In
1989, the UN called, in a unanimous
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resolution, for the elimination of all
high seas driftnets by 1992. Additional
information is available at https://
www.un.org/documents/ga/res/44/
a44r225.htm.
Although numerous conservation
efforts apply to the species, they do not
adequately reduce its risk of extinction
for the reasons discussed previously.
Extinction Risk Analysis
The best available information is
consistent with the species’ current
‘‘endangered’’ listing. The species
exhibits a global total index of nesting
female abundance of 32,060 females at
monitored beaches. This number is
lower than historical estimates of
nesting female abundance (n = 115,000,
Pritchard 1982; and n = 34,500, Spotila
1996), which did not include the large,
but then unknown, Gabon nesting
aggregation. Limited nesting female
abundance is a major source of concern
for most DPSs, whose small population
sizes place them in danger of stochastic
or catastrophic events that increase
extinction risk. The limited nesting
female abundance increases the
extinction risk of the species.
The species also exhibits declining
nesting trends for all but one of the
DPSs. With the exception of the DPS
with the smallest index of nesting
female abundance (i.e., SW Atlantic
DPS, with 27 nesting females), the DPSs
are declining at rates of 0.3 to 9.3
percent annually. Even low levels of
decline are a threat for DPSs with
limited nesting female abundance, and
nesting declines of approximately nine
percent (i.e., NW and SE Atlantic DPSs)
are unsustainable. Total declines of 97
and 99 percent have occurred within the
East Pacific and NE Indian DPSs,
respectively, since nesting was first
identified and quantified for these
populations. The declining trends in
nesting increase the extinction risk of
the species.
The species exhibits broad nesting
and foraging ranges. However,
metapopulation dynamics have likely
been reduced, with reductions in
abundance and the loss of some nesting
aggregations. The species also
demonstrates little genetic diversity,
relative to other sea turtle species.
Although the species demonstrates
some resilience to threats, overall we
find it to be at risk of extinction, due to
limited abundance and declining
nesting trends, which reflect the
cumulative impacts of threats that have
acted on the species in the past (and in
many cases continue to act on the
species).
Current threats continue to place the
species in danger of extinction. The
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primary threat to the species is bycatch
in commercial and artisanal, pelagic and
coastal, fisheries. Fisheries bycatch
reduces abundance by removing
individuals from the population.
Because several fisheries operate near
nesting beaches, productivity is also
reduced when nesting females are
prevented from returning to nesting
beaches. The harvest of eggs and turtles
is also a major threat to the species.
Illegal poaching occurs throughout the
range of the species, and harvest is legal
but poorly documented in some nations.
The loss and modification of nesting
habitat is another major threat, reducing
productivity and, in some instances,
abundance, when nesting females die as
a result of artificial lighting or
obstructions preventing them from
returning to sea. Predation results in the
loss of eggs and hatchlings, reducing
productivity of the species. Additional
threats that occur throughout the range
of the species include vessel strikes,
pollution, marine debris, oil and gas
exploration, and climate change.
Natural disasters and oceanographic
regime shifts are threats in some areas.
Though many regulatory mechanisms
are in place, they do not adequately
reduce the impact of these threats.
Based on our review of the best
available scientific and commercial
data, we find nothing that is
inconsistent with the leatherback
species’ current listing as an endangered
species. In sum, the best available
information is consistent with the
current listing status of the leatherback
sea turtle as an endangered species
throughout its range. The threatened
species definition does not apply
because the species is currently in
danger of extinction (i.e., at present),
rather than on a trajectory to become so
within the foreseeable future.
Final Determination
The Services determined that the best
available scientific and commercial
information would support recognizing
seven populations as DPSs (including
the NW Atlantic) because they meet the
discreteness and significance criteria for
DPSs. However, we found that—even
were they to be recognized and listed
separately—all DPSs meet the definition
of an endangered species because they
are in danger of extinction throughout
all of their ranges. The leatherback turtle
is currently listed throughout its range
as an endangered species. Replacing this
listing with seven endangered DPSs
would not be consistent with
Congressional guidance to use the
authority to list DPSs ‘‘sparingly’’ while
encouraging the conservation of genetic
diversity (see Senate Report 151, 96th
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Congress, 1st Session). Such guidance
clearly indicates that the Services have
some discretion to determine whether or
not to recognize DPSs that would
require disaggregating an existing listing
even where those populations can be
shown to meet the discreteness and
significance tests of the DPS Policy.
After determining that all seven
populations would have the same status
as the overall species, we next
considered whether there was any
reason to nevertheless replace the global
(range-wide) listing with individual
listings for the seven DPSs. We
conclude that disaggregating the global
listing is not warranted. It would be
inconsistent with Congressional
guidance and run counter to the
conservation purposes of the Act to
disaggregate the current listing into
DPSs, because those DPSs would have
the same listing status as the whole
currently. Disaggregating this listing
would bring about significant
complications and possible public
confusion without any meaningful
corresponding conservation benefit.
Replacing the range-wide listing with
seven DPSs having the same status
would not provide leatherback turtles
with an overriding conservation benefit,
as all members are currently protected
to the fullest extent under the ESA as an
endangered species. Section 7
consultations already consider the
effects of an action on individuals and
populations to determine whether a
Federal agency has insured that its
action is not likely to jeopardize the
continued existence of the species. Even
if the species were disaggregated into
DPSs, this change would not be
expected to result in different
substantive outcomes in consultations.
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In addition, focused conservation efforts
have been, and will continue to be,
applied at scales smaller than the
species-level. For example, FWS’
Marine Turtle Conservation Fund
provides funding to partners in foreign
nations to protect leatherback turtles
and their nesting habitats; projects
include efforts to monitor and protect
leatherback turtles in Indonesia and
Gabon (https://www.fws.gov/
international/wildlife-without-borders/
marine-turtle-conservation-fund.html).
Similarly, Pacific leatherback turtles are
highlighted under NMFS’ Species in the
Spotlight: Survive to Thrive initiative,
which directs attention and resources to
highly-at-risk species (https://
www.fisheries.noaa.gov/topic/
endangered-speciesconservation#species-in-the-spotlight).
For these reasons, the Services have
determined that replacing the existing
global listing with separate listings for
individual DPSs is not warranted.
Although the best available data
indicates that the populations meet the
criteria for significance and
discreteness, we find that it would not
further the purposes of the Act to
recognize and list seven DPSs separately
as endangered under the ESA. The
current global listing of the species
remains in effect.
We conclude that the petitioned
actions, to identify the NW Atlantic
population as a DPS and list it as a
threatened species under the ESA, are
not warranted. This is a final action,
and, therefore, we are not soliciting
public comments.
Peer Review
In December 2004, the Office of
Management and Budget (OMB) issued
PO 00000
Frm 00091
Fmt 4701
Sfmt 9990
48421
a Final Information Quality Bulletin for
Peer Review, establishing minimum
peer review standards, a transparent
process for public disclosure of peer
review planning, and opportunities for
public participation. The OMB Bulletin,
implemented under the Information
Quality Act (Pub. L. 106–554), is
intended to enhance the quality and
credibility of the Federal government’s
scientific information and applies to
influential or highly influential
scientific information disseminated on
or after June 16, 2005. To satisfy our
requirements under the OMB Bulletin,
we obtained independent peer review of
the Status Review Report by
independent scientists with expertise in
leatherback turtle biology, endangered
species listing policy, and related fields.
All peer reviewer comments were
addressed prior to the publication of the
Status Review Report and this finding.
References Cited
A complete list of references is
available upon request to the NMFS
Office of Protected Resources (see
ADDRESSES).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Samuel D. Rauch III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
Aurelia Skipwith,
Director, U.S. Fish and Wildlife Service.
[FR Doc. 2020–16277 Filed 8–7–20; 8:45 am]
BILLING CODE 3510–22–P
E:\FR\FM\10AUR2.SGM
10AUR2
Agencies
[Federal Register Volume 85, Number 154 (Monday, August 10, 2020)]
[Rules and Regulations]
[Pages 48332-48421]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2020-16277]
[[Page 48331]]
Vol. 85
Monday,
No. 154
August 10, 2020
Part II
Department of the Interior
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Fish and Wildlife Service
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50 Part 17
Department of Commerce
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National Oceanic and Atmospheric Administration
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50 CFR Parts 223 and 224
Endangered and Threatened Wildlife; 12-Month Finding on a Petition To
Identify the Northwest Atlantic Leatherback Turtle as a Distinct
Population Segment and List It as Threatened Under the Endangered
Species Act; Final Rule
Federal Register / Vol. 85, No. 154 / Monday, August 10, 2020 / Rules
and Regulations
[[Page 48332]]
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DEPARTMENT OF INTERIOR
Fish and Wildlife Service
50 Part 17
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DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
50 CFR Parts 223 and 224
[Docket No. 200717-0190]
RIN 0648-XF748
Endangered and Threatened Wildlife; 12-Month Finding on a
Petition To Identify the Northwest Atlantic Leatherback Turtle as a
Distinct Population Segment and List It as Threatened Under the
Endangered Species Act
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce; U.S. Fish and Wildlife
Service (USFWS), Interior.
ACTION: Notification of 12-month petition finding.
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SUMMARY: We, NMFS and USFWS, announce a 12-month finding on a petition
to identify the Northwest Atlantic population of the leatherback turtle
(Dermochelys coriacea) as a distinct population segment (DPS) and list
it as threatened under the Endangered Species Act (ESA). In response to
the petition, we completed a comprehensive status review of the
species, which also constitutes the 5-year review of the species, to
determine potential DPSs following the Policy Regarding the Recognition
of Distinct Vertebrate Population Segments Under the ESA and to perform
extinction risk analyses. Based on the best scientific and commercial
data available, including the Status Review Report, and after taking
into account efforts made to protect the species, we conclude that
seven populations would meet the discreteness and significance criteria
for recognition as DPSs, including the Northwest Atlantic population.
However, even if we were to list them separately, all seven DPSs would
meet the definition for endangered species (i.e., they are in danger of
extinction throughout all or a significant portion of their range). The
species is already listed as endangered throughout its range. We have
determined that the listing of DPSs is not warranted, and therefore we
do not propose any changes to the existing global listing.
DATES: This finding was made on August 10, 2020.
ADDRESSES: The Status Review Report are available on NMFS' website at
https://www.fisheries.noaa.gov/species/leatherback-turtle.
FOR FURTHER INFORMATION CONTACT: Jennifer Schultz, NMFS Office of
Protected Resources, (301) 427-8443, [email protected]. Persons
who use a Telecommunications Device for the Deaf (TDD) may call the
Federal Information Relay Service (FIRS) at 1-800-877-8339, 24 hours a
day and 7 days a week.
SUPPLEMENTARY INFORMATION:
Background
The leatherback turtle species as a whole was listed as an
endangered species (one determined to be threatened with worldwide
extinction) (35 FR 8491; June 2, 1970), under the Endangered Species
Conservation Act of 1969, the precursor statute to the ESA (16 U.S.C.
1531 et seq.). When the ESA was enacted in 1973, it specifically
provided for continuity with the lists previously in effect under the
Endangered Species Conservation Act. Section 4(c)(3) of the ESA
directed that species on the lists of endangered foreign or native
wildlife at the time the ESA took effect would be deemed ``endangered
species'' under the ESA without interruption. See 39 FR 1444 (January
9, 1974) (explaining transition provisions); 39 FR 1158, 1172 (January
4, 1974) (setting out the final list of ``endangered foreign
wildlife,'' including ``Turtle, Leatherback'' at 50 CFR 17.11).
On September 20, 2017, the Blue Water Fishermen's Association
petitioned NMFS and USFWS (together, the Services) to identify the
Northwest (NW) Atlantic leatherback turtle population as a DPS and to
list it as threatened under the ESA. On December 6, 2017, NMFS
published a ``positive'' 90-day finding in the Federal Register (82 FR
57565) announcing the determination that the petition presented
substantial information indicating that the petitioned action may be
warranted. At that time, NMFS also solicited information on leatherback
turtles and announced that it would commence, jointly with USFWS, a
status review of the entire listed species, pursuant to ESA section
4(b)(3)(A) and 50 CFR 424.14. The resulting Status Review Report
includes all information used to evaluate the petitioned actions and
explains the process followed by the Status Review Team (i.e., the
Team). The following summarizes that information; for additional
details, please see the Status Review Report (see ADDRESSES).
ESA Statutory, Regulatory, and Policy Provisions and Evaluation
Framework
Under the ESA, the term ``species'' includes any subspecies of fish
or wildlife or plants, and any DPS of any vertebrate fish or wildlife
which interbreeds when mature (16 U.S.C. 1532(16)). The Services
adopted a joint policy clarifying their interpretation of the phrase
``distinct population segment'' for the purposes of listing, delisting,
and reclassifying a species under the ESA (``Policy Regarding the
Recognition of Distinct Vertebrate Population Segments Under the
Endangered Species Act,'' 61 FR 4722 (Feb. 7, 1996; ``DPS Policy'').
The DPS Policy stipulates two elements that must be considered: (1)
Discreteness of the population segment in relation to the remainder of
the species to which it belongs; and (2) the significance of the
population segment to the species to which it belongs.
Section 3 of the ESA defines an endangered species as any species
which is in danger of extinction throughout all or a significant
portion of its range and a threatened species as one which is likely to
become an endangered species within the foreseeable future throughout
all or a significant portion of its range (16 U.S.C. 1532(6) and (20)).
Thus, we interpret an ``endangered species'' to be one that is
presently in danger of extinction. A ``threatened species,'' on the
other hand, is not presently in danger of extinction, but is likely to
become so within the foreseeable future (that is, within a specified
later time). In other words, the primary statutory difference between a
threatened and endangered species is the timing of when a species may
be in danger of extinction, either presently (endangered) or within the
foreseeable future (threatened). The ESA uses the term ``foreseeable
future'' to refer to the time over which identified threats are likely
to impact the biological status of the species. The duration of the
``foreseeable future'' in any circumstance is inherently fact-specific
and depends on the particular kinds of threats, the life-history
characteristics, and the specific habitat requirements for the species
under consideration. The existence of threats to a species and the
species' response to such threats are not, in general, equally
predictable or foreseeable. Hence, in some cases, the ability to
foresee a threat to a species is greater than the ability to foresee
the species' exact response, or the timeframe of such a response, to
that
[[Page 48333]]
threat. For purposes of making this 12-month finding, the relevant
consideration is whether the species' population response (i.e.,
abundance, productivity, spatial distribution, diversity) is
foreseeable, not merely whether the emergence of a threat is
foreseeable. The foreseeable future extends only as far as we are able
to reliably predict the species' population response to threats.
Pursuant to the ESA and our implementing regulations, we determine
whether a species is threatened or endangered based on any one or a
combination of the following ESA section 4(a)(1) factors or threats (16
U.S.C. 1533(a)(1), 50 CFR 424.11(c)):
1. The present or threatened destruction, modification, or
curtailment of its habitat or range;
2. Overutilization for commercial, recreational, scientific, or
educational purposes;
3. Disease or predation;
4. Inadequacy of existing regulatory mechanisms; or
5. Other natural or manmade factors affecting its continued
existence, which could include but are not limited to: Fisheries
bycatch; vessel strikes; pollution (including marine debris and
plastics, contaminants, oil and gas activities, and derelict fishing
gear); natural disasters; climate change; and oceanographic regime
shifts.
Section 4(b)(1)(A) of the ESA requires us to make listing
determinations based solely on the best scientific and commercial data
available after conducting a review of the status of the species and
after taking into account efforts being made by any State or foreign
nation or political subdivision thereof to protect the species'
existence (16 U.S.C. 1533(b)(1)(A)).
Approach to the Status Review
The Services convened a team of NMFS and USFWS biologists (i.e.,
the Team) to gather and review the best available scientific and
commercial data on the leatherback turtle, assess the discreteness and
significance of populations by applying the DPS Policy, evaluate the
extinction risk of any population segments that meet the DPS criteria,
and document all findings in a report (i.e., the Status Review Report).
Although the petitioner requested evaluation only of the NW Atlantic
leatherback population, we instructed the Team to perform a
comprehensive status review to identify and evaluate the status of all
potential DPSs.
The Team compiled information on leatherback turtle life history,
biology, ecology, demographic factors, and threats. This included the
information received in the petition and in response to the Federal
Register request associated with the 90-day finding (82 FR 57565;
December 6, 2017). The Team also requested leatherback nesting data
from beach monitoring programs. To evaluate recent abundance and
trends, unpublished nesting beach monitoring datasets were often the
best available data (i.e., most recent and relevant). The Team assessed
these data in terms of standardization (i.e., the use of standardized
methodology), consistency (i.e., consecutive seasonal data collection),
and duration of data collection (i.e., the number of years that data
were collected). When evaluating threats, peer-reviewed information,
specifically primary research with large sample sizes and long-term
sampling duration, was often the best available data. In some
locations, reports from governments or non-governmental organizations
and expert opinion constituted the best available information. The Team
also addressed the source and magnitude of any uncertainty and the
impact on its conclusions.
The Team evaluated the discreteness and significance of each
population and provided their evaluation of whether each population
would meet the criteria of the DPS Policy. The DPS Policy states that a
population of a vertebrate species may be considered discrete if it
satisfies one of the following conditions: (1) It is markedly separated
from other populations of the same taxon as a consequence of physical,
physiological, ecological, or behavioral factors (quantitative measures
of genetic or morphological discontinuity may provide evidence of this
separation); or (2) it is delimited by international governmental
boundaries within which differences in control of exploitation,
management of habitat, conservation status, or regulatory mechanisms
exist that are significant in light of section 4(a)(1)(D) of the ESA
(61 FR 4722, February 7, 1996). While the Team used the term ``DPS'' in
describing and discussing populations that they concluded meet the
requirements of discreteness and significance, it is important to note
that the DPS term is used throughout the Status Review Report for ease
of reference only. A DPS is formally recognized under the ESA only upon
a listing action by the Services, and the Services cannot delegate
authority to take formal listing actions to status review teams. The
information compiled by the Team must be reviewed by the Services,
which retain responsibility for making the listing determination after
complying with all the requirements of Section 4 of the ESA and
considering agency policies. Because we ultimately conclude for the
reasons discussed in this finding that it would not be appropriate to
disaggregate the existing global listing into DPSs, references in the
Status Review Report (and in this finding when we are reviewing the
information presented by the Team) must be understood as references to
potential or hypothetical DPSs only.
The Team evaluated significance in terms of the importance of the
population segment to the overall welfare of the species, such as: (1)
Persistence of the population segment in an unusual or unique
ecological setting; (2) evidence that loss of the population segment
would result in a significant gap in the range of the taxon; (3)
evidence that the DPS represents the only surviving natural occurrence
of a taxon that may be more abundant elsewhere as an introduced
population outside its historic range; or (4) evidence that the
population segment differs markedly from other populations of the
species in its genetic characteristics.
For each population segment that the Team determined would meet the
criteria of the DPS Policy (which the Team and we refer to as a ``DPS''
for ease of reference), the Team performed an extinction risk analysis,
which involved the evaluation of demographic factors and threats.
Demographic factors reflect the impact that operative threats have had
on the species. In some cases those threats or the impacts from the
threats are continuing in nature. The demographic factors included
abundance, productivity, spatial distribution, and diversity. Because
sea turtles spend the majority of their lives at sea, where they are
spread across vast distances, it is difficult to estimate total
abundance. However, the number of nesting females can be counted
directly, or estimated indirectly by counting the number of nests on
beaches, during a nesting season. Females nest more than once in a
season (i.e., clutch frequency, which is the average number of nests
per season) and do not nest every season (i.e., remigration interval,
which is the average number of years between successive nesting
seasons). To calculate the index of nesting female abundance at a
nesting beach, the Team summed the total number of nests over the most
recent remigration interval (i.e., a run-sum) and divided this number
by the clutch frequency. The Team performed these calculations only if
available data were recent (i.e., last year of the remigration interval
occurred in 2014 or more recently), consistent
[[Page 48334]]
(i.e., seasonal data collected for each year of the remigration
interval), and collected in a standardized manner (i.e., data
collection methods remained the same over the remigration interval), as
further detailed in the Status Review Report. To provide a total index
of nesting female abundance for each DPS, we summed the indices of
nesting female abundance for all monitored beaches used by that DPS.
The total index of nesting female abundance for each DPS is an index
(rather than a census) because not all nesting beaches met these
criteria. However, the nesting beaches that were not included were
generally unmonitored or not recently monitored because they host few
nesting females. Even where data were not sufficient to allow for a
calculation of the index of nesting female abundance, the Team provided
all available data to ensure the analysis would be as robust as
possible.
The Team evaluated the productivity for each DPS by evaluating
nesting trends (through trend analyses or bar graphs) and productivity
metrics. Where available data allowed it, they estimated the long-term
trend for individual beaches using a Bayesian state-space model of
stochastic exponential population growth (Boyd et al. 2017), where the
rate parameter describes the annual percent change in observed nest
counts (or female counts where applicable) over the period of data
collection. This is further explained in the Status Review Report. To
reflect current trends over approximately three remigration intervals,
the criteria for trend analyses were as follows: Nesting data (i.e.,
nest or nesting female counts) consistently collected over nine or more
years in a standardized manner (for that site), with the most recent
data collection in 2014 or later and with a minimum average number of
nests of 50 annually. The Team reported the median trend, along with
the standard deviation (sd), 95 percent credible interval (CI), and an
``f statistic'' which is the proportion of the posterior distribution
with the same sign as the median (i.e., the confidence that the trend
is positive or negative). When the data did not meet the criteria for
performing trend analyses, the Team provided bar graphs and/or
historical data in the Status Review Report. Based on the trend
analysis (where possible) and the best available historical data, the
Team characterized the nesting trend for each DPS as decreasing,
stable, or increasing. The Team also evaluated the following
productivity metrics (if available): Average size of nesting female;
nesting female survivorship; remigration interval; clutch size; clutch
frequency; internesting interval; incubation period; hatching success
(the proportion of eggs in a nest that produce live hatchlings); and
sex ratio. Each of these metrics contributes to the growth rate, or
reproductive potential, of the population.
For each DPS, the Team evaluated spatial distribution, which
included the number and location of nesting beaches and foraging areas,
as well as spatial structure (i.e., whether the DPS exists as a single
population or several subpopulations connected by metapopulation
dynamics). The Team also evaluated diversity, which like spatial
distribution, is a measure of resilience. In general, diverse
populations with broad spatial distributions and metapopulation
dynamics are more resilient to threats and environmental changes than
less diverse populations with narrow distributions.
For each DPS, the Team next evaluated each of the ESA Section
4(a)(1) factors (or ``threats'') as listed above (16 U.S.C. 1533(a)(1),
50 CFR 424.11(c)). For each threat, the Team used the best available
information to describe the threat, identify which life stages are
affected, and describe the impact to the DPS with as much specificity
as the best available information allowed to link the threat to the
demographic factor it affected. The best available data often allow
only for qualitative assessment. For each DPS, the Team identified the
primary threat(s) to its continued existence, as well as other threats.
The Team considered the impact of each threat individually, with the
primary threat(s) given the greatest weight, and all threats
cumulatively, to determine the extinction risk. To assess confidence in
the extinction risk determination, the Team identified any sources of
uncertainty and the impact of uncertainty on the conclusions. They
analyzed all threats assuming the DPS had lost ESA protections going
forward because a DPS would not receive such protections if it was not
listed under the ESA. For example, a DPS would not have benefits of
section 9 take prohibitions or section 7 consultations on actions that
may affect the DPS.
The Team performed an extinction risk assessment for each of the
seven DPSs by evaluating the demographic factors and threats, as
described above. Then, the Team voted, based on the best available
data, on whether the extinction risk of each DPS was high, moderate, or
low, following the definitions included in NMFS' internal guidance
document, ``Guidance on Responding to Petitions and Conducting Status
Reviews under the Endangered Species Act, Section II'' (i.e., NMFS'
Guidance; November 9, 2017) and in the Status Review Report.
After the Team completed its draft Status Review Report, the
Services met to review and discuss that document and conservation
efforts. The Services based our status determinations of the DPSs on
the best scientific and commercial data available (as compiled and
reflected in the Status Review Report) and after taking into account
efforts by States and foreign nation, or any political subdivision
thereof, to protect the species as mandated by the statute.
DPS Analysis
The following is a summary of the DPS analysis conducted by the
Team. For a detailed description of the Team's analyses of discreteness
and significance, please see the Status Review Report. As a starting
point, the Team considered seven leatherback populations that were
previously identified as regional management units (RMUs) by Wallace et
al. (2010) and recognized as subpopulations under the International
Union for Conservation of Nature (IUCN) Red List (https://www.iucnredlist.org/species/6494/43526147). The Team found that seven
leatherback populations met the discreteness and significance criteria
per the DPS Policy and identified the following potential DPSs:
Northwest (NW) Atlantic; Southwest (SW) Atlantic; Southeast (SE)
Atlantic; SW Indian; Northeast (NE) Indian; West Pacific; and East
Pacific.
Discreteness
The Team evaluated all populations for discreteness and determined
that each showed marked separation from the others as a consequence of
behavioral and physical factors. Behavioral factors, especially
returning to waters off a turtle's natal beach to breed, have prevented
interbreeding, resulting in reproductive isolation, as indicated by
genetic discontinuity.
Although some populations use the same foraging areas, tagging and
telemetry studies also demonstrate the discreteness of the populations
at nesting beaches. Physical factors, such as land masses, ocean
currents, and other oceanographic features, have established and
reinforced barriers to gene flow among the seven populations.
Genetic data provide the most compelling evidence for discreteness
among the seven populations. The most recent and comprehensive global
analysis of published and unpublished
[[Page 48335]]
mitochondrial deoxynucleic acid (mtDNA) sequence data (i.e., 28
haplotypes, which are unique sequences of mtDNA) evaluated samples
collected from 21 nesting sites representing key regions from all ocean
basins (Dutton et al. 2007; Dutton et al. 2013; Shanker et al. 2011;
Dutton and Shanker 2015); analyzing the evolutionary relationship of
these data revealed three distinct haplogroups (i.e., similar
haplotypes that cluster together, relative to other haplotypes) that
are geographically segregated across the Atlantic, Indian, and Pacific
Oceans (Dutton, unpublished data; NMFS and USFWS 2020). Early mtDNA
analyses indicated strong genetic discontinuity, globally
(FST = 0.415, P <0.001) and within ocean basins
(FST = 0.203 to 0.253, P <0.001; Dutton et al. 1999).
Wallace et al. (2010) combined these and other genetic data with
nesting, flipper tagging, and satellite telemetry data to identify
seven leatherback RMUs, which provided the starting point for our
identification of discrete populations.
From this starting point, the Team then evaluated more recent
genetic data. Subsequent genetic analyses confirmed genetic
discontinuity among the NW, SW, and SE Atlantic populations (Wallace et
al. 2010; Dutton et al. 2013; Carreras et al. 2013; Molfetti et al.
2013; Vargas et al. 2017). Elevated genetic differentiation at nuclear
DNA (FST = 0.211-0.86) indicates that males, like females,
likely return to the waters off their natal beaches to mate and that
male-mediated gene flow may not be as pronounced as previously thought
(Dutton et al. 2013; see Jensen et al. 2013). Nuclear (FST
>0.126, P <0.001; Dutton et al. 2013) and mtDNA (FST >0.061,
P = 0.05-0.001; Dutton et al. 2013; FST >0.061, P <0.01;
Vargas et al. 2017) analyses indicate genetic discontinuity between the
Atlantic populations and the SW Indian population. Preliminary mtDNA
results for leatherback turtles nesting at Little Andaman Island, India
(Shanker et al. 2011; Dutton and Shanker 2015), indicate that this
population is closely related to the extinct Malaysian population, with
which it shares common haplotypes. It is markedly different from the
South African nesting population, as well as those in the West Pacific
population (Dutton et al. 2007, 2013 and unpublished). Samples from
extant and extirpated nesting aggregations of the NE Indian population
(Shanker et al. 2011; Dutton and Shanker 2015; Dutton et al.
unpublished data) are genetically differentiated from the SW Indian
population (FST = 0.415, P <0.003; Dutton et al. 1999) and
the West Pacific population (X2 = 49.346, P = 0.002; Dutton
et al. 2007). There is genetic discontinuity between the West and East
Pacific populations, as demonstrated by significant genetic
differentiation between the samples from Solomon Islands in the western
Pacific and Mexico or Costa Rica in the eastern Pacific (FST
= 0.270 and 0.331, P <0.001; Dutton et al. 1999). Genetic discontinuity
among all seven populations provides evidence for marked separation
from the others and thus discreteness of each population.
Tagging and telemetry studies confirm marked separation of the
seven populations because nesting sites remain distant and isolated.
Nesting females of one population have not been tracked to, or observed
on, beaches used by another population, even though telemetry data
indicate shared use of foraging areas by different populations.
Telemetry studies demonstrate that females nesting on NW Atlantic
beaches move throughout most of the North Atlantic from the Equator to
about 50[deg] N latitude (Ferraroli et al. 2004; Hays et al. 2004;
James et al. 2005a; James et al. 2005b; 2005c; Eckert 2006a; Eckert et
al. 2006b; Hays et al. 2006; Doyle et al. 2008; Evans 2008; Dodge et
al. 2014; Fossette et al. 2014; Aleksa 2017; Aleksa et al. 2018).
Turtles originating from beaches of the NW Atlantic appear to mix at
foraging areas throughout the North Atlantic Ocean (Fossette et al.
2014), but their movements rarely extend into waters south of the
Equator. Tagging studies further support the connectivity within and
among nesting beaches and foraging areas of the North Atlantic Ocean
(Tro[euml]ng et al. 2004; Br[auml]utigam and Eckert 2006;
Chac[oacute]n-Chaverri and Eckert 2007; Turtle Expert Working Group
(TEWG) 2007; S[ouml]nmez et al. 2008; Dutton et al. 2013b; Horrocks et
al. 2016), but turtles tagged in the North Atlantic Ocean have never
been found on nesting beaches in Brazil (SW Atlantic population) or
Africa (SE Atlantic population). In the South Atlantic Ocean, post-
nesting females tracked from nesting beaches in Gabon and Brazil use
the same foraging areas, including waters off SW Africa, in the south
equatorial Atlantic and off SE Brazil and Uruguay (Almeida et al. 2011;
Witt et al. 2011). Turtles incidentally captured in fisheries off South
America (Billes et al. 2006, L[oacute]pez-Mendilaharsu et al. 2009)
also demonstrate that turtles originating from the SW and SE Atlantic
Ocean beaches share foraging areas. Despite such mixing at foraging
areas, there is no evidence for the shared use of nesting beaches.
Genetic data indicate that turtles return to their natal beaches to
nest on opposite sides of the Atlantic Ocean (Dutton et al. 2013;
Vargas et al. 2017), and no tag recoveries contradict these data.
In the Indian Ocean, telemetry studies have been conducted at South
African nesting beaches in the SW Indian Ocean (Hughes et al. 1998;
Luschi et al. 2006; Robinson et al. 2016) and at Andaman Islands
nesting beaches in the NE Indian Ocean (Namboothri et al. 2012;
Swaminathan et al. 2019). South African nesting females showed diverse
movements that were highly influenced by complex oceanographic currents
and features that lead them to foraging destinations in the South
Atlantic Ocean, SW Indian Ocean, and Mozambique Channel (Hughes et al.
1998, Luschi et al. 2006, Robinson et al. 2016). About half of the 10
post-nesting females tagged at the Andaman Islands moved westward: Two
individuals reached the Mozambique Channel; the other half moved
southeastward, past the Indonesian islands of Sumatra and Java, with
one leatherback reaching an apparent foraging ground off NW Australia
before transmissions stopped (Namboothri et al. 2012; Swaminathan et
al. 2019). Despite overlap in one foraging area (i.e., reaching the
Mozambique Channel), tagging data do not indicate movement between the
distant nesting beaches.
Within the Pacific Ocean, nearly all turtles tracked from East
Pacific nesting beaches moved southward across the Equator to forage in
open-ocean waters of the SE Pacific Ocean or in the coastal waters of
Central America, Peru, and Chile. The movements of post-nesting females
from the West Pacific Ocean are dependent on the season in which they
nest, with winter-nesting females predominantly tracked into the
Southern Hemisphere and summer-nesting females foraging in diverse
coastal and oceanic ecosystems throughout the northern Indo-Pacific
region (Benson et al. 2011). Telemetry data indicate little or no
overlap with foraging destinations utilized by nesting females of the
East and West Pacific populations (Bailey et al. 2012; Benson et al.
2011). However, a genetic study of bycaught turtles off the coast of
Chile and Peru indicated that 15 percent of leatherback turtles
originated from West Pacific nesting beaches (Donoso and Dutton 2010),
suggesting that foraging overlap may be more prevalent than estimated
by telemetry data. Still, there is no genetic evidence for contemporary
interbreeding between the two populations (Dutton et al. 2007), and
telemetry and tagging data do not indicate movement between the distant
[[Page 48336]]
nesting beaches. Thus, flipper tagging and satellite telemetry data
support the marked separation, and thus discreteness, of the seven
populations at their nesting beaches.
Physical factors likely shape and reinforce the behavior patterns
that result in reproductive isolation. Though the species has a global
range, with foraging areas extending into high latitudes, nesting
mainly occurs on tropical or subtropical beaches. Post-hatchling
dispersal is determined by the ocean currents they encounter off
nesting beaches. While adults move throughout tropical and temperate
waters irrespective of ocean currents, both males and females return to
the waters off their natal nesting beach to mate. This natal homing is
somewhat flexible, (Dutton et al. 2013; Jensen et al. 2013), creating
reproductive isolation only among distant nesting sites, which may also
be physically separated from one another by land masses and
oceanographic barriers to gene flow. For example, leatherback turtles
in the Atlantic Ocean are physically separated from those in the
Pacific Ocean by the Americas. Though leatherback turtles have greater
cold tolerance than other sea turtles, they do not appear to venture
into latitudes greater than 47[deg] S or 71[deg] N (Eggleston 1971;
Eckert et al. 2012). Therefore, the low latitude and cold waters of the
Cape Horn Current likely prevent movement between the Atlantic and
Pacific Oceans. Within ocean basins, nesting beaches of the discrete
populations are separated by long distances of uninterrupted deep water
(e.g., the East Pacific Barrier and the mid-Atlantic Barrier). While
leatherback turtles clearly cross these open-ocean barriers to reach
distant foraging areas, they do not appear to do so for nesting and
breeding, but rather return to their natal region to breed and nest
(Barragan et al. 1998; Dutton et al. 1999; Barragan and Dutton 2000;
Dutton et al. 2013). Within ocean basins, currents shape post-
hatchlings' movement patterns, which they may retain as adults (e.g.,
Fossette et al. 2010; Benson et al. 2011). The NW Atlantic leatherback
population appears to be physically separated from the SE and SW
Atlantic populations by the current systems of the South and North
Atlantic Gyres, respectively. NW Atlantic leatherback nesting beaches
are adjacent to northward moving currents (e.g., Gulf Stream).
Leatherback hatchlings from these nesting beaches, therefore, are
transported northward, remaining in the North Atlantic Ocean. Those
that survive return to their nesting beaches as adults, completing
their life stages within the North Atlantic (Fossette et al. 2010;
Chambault et al. 2017). The SE and SW Atlantic populations are
similarly retained in the South Atlantic Ocean by the South Atlantic
Gyre and the Benguela Current, which flows northward along the SE coast
of Africa, restricting movement into the Indian Ocean. Within the
Indian Ocean, the Somali Current runs between the nesting beaches of
the SW and NE Indian populations. The NE Indian and West Pacific
populations likely became isolated as a result of exposed land barriers
between Indonesia, New Guinea, and the Philippines as a result of low
sea levels within the past 6,000 years (Barber et al. 2000). Seasonal
monsoons may also play a contemporary role by altering current
directions and hatchling dispersal patterns (Benson et al. 2011; Gaspar
et al. 2012). Thus, physical factors have likely helped to shape, or at
least reinforce, the reproductive isolation among distant nesting
beaches.
Based on these data, the Team concluded that the seven populations
demonstrate discreteness, or marked separation from each other, due to
behavioral and physical factors. These are the NW Atlantic, SW
Atlantic, SE Atlantic, SW Indian, NE Indian, West Pacific, and East
Pacific populations.
Significance
Each of the discrete populations is significant to the species
because the loss of any one would result in a significant gap (i.e., a
half or quarter of an ocean basin) in the range of the species. Several
populations also persist in unique ecological settings. Each population
likely possesses unique genetic characteristics and local adaptations
as a result of thousands of years of reproductive isolation, but none
have yet been identified because all genetic studies have involved
neutral markers. Therefore, the Team did not rely on evidence of unique
genetic characteristics and local adaptations for its significance
finding.
A loss of the NW Atlantic population would result in a gap (i.e.,
the entire North Atlantic Ocean) of the nesting and foraging range of
the species. If the NW Atlantic population were extirpated, it is
unlikely that leatherback turtles from other populations would
recolonize the North Atlantic Ocean in an ecological time frame (i.e.,
tens to hundreds of years), leaving a significant gap in the range of
the species. Extirpation of this population would also significantly
reduce the genetic diversity of the species, as reflected by the
possession of several unique haplotypes. Leatherback turtles of the NW
Atlantic Ocean also occur in a unique ecological setting; this is the
only DPS that regularly forages at high latitudes. Sightings have been
documented as far north as Norway and Iceland (Brongersma 1972; Goff
and Lien 1988; Carriol and Vader 2002; McMahon and Hayes 2006; Eckert
et al. 2012). Such high latitude foraging is likely facilitated by the
warm Gulf Stream, which meets cold water currents to create highly
productive foraging areas. The Team concluded that the NW Atlantic
population is biologically significant to the species.
In the SW Atlantic Ocean, leatherback turtles only nest in a small
area of the coastline of Brazil. All other nesting in South America
occurs above the Equator or on the Pacific Coast. Therefore, the loss
of this population would result in a gap of the nesting range of the
species (i.e., the SW Atlantic coast). Although SE Atlantic leatherback
turtles forage off the coasts of Brazil, Argentina, and Uruguay, they
do not breed there. Rather, they return to the waters off western
Africa to mate (Vargas et al. 2017). Therefore, if the SW Atlantic
population were extirpated, it is unlikely that leatherback turtles
from other populations would recolonize this region, leaving a
significant gap in the nesting range of the species. The extirpation of
this population would also significantly reduce the genetic diversity
of the species, as reflected by the possession of unique haplotypes and
high genetic diversity, despite the small population size (Vargas et
al. 2017). The SW Atlantic population is biologically significant to
the species.
Leatherback turtles of the SE Atlantic population nest in West
Africa and forage in the South Atlantic Ocean. This population is much
more abundant than the SW Atlantic population, which also forages in
the South Atlantic Ocean. Therefore, the loss of this population would
result in a gap of the nesting range of the species (i.e., western
Africa) and a significant reduction in the abundance of leatherback
turtles foraging throughout the South Atlantic Ocean. The extirpation
of this population would also significantly reduce the genetic
diversity of the species, as reflected by the possession of unique
haplotypes. The Team concluded that the SE Atlantic population is
biologically significant to the species.
In the SW Indian Ocean, leatherback turtles only nest in a small
area along the South African and Mozambican coastlines. No other
leatherback turtles nest in eastern Africa or in other areas throughout
the entire western Indian Ocean. Therefore, the loss of this population
would result in a gap of the
[[Page 48337]]
nesting range of the species (i.e., the SW Indian Ocean). The SW Indian
population also occurs in a unique ecological setting: It is the only
population to nest on temperate beaches. The warm Agulhas Current,
adjacent to the nesting beaches, likely facilitates their high-latitude
nesting. The Team concluded that the SW Indian population is
biologically significant to the species.
Leatherback turtles nest in small numbers in the NE Indian Ocean.
These nesting sites are separated from other Indian Ocean nesting sites
by at least 5,000 km. Although western Pacific nesting sites are
closer, males and females return to the waters off their natal beaches
to breed, preventing interbreeding among NE Indian and West Pacific
populations. Therefore, the loss of this population would result in a
gap of the nesting range of the species (i.e., the NE Indian Ocean).
The extirpation of this population would also significantly reduce the
genetic diversity of the species, as reflected by the possession of
unique haplotypes. The Team concluded that the NE Indian population is
biologically significant to the species.
West Pacific leatherback turtles nest in small numbers primarily in
four nations of the West Pacific Ocean. These nesting sites are
separated from East Pacific nesting sites by over 10,000 km. Though NE
Indian nesting sites are closer in distance, male and female philopatry
prevents interbreeding. Therefore, the loss of this population would
result in a gap of the nesting range of the species (i.e., the West
Pacific Ocean). The loss of this population would also result in a gap
of the foraging range of the species (i.e., the North Pacific Ocean).
The extirpation of this population would also significantly reduce the
genetic diversity of the species, as reflected by the possession of
unique haplotypes. The West Pacific population is ecologically unique
in two ways: It is the only population to forage in both hemispheres;
and it nests year-round, with nesting peaks in the summer and winter.
The Team concluded that the West Pacific population is biologically
significant to the species.
Leatherback turtles nesting on eastern Pacific coastlines also
forage in the East Pacific Ocean. A loss of this population would
result in a gap of the nesting range of the species (i.e., the East
Pacific Ocean). Though West Pacific leatherback turtles may forage off
the coasts of Peru and Chile, they do not breed there (Donoso and
Dutton 2010). Therefore, if the East Pacific population were
extirpated, it is unlikely that leatherback turtles from other
populations would recolonize this region, leaving a significant gap in
the nesting range of the species. The extirpation of this population
would also significantly reduce the genetic diversity of the species,
as the population possess several unique haplotypes. The East Pacific
population is unique in having the smallest nesting female size, clutch
size, and egg size of all populations, possibly reflecting unique
foraging conditions that are subject to oceanographic regime shifts
(e.g., the El Ni[ntilde]o Southern Oscillation, or ENSO). The Team
concluded that the East Pacific population is biologically significant
to the species.
DPS Summary
The Team found that seven populations met the definition for
discreteness. These populations are markedly separated as a result of
the behavioral factors of movement (as demonstrated by satellite
telemetry and flipper tagging studies) and philopatry, which has led to
reproductive isolation (as demonstrated by genetic discontinuity). They
are also physically separated by land masses, oceanographic features,
and currents. The Team found these seven populations to be significant
to the species because the loss of any one of them would result in a
significant gap in the range of the species as well as a significant
loss of genetic diversity, reducing the evolutionary potential of the
species. Some populations also occur in a unique ecological setting.
Thus, after reviewing the best available information, the Team
identified the following populations as potential DPSs: NW Atlantic, SW
Atlantic, SE Atlantic, SW Indian, NE Indian, West Pacific, and East
Pacific. The Team defined the potential DPSs as leatherback turtles
originating from nesting beaches within the boundaries for each DPS.
The range of each DPS, which also includes foraging areas, thus extends
beyond the nesting boundaries for most DPSs, and may overlap
extensively with the range of another DPS. The boundaries are based on
the best available genetic, telemetry, and observational data. When
such data were not available, the Team used information on possible
barriers to gene flow, such as oceanographic features. For ease of use,
the Team applied political boundaries when this did not conflict with
biological or oceanographic data. Additional information on the
boundaries is available in the following sections, which summarize the
extinction risk analysis for each DPS, and in the Status Review Report.
NW Atlantic DPS
The Team defined the NW Atlantic DPS as leatherback turtles
originating from the NW Atlantic Ocean, south of 71[deg] N, east of the
Americas, and west of Europe and northern Africa; the southern boundary
is a diagonal line between 5.377[deg] S, 35.321[deg] W and 16.063[deg]
N, 16.51[deg] W. The northern boundary reflects a straight latitudinal
line based on the northernmost documented occurrence of leatherback
turtles (Brongersma 1972; Goff and Lien 1988; Carriol and Vader 2002;
McMahon and Hayes 2006; Eckert et al. 2012). The southern boundary is a
diagonal line between the elbow of Brazil, where the Brazilian current
begins and likely restricts the nesting range of this DPS, and the
northern boundary of Senegal. The boundary between Senegal and
Mauritania was chosen because the SE Atlantic DPS does not appear to
nest above this boundary (Fretey et al. 2007).
The range of this DPS (i.e., all areas of occurrence) extends
throughout the North Atlantic Ocean, including the Caribbean Sea, Gulf
of Mexico (GOM), and Mediterranean Sea. Available data indicate that
the NW Atlantic DPS occurs (at varying levels of frequency) in the
waters of the following nations or territories: Albania, Algeria,
Anguilla, Antigua and Barbuda, Aruba, Azores, Bahamas, Barbados,
Belize, Bermuda, Bonaire, Bosnia and Herzegovina, Brazil, British
Virgin Islands, Canada, Cape Verde, Cayman Islands, Colombia, Costa
Rica, Croatia, Cuba, Cura[ccedil]ao, Cyprus, Denmark, Dominica,
Dominican Republic, Egypt, France, French Guiana, Greece, Greenland,
Grenada, Guadeloupe, Guatemala, Guyana, Haiti, Honduras, Iceland,
Ireland, Israel, Italy, Jamaica, Lebanon, Libya, Madeira, Malta,
Martinique, Mauritania, Mexico, Montenegro, Montserrat, Morocco,
Netherlands Antilles, Nicaragua, Norway, Panama, Portugal, Slovenia,
Spain, St. Barthelemy, St. Eustatius, St. Kitts and Nevis, St. Lucia,
St. Maarten, St. Pierre and Miquelon, St. Martin, St. Vincent and the
Grenadines, Suriname, Sweden, Syria, Trinidad and Tobago, Tunisia,
Turkey, Turks and Caicos Islands, United Kingdom, United States
(including Puerto Rico and the U.S. Virgin Islands (USVI), Venezuela,
and Western Sahara.
All nesting in this DPS occurs in the NW Atlantic Ocean,
concentrated from the southeast United States throughout the Wider
Caribbean Region (Dow et al. 2007). Leatherback nesting in the NW
Atlantic can be grouped into several broad geographical areas,
including the
[[Page 48338]]
U.S. mainland (primarily Florida), North Caribbean (including USVI and
Puerto Rico), West Caribbean (Honduras to Colombia), and Southern
Caribbean/Guianas (Venezuela to French Guiana; TEWG 2007). The largest
nesting aggregations occur in Trinidad, French Guiana, and Panama. The
northern-most confirmed nesting occurs in North Carolina, but there has
been a crawl recorded as far north as Assateague Island National
Seashore, Maryland (Rabon et al. 2003). No nesting occurs in the
Mediterranean Sea (Casale and Margaritoulis 2010).
Nesting occurs on unobstructed, high-energy beaches with either a
deep water oceanic approach or a shallow water approach with mud banks,
but without coral or rock formations (TEWG 2007). The main
characteristics of leatherback nesting beaches include coarse-grained
sand; steep, sloping littoral zone; obstacle-free approach; proximity
to deep water; and oceanic currents along the coast (Hendrickson and
Balasingam 1966 in Eckert et al. 2015). During the nesting season,
adult females and males inhabit the waters off nesting beaches. During
a nesting season, females generally stay within about 100 km of their
nesting beaches, remaining close to the coast on the continental shelf,
and engaging in shallow dives (Eckert et al. 2012). Intra-seasonal
movement of greater than 100 km also occurs, especially between French
Guiana and Suriname (Fossette et al. 2007; Georges et al. 2007), Panama
and Costa Rica (Chac[oacute]n-Chaverri and Eckert 2007), and among
Caribbean nesting beaches, including those on Trinidad (Brautigam and
Eckert 2006; Georges et al. 2007; Horrocks et al. 2016). Adult males
migrate from temperate foraging areas in the North Atlantic Ocean to
waters off nesting beaches, typically arriving before the nesting
season and remaining for the majority of the season (James et al.
2005b; Doyle et al. 2008; Dodge et al. 2014).
Foraging areas of the NW Atlantic DPS include coastal and pelagic
waters of the North Atlantic Ocean (Eckert et al. 2012; Saba 2013;
Shillinger and Bailey 2015). These waters include the GOM, North
Central Atlantic Ocean, northwestern Atlantic shelf waters of the
United States and Canada, waters along the southeastern U.S. coast, the
Mediterranean Sea, and the northeastern Atlantic shelf waters of Europe
and northwestern Africa (TEWG 2007). Some post-nesting females also
remain in tropical waters to forage (Fossette et al. 2010). This DPS is
mostly commonly associated with open-ocean and coastal shelf foraging
areas off Nova Scotia (Canada), northeastern United States, GOM,
northwestern Europe, and northwestern Africa (James et al. 2005a,
2006b, 2007; Eckert 2006; Eckert et al. 2006; Fossette et al. 2010a;
Fossette et al. 2010b; Dodge et al. 2014; Stewart et al. 2016; Aleksa
et al. 2018). Fossette et al. (2014) analyzed available satellite
telemetry data from 1995 to 2010 on post-nesting females (n = 93) as
well as males (n = 4), females (n = 8), and a juvenile (n = 1) from
foraging grounds throughout the Atlantic Ocean. They found widespread
use of the North Atlantic Ocean (Fossette et al. 2014). High-use areas
mainly occurred in the central (25 to 50[deg] N, 50 to 30[deg] W) and
eastern Atlantic Ocean, in particular in the waters offshore Western
Europe, around Cape Verde (year-round) and the Azores (October to
March; Fossette et al. 2014). Fossette et al. (2014) found that
seasonal high-use areas also occurred along the eastern U.S. coast
(April to June and October to December) and off Canada (July to
December). The GOM is also a high-use foraging area, with a peak in the
northeast GOM during August and September (Aleksa et al. 2018).
Overall, leatherback turtles of the North Atlantic population appear to
have a diverse array of foraging habitat available.
Abundance
The total index of nesting female abundance for the NW Atlantic DPS
is 20,659 females. The nesting beaches with the greatest abundance have
been included in this index, and most beaches with an unquantified
number of nests likely host few nesting females. We based this index on
24 nesting aggregations in 10 nations: Trinidad and Tobago (n =
11,324), French Guiana (n = 2,519), Panama (n = 2,251), United States
(n = 1,694), Costa Rica (n = 1,306), Suriname (n = 698), Grenada (n =
499), Venezuela (n = 215), Guyana (n = 76), and Nicaragua (n = 10).
With the possible exception of Colombia, our total index does not
include 31 unquantified but likely small nesting aggregations for which
data are not available. It also does not include outdated data
published by Dow et al. (2007), which includes binned crawls,
categorized as less than 25, 25 to 100, 100 to 500, 500 to 1000, or
unknown abundance. Crawls or emergences (measured as females or tracks
on beaches) include both successful egg-laying and unsuccessful
nesting, so the number of crawls represents approximately two to 10
times the number of nests (Dow et al. 2007). Because the Dow et al.
data, which are more than 10 years old and do not provide the number of
actual nests, may not be representative of recent nesting trends, we
did not include them in our total index. To calculate the indices of
nesting female abundance, we added the number of nests over the last 3
years (representing the most recent remigration interval; Eckert et al.
2012) and divided by the clutch frequency (site-specific values or,
when such values were not available, the average of the site-specific
values, i.e., 5.5 clutches per season).
Our total index of nesting female abundance is based on the best
available data for this DPS. It is the most robust estimate of nesting
females at this time because it only includes available nesting data
from recently and consistently monitored nesting beaches. Our total
index does not include data from beaches where we were unable to
quantify the number of nesting females, either due to the lack of
recent or available nesting data or because only crawl data were
reported (often on smaller nesting beaches). Scattered nesting may
occur on beaches throughout the region, but because these beaches are
not monitored, or have not been recently monitored, recent data are not
available.
Nesting in the NW Atlantic DPS is characterized by many small
nesting beaches. Large nesting aggregations are rare; only about 10
leatherback nesting beaches in the Wider Caribbean Region (about two
percent of the DPS's total nesting sites) host more than 1,000 crawls
annually (Dow Piniak and Eckert 2011). Only one site, Grande Riviere in
Trinidad, hosts more than 5,000 nesting females, representing 29
percent of the total index of nesting female abundance. Relatively
large nesting aggregations are also found in Matura (Trinidad),
Chiriqui Beach (Panama), and Cayenne and Remire Montjoly (French
Guiana). In contrast, most known nesting beaches support a small
nesting female abundance; 71 percent of the total nesting sites record
annual crawls of less than 100 (Dow Piniak and Eckert 2011). The number
of nesting females is unquantified at 31 beaches (i.e., the majority of
nesting sites for the DPS). However, for the reasons identified above,
most of those sites have small abundance levels as inferred from the
numbers of crawls estimated by Dow et al. (2007). Therefore, our total
index of nesting female abundance represents the most robust estimate
allowed by the best available data and includes the majority of nesting
females because the largest nesting aggregations were included. The
data regarding additional nesting aggregations are not sufficiently
recent, specific, or reliable for inclusion, and the contribution of
these nesting
[[Page 48339]]
aggregations to the total index is expected to be small.
Our total index of nesting female abundance is similar in
comparison to other published estimates. TEWG (2007) estimated the
abundance of NW Atlantic leatherback turtles using nesting data from
2004 and 2005. At that time, the number of adult females (equating to
total index of nesting female abundance in our analysis) was estimated
to be approximately 18,700 (range 10,000 to 31,000). While a wide range
was provided, the point estimate in TEWG (2007) is similar to, albeit
slightly lower than, our total index of 20,659 nesting females. The
most recent, published IUCN Red List assessment for the NW Atlantic
Ocean subpopulation estimated a total of 20,000 mature individuals (The
NW Atlantic Working Group 2019). Our total index, which only includes
nesting females, exceeds their estimate, likely due to our use of a 3-
year remigration interval, which has increased at some locations in
recent years (e.g., 4.5 years at St. Croix; K.R. Stewart, The Ocean
Foundation and C. Lombard, USFWS, pers. comm., 2019).
We conclude that the total index of nesting females for the NW
Atlantic DPS is 20,659 females. The nesting beaches with the greatest
abundance have been included in our total index, and most beaches with
an unquantified number of nests likely host few nesting females.
Current nesting female abundance is not at a level where stochastic or
environmental changes would have catastrophic impacts, but the
abundance at several nesting sites with previously high density has
declined drastically. However, as we discuss below, a declining nest
trend and several existing threats will likely continue to reduce this
abundance.
Productivity
The NW Atlantic DPS exhibits decreasing nest trends at nesting
aggregations with the greatest indices of nesting female abundance.
Though some nesting aggregations indicate increasing trends, most of
the largest ones demonstrate declining nest trends. We evaluated nest
trends by using nest count data consistently collected using a
standardized approach for at least 9 years, with the last year of data
in 2014 or more recently and with an average of more than 50 nests
annually. When data did not meet these criteria, we evaluated bar
graphs provided in the Status Review Report to consider all available
data. Thus, these data are representative of the DPS because they
include the largest nesting aggregations. With the possible exception
of Colombia, nesting aggregations for which data are not available are
likely small. Significant declines have been observed at nesting
beaches with the greatest historical or current nesting female
abundance, most notably in Trinidad and Tobago (Grande Riviere, Fishing
Pond, and Tobago), Suriname, French Guiana (Awala-Yalimapo), Florida,
and Costa Rica (Tortuguero). Therefore, these nest trends represent the
best available data for this DPS.
In Trinidad and Tobago, trends in annual nest counts were largely
negative between 2009 and 2017, the years for which data were
available. For Trinidad, we analyzed trends for three separately
monitored beaches, including Grande Riviere, Matura, and Fishing Pond.
The long-term trend was negative for Grande Riviere (median = -6.9
percent; sd = 17.4 percent; 95 percent CI = -43.8 to 26.9 percent; f =
0.682; mean annual nests = 13,272), positive for Matura (median = 1.8
percent; sd = 15.1 percent; 95 percent CI = -29.2 to 33.0 percent; f =
0.561; mean annual nests = 7,359), and negative for Fishing Pond
(median = -19.3 percent; sd = 15.1 percent; 95 percent CI = -49.8 to
12.0 percent; f = 0.916; mean annual nests = 3,892). For Tobago, the
median trend was -0.9 percent annually (sd = 11.3 percent; 95 percent
CI = -25.0 to 21.5 percent; f = 0.540; mean annual nests = 452).
For French Guiana, we analyzed nest count data from 2002 to 2017
for Awala-Yalimapo beach in the west and data from 1999 to 2017 for
Cayenne and Remire Montjoly beaches in the east. There was a steep
decline at Awala-Yalimapo, with a median trend of -19.4 percent
annually (sd = 12.2 percent; 95 percent CI = -43.2 to 6.0 percent; f =
0.942; mean annual nests = 3,200). In contrast to Awala-Yalimapo, nest
counts at Cayenne and Remire Montjoly increased by 2.8 percent annually
(sd = 12.9 percent; 95 percent CI = -24.9 to 27.9 percent; f = 0.596;
mean annual nests = 3,498). In addition, leatherback nesting occurred
on remote beaches in western French Guiana until 2013 (e.g., a high of
4670 nests was found in 2003, with 1,270 mean annual nests from 2002 to
2013), but we were unable to analyze trends because monitoring on these
remote beaches has been reduced since approximately 2010 due to
significant beach erosion and the disappearance of some previously
monitored beaches.
Suriname, Grenada, and Panama each had a single time series
sufficient for trend analysis. For Suriname, we combined datasets from
two beaches, Galibi and Braamspunt, which were monitored between 2001
and 2017. Total nests in Suriname declined by -14.6 percent annually
(sd = 9.6 percent; 95 percent CI = -36.4 to 4.5 percent; f = 0.953;
mean annual nests = 4,586). In Grenada, data on the number of nesting
tracks were collected on Levera beach between 2002 and 2018. There was
a 7.1 percent annual increase in tracks at Levera during that period
(sd = 8.7 percent; 95 percent CI = -10.5 to 25.3 percent; f = 0.827;
mean annual tracks = 895). In Panama, the nest counts at Chiriqui beach
increased by 0.8 percent annually (sd = 7.0 percent; 95 percent CI = -
14.1 to 14.6 percent; f = 0.557; mean annual nests = 4,463) between
2004 and 2017.
In Costa Rica, the four beaches for which we had sufficient data to
analyze annual nest count trends mostly exhibited declining trends.
Tortuguero experienced the steepest decrease, with a median trend of -
10.9 percent annually (sd = 4.2 percent; 95 percent CI = -19.5 to 2.2
percent) for data collected between 1995 and 2017. Nest counts
decreased by -3.8 percent annually at Pacuare beach (sd = 9.3 percent;
95 percent CI = -22.6 to 16.9 percent) between 2004 and 2017, but
increased by 1.8 percent annually (sd = 6.0 percent; 95 percent CI = -
10.8 to 14.2 percent) at the nearby Pacuare Nature Reserve between 1991
and 2017. Nest counts at Estacion la Tortuga deceased slightly, with a
median trend of -0.5 percent annually (sd = 7.0 percent; 95 percent CI
= -15.7 to 13.1 percent) between 2002 and 2017.
For the United States, we analyzed annual nest count trends for
Florida (statewide data collected between 2008 and 2017), three beaches
in Puerto Rico, including Culebra (1984 to 2017), Luquillo-Fajardo
(1996 to 2017), and Maunabo (1999 to 2017), and Sandy Point National
Wildlife Refuge in St. Croix, USVI (1982 to 2017). The median trend for
Florida was a decline of -2.1 percent annually (sd = 13.0 percent; 95
percent CI = -28.3 to 25.5 percent; f = 0.582; mean annual nests =
1,288). Culebra nests decreased by -3.7 percent annually (sd = 5.3
percent; 95 percent CI = -14.9 to 6.8 percent; f = 0.791; mean annual
nests = 153), while nests increased by 15.9 percent annually at
Luquillo-Fajardo (sd = 5.5 percent; 95 percent CI = -7.1 to 15.3
percent; f = 0.805; mean annual nests = 283) and by 7.7 percent
annually at Maunabo (sd = 4.9 percent; 95 percent CI = -2.7 to 17.4
percent; f = 0.945; mean annual nests = 161). In St. Croix, nests
increased by 1.7 percent annually (sd = 4.6 percent; 95 percent CI = -
7.8 to 10.7 percent; f = 0.660; mean annual nests = 399).
These trend data are similar to other recent findings, adding
further
[[Page 48340]]
confidence in declining trends at multiple large nesting aggregations.
Because of concerns about declining nest counts throughout the region,
the National Fish and Wildlife Foundation (NFWF) convened a NW Atlantic
Leatherback Working Group (i.e., the Working Group) to assess recent
nesting data and complete a region-wide trend analysis (NW Atlantic
Leatherback Working Group 2018). The trend analyses conducted by the
Working Group used leatherback nesting data from 23 sites from 14
different nations with at least 10 years of data with consistent
within-site methodology, analyzing data for three time periods: 1990 to
2017, 1998 to 2017, and 2008 to 2017. Our approach to trend analyses
was similar to that used by the Working Group in that both approaches
involved Bayesian analyses of data meeting set criteria. However, the
Team decided against aggregating the data over the DPS due to
incongruity of data collection methods, collection dates and duration,
and reporting. Despite these differences, the overall conclusion was
the same--an overall declining nest trend.
The Working Group found that regional, abundance-weighted trends
were negative for all three time periods and became more negative in
the more recent time series (NW Atlantic Leatherback Working Group
2018). Specifically, overall nesting trends decreased at -4.21 percent
annually from 1990 to 2017 and at -5.37 percent annually from 1998 to
2017, with the most notable decrease (-9.32 percent annually) occurring
during the most recent time frame of 2008 to 2017. While site-level
trends showed variation within and among sites and across the time
periods, overall the sites also reflected the same regional pattern:
More negative trends were apparent during the most recent time frame.
Seven sites had significant positive nesting trends from 1990 to 2017,
but no sites exhibited significant positive trends from 2008 to 2017.
The significant decline observed at Awala-Yalimapo, French Guiana (-
12.95 percent annually from 1990 to 2017, -19.05 percent annually from
1998 to 2017, and -31.26 percent annually from 2008 to 2017), drove the
regional results, but similar significant declines were found at other
nesting beaches for the longer time period, including: St. Kitts and
Nevis (-12.43 percent annually), Tortuguero, Costa Rica (-10.42 percent
annually), Suriname (-5.14 percent annually), and Culebra, Puerto Rico
(-4.61 percent annually). It should be noted that the other nesting
beach in French Guiana (Cayenne) demonstrated an increasing trend (7.44
percent annually from 1990 to 2017 and 8.19 percent annually from 1998
to 2017). However, it exhibited a decreasing trend (-14.21 percent
annually) from 2008 to 2017. While nesting increased over time at
Cayenne, this increase has apparently not resulted from females
shifting from Awala-Yalimapo, as turtles that nest at Cayenne are
genetically distinct (Molfetti et al. 2013) and females tagged in
Awala-Yalimapo are not seen in Cayenne or vice versa (NW Atlantic
Leatherback Working Group 2018).
These modeling results demonstrate that there has been a decline in
NW Atlantic nesting from 1990 to 2017, with the most significant
decreases occurring from 2008 to 2017. Some nesting beaches
demonstrated positive trends for the longer time period. However, none
showed significant increases over the most recent time period. The
cause for the decline is uncertain, but the Working Group identified
anthropogenic sources (e.g., fisheries bycatch), habitat losses, and
changes in life history parameters (such as remigration interval) as
potential drivers of the regional decline. While these results were
taken into consideration by the Team when evaluating the extinction
risk of the NW Atlantic DPS, the Team also performed its own trend
analysis of the data provided to the Team so that the trends were
calculated in a manner consistent with other DPSs. Regardless, both
trend analyses conclude that the NW Atlantic DPS is experiencing a
significant decline in nesting.
In-water abundance studies of leatherback turtles are rare.
Archibald and James (2016) assessed the relative abundance of turtles
at a foraging area off Nova Scotia, Canada, from 2002 to 2015. This
study evaluated opportunistic sightings per unit effort and found a
mean density of 9.8 turtles per 100 km\2\, representing the highest in-
water density of leatherback turtles reported to date. Archibald and
James (2016) concluded that the relative abundance of foraging
leatherback turtles off Canada exhibited high inter-annual variability
but, overall, showed a stable trend from 2002 to 2015. The authors
reported that (at that time) these results were consistent with the
stable or, in some cases, increasing trends reported for contributing
NW Atlantic nesting beaches over the last decade (Dutton et al. 2005;
Girondot et al. 2007; Fossette et al. 2008; McGowan et al. 2008;
Stewart et al. 2011; Rivas et al. 2015). While there were no
indications of a decreasing trend, the results should be interpreted
with caution because of the small study area, opportunistic data
collection, availability bias variance, and lack of understanding of
the relative density outside the study area (Archibald and James 2016).
Despite the declining trend in nesting, productivity parameters for
the DPS are similar to the species' averages (though some may be
declining, as we discuss below). While there is some variation, most
productivity parameters are relatively consistent throughout the DPS.
The overall survival rate for nesting females is relatively high at 85
percent (Pfaller et al. 2018), with mean estimates of 0.70 to 0.99 in
French Guiana (Rivalan et al. 2005, 2008), 0.89 in St. Croix (Dutton et
al. 2005), and 0.89 to 0.96 on the Atlantic coast of Florida (Stewart
et al. 2007, 2014). Remigration intervals range from 1 to 11 years
(Schulz 1975; Boulon et al. 1996; Chevalier and Girondot 1998;
Hilterman and Goverse 2007; Eckert et al. 2012; Stewart et al. 2014;
Rivas et al. 2016; Garner et al. 2017). In St. Croix and St. Kitts, the
median remigration interval appears to be increasing (4.5 years; K.R.
Stewart, The Ocean Foundation and C. Lombard, USFWS, pers. 2019; K.M.
Stewart, Ross University School of Veterinary Medicine and St. Kitts
Sea Turtle Monitoring Network, pers. comm., 2019). Averaging all
available data, the mean remigration interval for the DPS is 2.7 years,
rounded to 3 years for use in our calculation of the index of nesting
female abundance. Average clutch frequency per nesting season ranges
from 3.6 to 8.3 throughout the region, with an overall mean of 5.5
nests per season, interspersed with 9 to 10 day internesting intervals
(Eckert et al. 2015; Garner et al. 2017). Recent records indicate that
nesting females deposit 80 to 88 eggs per clutch. However, an early
study by Carr and Ogren (1959) reported only 67 eggs per clutch.
Hatching success is highly variable for nests that remain in situ, even
for those that are viable and do not experience significant inundation
or predation, with estimates as low as 8.9 percent in Costa Rica
(Tro[euml]ng et al. 2007) and 10.6 percent in Suriname (Hilterman and
Goverse 2007) and as high as 93.4 percent in Florida (Perrault et al.
2012). Overall, hatching success is estimated at approximately 50
percent (Eckert et al. 2012). Hatchling sex ratios often exhibit a
female bias, but less so than for other sea turtle species, with
estimated production of anywhere from 30 to 100 percent females in
Suriname, Tobago, Colombia, and Costa Rica (Mrosovsky et al. 1984;
Dutton et al. 1985; Godfrey et al. 1996; Leslie et al. 1996; Mickelson
and
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Downie 2010; Pati[ntilde]o-Mart[iacute]nez et al. 2012). However, the
proportion of females documented in foraging individuals and strandings
ranges from 57 to 70 percent (Murphy et al. 2006; James et al. 2007;
TEWG 2007), and the ratio of females to males during an individual
breeding season is thought to be closer to 1:1 (Stewart and Dutton
2014).
We conclude that the DPS exhibits a declining nest trend. In
addition, there are indications of decreased productivity within the
DPS. In St. Croix, one of the most thoroughly monitored nesting beaches
in this DPS, the data from 1981 to 2010 indicate that hatching success
and clutch frequency are declining and remigration intervals are
increasing (Garner et al. 2017). Overall, we have a high degree of
confidence in the decreasing nest trend and productivity metrics for
this DPS, due to the large amount of data available from the largest
nesting aggregations. We acknowledge that data are not available from
all nesting beaches, but the data that we have relied upon is the best
available and meets established standards. The declining trends reflect
reduced nesting female abundance. In addition, longer remigration
intervals and/or reduced clutch frequencies may play a role in this
decline. The decline reflects a reduction in productivity that places
the DPS at risk given the magnitude and duration of the decreasing
trend.
Spatial Distribution
The DPS has a broad spatial distribution for both foraging and
nesting. There is significant genetic population structure, with
subpopulations connected via various levels of gene flow and
metapopulation dynamics. Tagging and telemetry studies indicate
considerable mixing of leatherback turtles among nesting beaches and at
multiple foraging areas throughout the North Atlantic Ocean.
Nesting is widespread throughout the NW Atlantic beaches, occurring
primarily as scattered, small aggregations throughout the Wider
Caribbean, but with larger concentrations of nesting activity at
certain sites in Trinidad, French Guiana, Suriname, Trinidad, Colombia,
Panama, Costa Rica, Puerto Rico, St. Croix, and Florida (Horrocks et
al. 2016).
Genetic sampling in the NW Atlantic DPS has been generally
extensive with good coverage of large populations in this region.
However, sampling from some smaller Caribbean nesting aggregations is
absent, and there are gaps in sampling or analysis for nesting sites
along the coasts of South and Central America (e.g., Guyana, Venezuela,
Colombia, and Panama). A comprehensive survey of genetic population
structure in the Atlantic Ocean included large sample sizes from five
nesting populations representative of the DPS and analysis of longer
mtDNA sequences in combination with an array of 17 nuclear
microsatellite DNA loci (Roden and Dutton 2011; Dutton et al. 2013).
The microsatellite data revealed fine-scale genetic differentiation
among neighboring subpopulations (Dutton et al. 2013): Trinidad, French
Guiana/Suriname, Florida, Costa Rica, and St. Croix. The mtDNA data
failed to find significant differentiation between Florida and Costa
Rica or between Trinidad and French Guiana/Suriname. However, Dutton et
al. (2013) show that the mtDNA sequence variation had relatively low
statistical power to detect fine scale structure compared to the
microsatellite DNA loci. The mtDNA homogeneity between Costa Rica and
Florida, with differentiation demonstrated at nuclear DNA loci,
suggests that Costa Rica may be the source of founders for the Florida
population via one or multiple recent colonization events, likely
indicating historic connectivity rather than ongoing demographic
connectivity (Dutton et al. 2013). Likewise the French Guiana/Suriname
and Trinidad populations were undifferentiated with mtDNA likely
indicating historic connectivity. However, microsatellite DNA reveal
fine-scale genetic structure that is consistent with tagging studies
demonstrating a lack of nesting female movement between the two nesting
aggregations (TEWG 2007). Significant genetic differentiation has also
been reported for Martinique and Guadeloupe and the mainland French
Guiana rookery (Molfetti et al. 2013). St. Croix likely represents a
broader Northern Caribbean subpopulation of the NW Atlantic population
that includes multiple neighboring island nesting aggregations in the
USVI and Puerto Rico. However, sampling and analysis would be required
to determine extent of fine scale structuring (NMFS unpublished data;
Dutton et al. 2013). The Costa Rica (Tortuguero and Gandoca) and Guiana
(French Guiana and Suriname) nesting aggregations are distinct
subpopulations based on microsatellite and mtDNA results (Dutton et al.
2013), but information on tag returns indicates movement of nesting
females between adjacent beaches of Panama, Colombia, Venezuela and
Guyana. Therefore, these nesting aggregations have ``fuzzy''
boundaries, likely a result of flexible natal homing. Nesting females
use beaches up to 400 km apart between nesting seasons (Tro[euml]ng et
al. 2004; Chac[oacute]n-Chaverri and Eckert 2007) and up to 463 km
apart within the same nesting season (Stewart et al. 2014). Additional
sampling of the remaining nesting sites will be required to determine
the extent of fine-scale structuring within the NW Atlantic DPS.
However, the available science indicates significant substructure
within the DPS.
Tagging studies indicate individual movement and gene flow among
nesting aggregations. This is facilitated by the species' flexible
natal homing, i.e., philopatry to a region, rather than a specific
beach. In adjacent nesting sites in French Guiana and Suriname, five to
six percent of nesting females were observed to shift from one site to
the other within a season (TEWG 2007), while Schulz (1971) reported
this proportion to be slightly higher at 8.5 percent. In contrast, 35
percent of nesting females in Gandoca, Costa Rica, were estimated to
nest at sites other than the study site during an individual season
(Chac[oacute]n-Chaverri and Eckert 2007). The predisposition of nesting
females to stray within a nesting season may be influenced by the
proximity of alternative nesting sites within a range of approximately
200 km (Horrocks et al. 2016). However, even within a given nesting
season, females have been observed to move as far as 369 km (Grenada),
463.5 km (Florida), and 532 km (Dominica) from their original location
(Horrocks et al. 2016). Among nesting seasons, interchange between
nesting locations also appears to be frequent and wide-ranging, with
maximum distance separating two nesting sites for an individual female
recorded as 1,849 km over an 8-year span (Horrocks et al. 2016).
Genetic studies have revealed that turtles from different nesting
aggregations use the same foraging areas. Analyzing 684 longline
bycatch samples from across the NW Atlantic in a mixed stock analysis
and microsatellite assignment, Stewart et al. (2016) found that
leatherback turtles from Costa Rica were caught in a higher proportion
in the GOM (43 percent) compared to the Northeast Distant fishing zone,
an area in the northwestern Atlantic Ocean (6 percent), while turtles
from Trinidad and French Guiana comprised 54 percent of bycatch in the
GOM and 93 percent in the Northeast Distant fishing zone. A study of
turtles foraging off Nova Scotia, Canada, similarly assigned most (82
percent) of the 288 sampled turtles to Trinidad (n = 164) and French
[[Page 48342]]
Guiana (n = 72), with 15 percent (n = 44) from Costa Rica, and the
remainder from St. Croix (n = 7) and Florida (n = 1; Stewart et al.
2013). These proportions generally represent the relative population
sizes for these breeding populations. Microsatellite DNA assignment of
wild captured or stranded males (n = 122) throughout the NW Atlantic
and Mediterranean found that all males originated from NW Atlantic
nesting aggregations (Trinidad: 55 percent, French Guiana: 31 percent,
and Costa Rica: 14 percent; Roden et al. 2017). No turtles were
identified from St. Croix or Florida. One turtle that stranded in
Turkey was assigned to French Guiana, while strandings in France were
assigned to Trinidad or French Guiana (Roden et al. 2017).
The mixing of nesting aggregations at foraging areas is also
supported by several tagging and/or satellite telemetry projects,
conducted in U.S. waters (Murphy et al. 2006; LPRC 2014; Dodge et al.
2014, 2015; Aleksa et al. 2018), Canada (James et al. 2005a, 2005b,
2005c, 2006b, 2007; Bond and James 2017), Atlantic Europe and
Mediterranean (Doyle et al. 2008; Sonmez et al. 2008), and on nesting
beaches of various nations (Hildebrand 1987; Hays et al. 2004;
Ferraroli et al. 2004; Eckert 2006; Eckert et al. 2006; Hays et al.
2006; TEWG 2007; Sonmez et al. 2008; Evans et al. 2008; Fossette et al.
2010a, 2010b; Richardson et al. 2012; Bailey et al. 2012; Stewart et
al. 2014; Fossette et al. 2014; Horrocks et al. 2016; Chambault et al.
2017). For instance, turtles from Nova Scotian foraging grounds were
tracked to nesting areas off Colombia, Trinidad, Guyana, and French
Guiana (Bond and James 2017). The reverse has also been demonstrated:
some leatherback turtles from the western Atlantic undertake annual
migrations to Canadian waters to forage (James et al. 2005c),
exemplified by post-nesting adults tracked to the waters off Nova
Scotia from a variety of nesting locations, including French Guiana and
Trinidad (Fossette et al. 2014), Costa Rica, Panama (Evans et al.
2008), and Anguilla (Richardson et al. 2012). The eastern and western
GOM also provide foraging areas for this DPS (Aleksa et al. 2018), as
observed from tracks of post-nesting turtles from Florida (Hildebrand
1987), Costa Rica (Tortuguero, Gandoca), and Panama (Chiriqu[iacute]
Beach; Evans et al. 2008; Evans et al. 2012). Evans et al. (2008)
suggested that the GOM may represent a significant foraging ground for
leatherback turtles from the Caribbean coast of Central America.
High use foraging areas may be identified through available
telemetry data, but the migration routes to those areas may vary.
Ferraroli et al. (2004) tracked leatherback turtles from French Guiana
and found turtles dispersed widely throughout the North Atlantic but
mostly followed two dispersion patterns: (1) Moving north to the Gulf
Stream area, where they started following the general ocean
circulation; and (2) traveling east, swimming mostly against the North
Equatorial Current. Fossette et al. (2014) found a relatively broad
migratory corridor when turtles departed their nesting sites in French
Guiana/Suriname, and their movements overlapped with turtles from
Grenada and Trinidad. Fossette et al. (2010a, 2010b) found that turtles
tracked from nesting beaches in French Guiana, Suriname, and Grenada
and turtles caught in waters off Nova Scotia and Ireland displayed
three distinct migration strategies: (1) Heading northwest to fertile
foraging areas off the Gulf of Maine, Canada, and GOM; (2) crossing the
North Atlantic Ocean to areas off western Europe and Africa; and (3)
residing between northern and equatorial waters. Essentially, tagging
data coupled with satellite telemetry data indicate that leatherback
turtles of the NW Atlantic DPS use the entire North Atlantic Ocean for
foraging and migration (TEWG 2007).
Although adults forage at multiple areas throughout the North
Atlantic Ocean (Fossette et al. 2014), the range of juvenile
leatherback turtles may be more restricted. Using an active movement
model, Lalire and Gaspar (2019) found that most juveniles originating
from nesting beaches in French Guiana and Suriname cross the Atlantic
Ocean at mid-latitudes with north-south seasonal migrations; after
several years, they reach the coasts of Europe and North Africa. Eckert
(2002) reviewed the records of nearly 100 sightings of juvenile (less
than 100 cm curved carapace length (CCL)) leatherback turtles and
determined they are generally found in waters warmer than 26 [deg]C,
suggesting that the first portion of their life is spent in tropical
and subtropical waters. After exceeding 100 cm CCL, distribution
extends into cooler waters (as low as 8 [deg]C), which is considered to
be the primary habitat for the species (Eckert 2002).
The wide distribution of nesting and foraging areas likely buffers
the DPS against local catastrophes or environmental changes. The fine-
scale population structure, with movement of individuals and genes
among nesting aggregations, indicates that the DPS has the capacity to
withstand other catastrophic events.
Diversity
The NW Atlantic DPS exhibits spatial diversity, as demonstrated by
insular and continental nesting, multiple diverse foraging areas, and
moderate genetic diversity. The DPS nests along both continental and
insular coastlines. Nesting beach habitat also shows considerable
diversity, ranging from coarse-grained, sandy beaches to silty,
ephemeral shorelines whose dynamics are influenced by estuarine input.
The breadth and, in some cases, transiency, of suitable nesting habitat
in the western North Atlantic may contribute to consistent, low-level
flexibility in natal homing, both within and among reproductive seasons
(Br[auml]utigam and Eckert 2006), and this flexibility is thought to
surpass that of other sea turtle species (TEWG 2007).
This DPS exhibits some temporal variation in nesting. Nesting
generally begins in March or April, peaks in May or June, and ends in
July or August (Eckert et al. 2012). In French Guiana, a second small
nesting peak was documented in Awala-Yalimapo during December and
January. However, the number of nests deposited during that time frame
decreased from 700 in 1986/1987 to 40 in 1992/1993, and now only a
small number of individuals are observed to nest during that time
(Girondot et al. 2007). Some evidence indicates that the timing of
nesting may be modulated by environmental characteristics distant from
the nesting beach, such as water temperatures at foraging grounds
(Neeman et al. 2015).
The foraging strategies are also diverse, with turtles using
coastal and pelagic waters throughout the entire North Atlantic Ocean
(Fossette et al. 2014). Foraging habitats include temperate waters of
the GOM, North Central Atlantic Ocean, northwestern shelf (United
States and Canada), southeastern U.S. coast, the Mediterranean Sea, and
northeastern shelf (Europe; TEWG 2007). Some post-nesting females also
remain in tropical waters (Fossette et al. 2010). Overall, leatherback
turtles in the North Atlantic Ocean appear to have a diverse array of
foraging habitat available.
Genetic diversity of the DPS is moderate, with six mtDNA haplotypes
(Dutton et al. 2013). In St. Croix, a unique haplotype occurs at high
frequency. The Florida and Costa Rica nesting aggregations each possess
one unique, low frequency haplotype.
Based upon this information, we conclude that nesting location and
habitat are diverse, providing some level of resilience against short-
term spatial and temporal changes in the
[[Page 48343]]
environment. However, high-abundance nesting occurs only at a few
locations (e.g., Trinidad, French Guiana, and Panama). The foraging
diversity likely provides resilience against local reductions in prey
availability or catastrophic events, such as oil spills, by limiting
exposure to a limited proportion of the total population. Moderate
genetic diversity may provide the DPS with the raw material necessary
for adapting to long-term environmental changes, such as cyclic or
directional changes in ocean environments due to natural and human
causes (McElhany et al. 2000; NMFS 2017). We conclude that such
diversity provides some level of resilience to threats for this DPS.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
Destruction and modification of leatherback turtle nesting habitat
results from a variety of activities including coastal development and
construction; beach erosion and inundation; placement of erosion
control and nearshore shoreline stabilization structures and other
barriers to nesting; beachfront lighting; vehicular and pedestrian
traffic; beach sand placement; sand extraction; removal of native
vegetation; and planting of non-native vegetation (Lutcavage et al.
1997; Bouchard et al. 1998; USFWS 1999; Dow et al. 2007; Eckert et al.
2012; NMFS and USFWS 2013). As a result, most nesting beaches are
severely degraded by such activities that continue to cause adverse
impacts throughout the range of the DPS.
Coastal Development and Construction
In many areas, nesting habitat is under constant threat from
coastal development and construction (Dow et al. 2007; Crespo and Diez
2016; Flores and Diez 2016). Coastal development impacts include
construction of buildings and pilings on the beach; increased erosion;
artificial lighting; pollution; recreational beach equipment and other
obstacles on the beach; beach driving; increased human disturbance; and
mechanized beach cleaning (Lutcavage et al. 1997; USFWS 1999; Hernandez
et al. 2007; Dow et al. 2007; Trinidad and Tobago Forestry Division et
al. 2010; Flores and Diez 2016). Driftwood found on nesting beaches
also has the potential to alter nesting beach habitat and obstruct
nesting females and hatchlings, as seen in Gandoca, Costa Rica
(Chac[oacute]n-Chaverri and Eckert 2007). These threats impact nesting
habitat by reducing the amount and quality of suitable beaches,
preventing or deterring nesting females from using optimal locations,
destroying nests, eggs, and hatchlings, and preventing hatchlings from
successfully reaching the ocean (USFWS 1999; Chac[oacute]n-Chaverri and
Eckert 2007; Hernandez et al. 2007; Witherington et al. 2014).
Development involving the construction of tall buildings and clearing
of vegetation can also alter sand temperatures and skew sex ratios
(Gledhill 2007).
Development occurs to varying extents throughout the range of the
DPS, but most leatherback nesting occurs in proximity to some coastal
development. The Florida shoreline is extensively developed outside
wildlife refuges (Witherington et al. 2011). In Grenada, nearly 20
percent of all nests surveyed from 2001 to 2005 occurred in an area
affected by development, resulting in ongoing run-off onto nesting
beaches (Maison et al. 2010). In Trinidad, increasing rural and
commercial beachfront development is a concern, especially on the east
coast where the main nesting beaches are located (Trinidad and Tobago
Forestry Division et al. 2010), including Grande Riviere, the largest
nesting aggregation of this DPS. Likewise, several Tobago beaches are
densely developed for commercial tourism, resulting in reduced turtle
access to potential nesting sites due to buildings, umbrellas, and
other recreational equipment (Trinidad and Tobago Forestry Division et
al. 2010). Development in Puerto Rico, in particular Playa Grande-El
Paraiso (i.e., Dorado Beach, which is considered to be the most
important nesting beach in Puerto Rico), is also a notable concern
(Crespo and Diez 2016; Flores and Diez 2016). There, ecosystems
continue to be threatened by coastal development, even though the
coastal zone is protected by the Maritime-Terrestrial Zone designation
(i.e., Coastal Public Trust Lands; Flores and Diez 2016).
Coastal development likely influences leatherback nest placement
and subsequent nest success, which is the percentage of nesting
attempts (i.e., emergences onto the beach) that result in eggs being
deposited. On Margarita Island, Venezuela, Hernandez et al. (2007)
found that leatherback nesting aggregated towards the portions of the
beach with fewer risk factors, such as light pollution and
concentrations of beach furniture. This change in nesting behavior
resulted in females nesting in less optimum areas (e.g., areas with
lower hatching success), thus affecting the reproductive potential of
leatherback turtles in this region.
The magnitude of development is also changing in some areas, where
nest placement and success may be affected in the future. For instance,
the area around Cayenne, French Guiana, is undergoing increased
urbanization and recreational use (Fossette et al. 2008). In recent
years, nesting has increased at Cayenne and eastern beaches compared to
the western Awala-Yalimapo beaches (R[eacute]serve Naturelle de l'Amana
data in Berzins 2018 and KWATA data in Berzins 2018). As such, more
nesting in French Guiana is exposed to coastal development and the
associated threats, and these threats are likely to continue and
increase.
Beach Erosion and Inundation
While erosion is often intensified due to anthropogenic influences,
natural features in some areas result in high erosion rates and
unstable beaches, thus affecting leatherback nesting. For instance, the
Maroni River influence in the Guianas (French Guiana especially) has
resulted in highly dynamic and unstable beaches, with shifting mudflats
making nesting habitat unsuitable (Crossland 2003; Goverse and
Hilterman 2003; Fossette et al. 2008). Beaches are often created and
lost along the coast of French Guiana (Kelle et al. 2007). For example,
remote beaches in western French Guiana experience significant beach
erosion and several disappeared, reducing or preventing monitoring (and
likely nesting). In Suriname, Braamspunt Beach at the mouth of the
Suriname River is moving west, out of the established Wia Wia Nature
Reserve and may disappear in the next several years (M. Hiwat, WWF,
pers. comm., 2018). This is significant in that Braamspunt is currently
the main nesting beach in Suriname. The second highest nesting area in
Suriname, Galibi Beach, is also experiencing significant erosion and
becoming narrower. Similar beach erosion is occurring in Guyana, as
well as in Trinidad and Tobago (Reichart et al. 2003; Trinidad and
Tobago Forestry Division et al. 2010). At some Trinidad and Tobago
nesting sites (e.g., Fishing Pond, Matura, Grande Riviere, and Great
Courland Bay), rivers emerge onto nesting beaches and create additional
erosion during the nesting season (Godley et al. 1993; Lee Lum 2005),
intensifying nest loss (up to 35 percent of nests; Trinidad and Tobago
Forestry Division et al. 2010).
Seasonal erosion also occurs at most Caribbean nesting beaches. A
survey of Wider Caribbean Regions found that erosion/accretion was the
highest threat to nesting habitat (Dow et al. 2007). For example, at
Playa Gandoca, Costa Rica, erosion from strong coastal drift currents
is thought to be one of the largest obstacles to hatching success,
destroying greater than 10 percent of all
[[Page 48344]]
nests laid in some years (Chac[oacute]n-Chaverri and Eckert 2007). In
2006 and 2007, coastal erosion and inundation accounted for 33 to 42
percent of nest loss in southern Panama and 29 to 48 percent on
Caribbean Colombia beaches (Pati[ntilde]o-Mart[iacute]nez et al. 2008).
Inundation of nests is also a concern. Leatherback turtles
generally nest closer to the water than other sea turtles (Caut et al.
2010). If nests are laid too close to the high tide line, they are
subjected to erosion and inundation, which can result in egg mortality
from suffocation or curtailed embryonic development (Chac[oacute]n-
Chaverri and Eckert 2007; Caut et al. 2010). This inundation phenomenon
occurs on multiple nesting beaches and is particularly of concern in
areas with high tidal influence and dynamic coastlines. On Krofajapasi
beach in Suriname, 31.6 percent of nests laid by females were below the
spring high tide level and determined to be ``doomed'' clutches (Dutton
and Whitmore 1983). Similarly, in Gandoca, Costa Rica, 37 percent of
nests from 1990 to 2004 were laid in the low tide zone and would have
been inundated if not relocated (Chac[oacute]n-Chaverri and Eckert
2007). In St. Croix, 43 percent of the nests (with a range of 25 to 68
percent) were considered to be ``doomed'' each season (McDonald-Dutton
et al. 2001), but beginning in 1983, all doomed clutches were relocated
to improve hatching success (Dutton et al. 2005). Without intervention,
these nests would likely have been lost. On Awala-Yalimapo, French
Guiana, 27 of 89 nests were overlapped by tide at least once during the
incubation period, and the hatching success was on average
significantly lower in overwashed nests (Caut et al. 2010). Observed
mortality was 100 percent in the intertidal zone at sites along the
coasts of Panama and Colombia, with an overall nest loss by erosion and
inundation ranging from 16 to 48 percent among three major nesting
sites (Pati[ntilde]o-Mart[iacute]nez et al. 2008). While levels of
inundation and resulting declines in hatching success have been noted
at multiple sites throughout the range of the NW Atlantic DPS, the
specific impacts of inundation may be variable. Hilterman and Goverse
(2007) noted that leatherback nests can tolerate relatively high levels
of inundation, so hatching may still be successful despite proximity to
the tide line. Because of this, and because it may affect natural sex
ratios (Mrosovsky and Yntema 1980), the relocation of nests susceptible
to inundation was abandoned in 2002 in Suriname (Hilterman and Goverse
2007); only nests directly threatened by beach erosion are relocated,
under certain circumstances. Other nations still relocate nests to
reduce the impacts of erosion. However, as mentioned, such practices
may result in cooler nests and affect sex ratios (Spanier 2008). While
eggs relocated to hatcheries could have been lost under natural
circumstances, due to coastal erosion and inundation in some areas
(Dutton and Whitmore 1983, Chac[oacute]n-Chaverri and Eckert 2007),
hatching success in relocated nests is often lower than in situ nests
(Revuelta et al. 2014; Valentin-Gamazo et al. 2018; Florida Department
of Environmental Protection unpublished data 2018).
Such naturally dynamic areas make it difficult to protect nesting
beach habitat and accurately assess leatherback nesting trends. This is
particularly noteworthy given that nesting females use high energy,
erosion-prone beaches, which often result in high nest loss
(Chac[oacute]n-Chaverri and Eckert 2007; TEWG 2007; Spanier 2008;
Trinidad and Tobago Forestry Division et al. 2010). However,
leatherback turtles in the Guianas seem to have adapted to this
constant geomorphological change of beaches. When new beaches develop,
they may be colonized within months by nesting females, who take
advantage of the fresh, clean sand (or seashells, in Guyana) and
absence of entangling or deep-rooted beach vegetation (TEWG 2007).
Nest site selection by leatherback turtles is still poorly
understood (Maison et al. 2010), but nesting females may be changing
their nesting patterns due to erosion. Spanier (2008) found that
nesting females at Playa Gandoca, Costa Rica, appear to actively select
nest sites that are not undergoing extensive erosion, with slope
considered to be the cue for site selection. A similar result was found
on Grande Riviere, Trinidad, with a nesting shift from east to west
throughout the season as an apparent response to erosion on the eastern
end of the nesting beach (Lee Lum 2005). Further, Maison et al. (2010)
studied nest placement in Grenada and discovered that leatherback
turtles seemed to respond to the accretion of the north facing beach
and erosion of the east facing beach in 2005 by nesting more often on
the north facing beach. If erosion is increasing in existing nesting
locations, nesting may occur in areas with lower success rates, thus
affecting productivity. In addition, leatherback nests are deeper than
those of other sea turtles; water content and salinity typically
increase with depth, leading to a decrease in sea turtle hatching
success (Foley et al. 2006).
Erosion Control, Nearshore Shoreline Stabilization Structures, and
Other Barriers
A widespread strategy to reduce coastal erosion is to construct
erosion control structures. However, these structures reduce the amount
of available nesting habitat. Also, when beachfront development occurs,
the site is often engineered to protect the property from erosion. This
type of shoreline engineering, collectively referred to as beach
armoring, includes sea walls, rock revetments, riprap, sandbag
installations, groins and jetties. Beach armoring can result in
permanent loss of a nesting beach through accelerated erosion and
prevention of natural beach/dune accretion. These impacts can prevent
or hamper nesting females from accessing suitable nesting sites (USFWS
1999). Clutches deposited seaward of these structures may be inundated
at high tide or washed out entirely by increased wave action near the
base of the erosion control structures. As these structures fail and
break apart, they spread debris on the beach, thus creating additional
impacts to hatchlings and nesting females.
In the southeastern United States, numerous erosion control
structures that create barriers to nesting have been constructed. In
Florida, the total amount of existing and potential future armoring
along the coastline is approximately 24 percent (164 miles; FDEP, pers.
comm., 2018). This assessment of armoring does not include other
structures that are a barrier to sea turtle nesting, such as dune
crossovers, cabanas, sand fences, and recreational equipment.
Additionally, jetties have been placed at many ocean inlets in the
United States to keep transported sand from closing the inlet channel.
The installation of jetties resulted in lower loggerhead and green
turtle nesting density updrift and downdrift of the inlets, leading
researchers to propose that beach instability from both erosion and
accretion may discourage turtle nesting (Witherington et al. 2005).
Leatherback nesting near jetties and inlets is low, possibly reflecting
their avoidance of such areas. There are some efforts, such as the
Coastal Construction Control Line Program, that provide protection for
Florida's beaches and dunes while allowing for continued use of private
property. However, armoring structures on and adjacent to the nesting
beach continue to be permitted and constructed on the nesting beaches
of Florida, as in other nations where the DPS nests.
Due to erosion, beach nourishment is a frequent activity in some
developed
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areas, and many beaches are on a periodic nourishment schedule. Beach
nourishment may result in direct burial and disturbance to nesting
females, if conducted during the nesting season. It may also result in
changes in sand density, beach hardness, beach moisture content, beach
slope, sand color, sand grain size, sand grain shape, and sand grain
mineral content, if the placed sand is dissimilar from the original
beach sand (Nelson and Dickerson 1988; USFWS 1999). These changes can
affect nest site selection, digging behavior, incubation temperature
(and hence sex ratios), gas exchange parameters within incubating
nests, hydric environment of the nest, hatching success and hatchling
emerging success (Lutcavage et al. 1997; Steinitz et al. 1998; Ernest
and Martin 1999; USFWS 1999; Rumbold et al. 2001; Brock et al. 2009).
On severely eroded sections of beach, where little or no suitable
nesting habitat previously existed, beach nourishment has been found to
result in increased nesting (Ernest and Martin 1999). However, on most
beaches in the southeastern United States, nesting success typically
declines for the first year or two following nourishment, even though
more nesting habitat is available for turtles (Trindell et al. 1998;
Ernest and Martin 1999; Herren 1999; Brock et al. 2009). Further,
nourishment projects result in heavy machinery, pipelines, increased
human activity and artificial lighting on the project beach, further
affecting nesting females and beach habitat. Overall, the impacts of
beach nourishment to this DPS are not as widespread as other threats to
nesting habitat, as Dow et al. (2007) found that only four nations
(Anguilla, Cuba, Mexico, and United States) reported frequent or
occasional beach nourishment.
Artificial Lighting
Coastal development also contributes to habitat degradation by
increasing light pollution, which can result in hatchling and nesting
female disorientation, altering behavior and leading to mortality. In
Florida, from 2013 to 2017, a total of 341 leatherback nests
(representing the whole or majority of hatchlings in the nest) and five
nesting females were disoriented (FWC unpublished data 2018).
Artificial lighting ranked as the third highest threat to nesting/
hatching turtles in the Wider Caribbean Region (Dow et al. 2007). For
example, urban development is significant in Puerto Rico, with light
pollution (as well as coastal erosion and deforestation) occurring near
leatherback nesting beaches (Crespo and Diez 2016). Fortunately, some
of the major nesting beaches in this DPS are located in comparatively
remote areas, and large-scale development is currently less of an issue
there (Trinidad and Tobago Forestry Division et al. 2010; NMFS and
USFWS 2013). That said, even within the same country, light pollution
is variable. Fossette et al. (2008) reported that in French Guiana,
light pollution from residential areas is a problem at Cayenne Beach,
but it is not an issue at Awala-Yalimapo. Similarly, lighting is not a
significant problem on nesting beaches in Trinidad, but is a concern in
Tobago (Trinidad and Tobago Forestry Division et al. 2010). With the
risk of increased development in some of these relatively remote areas,
additional light pollution is anticipated, and disorientation of
hatchlings and adults from such lighting may become a bigger problem.
In Costa Rica, beachfront lighting is increasing and may become
problematic at Gandoca Beach (Chac[oacute]n-Chaverri and Eckert 2007)
and Tortuguero (de Haro and Tro[euml]ng 2006).
Light pollution has been managed to some extent (Witherington et
al. 2014). Lighting in Florida is regulated by multiple rules and
regulations including Florida statutes, the Florida Building Code, and
local lighting ordinances (Witherington et al. 2014). In addition, the
Florida Department of Transportation and local governments have adopted
lighting-design standards. A total of 82 municipalities in Florida have
adopted lighting ordinances to minimize the impact of lighting on
adjacent sea turtle nesting beaches (Witherington et al. 2014).
However, compliance and enforcement is lacking in some areas. Further,
lighting away from areas covered by beachfront ordinances is
unregulated, resulting in urban glow. Although outreach and
conservation programs control the impacts of lighting in some other
locations, such as Costa Rica, Mexico, and Puerto Rico (Lutcavage et
al. 1997; Crespo and Diez 2016), a majority of nations do not have
regulations in place.
Sand Extraction
Extracting sand from nesting beaches for construction projects has
a detrimental effect on the amount of available nesting beach habitat
and also accelerates erosion (resulting in the aforementioned
associated impacts). Sand mining occurs in most Wider Caribbean nations
to varying extent and frequency (Dow et al. 2007). In particular, beach
sand mining has been extensive at Matura Bay and Blanchisseuse in
Trinidad (Trinidad and Tobago Forestry Division et al. 2010). Some
nations regulate sand mining: In St. Lucia, the Conservation and
Management Act of 2014 requires a certificate of environmental approval
for projects removing sand from nesting beaches.
Removal of Native Vegetation
In some nations, upland deforestation and the resultant deposition
of debris and garbage can destroy or modify nesting beaches. The debris
can block access of gravid (pregnant) females and fatally trap emergent
hatchlings (Chac[oacute]n-Chaverri and Eckert 2007). The accumulation
of logs reduces the amount of available nesting habitat, possibly
forcing leatherback females to nest in suboptimal locations (TEWG
2007). Deforestation due to coastal development is a notable concern in
Puerto Rico (Crespo and Diez 2016).
Vehicular Traffic
Beach driving also occurs in most nations throughout the range of
this DPS (Chac[oacute]n-Chaverri and Eckert 2007; Dow et al. 2007;
Trinidad and Tobago Forestry Division et al. 2010). In the United
States, vehicular driving is allowed on certain beaches in Florida
(e.g., Duval, St. Johns, and Volusia Counties). Beach driving reduces
the quality of nesting habitat in several ways. Vehicle ruts on the
beach can prevent or impede hatchlings from reaching the ocean
following emergence from the nest (Mann 1977; Hosier et al. 1981; Cox
et al. 1994; Hughes and Caine 1994). Sand compaction by vehicles
hinders nest construction and hatchling emergence from nests (Mann
1977; Gledhill 2007). Vehicle lights and vehicle movement on the beach
after dark can deter females from nesting and disorient hatchlings.
Additionally, vehicle traffic contributes to erosion, especially during
high tides or on narrow beaches where driving is concentrated on the
high beach and foredune.
Vegetation
Beach vegetation (native and non-native) can affect turtle nesting
productivity by obstructing nest construction and potentially drying
the sand (resulting in egg chamber collapse). Vegetation can form
impenetrable root mats that can invade and desiccate eggs and affect
developing embryos, impede hatchling emergence, and trap hatchlings
(Conrad et al. 2011). Non-native vegetation has invaded many coastal
areas and often outcompetes native plant species (USFWS 1999). The
occurrence of exotic vegetation (or loss of native vegetation) was
recognized as a medium-ranked threat in many Wider Caribbean nations
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(Dow et al. 2007). The Australian pine (Casuarina equisetifolia) is
particularly harmful to sea turtles (USFWS 1999). Australian pines
cause excessive shading of the beach that would not otherwise occur.
Studies of loggerhead turtles in Florida suggest that nests laid in
shaded areas are subjected to lower incubation temperatures, which may
alter the natural hatchling sex ratio (Marcus and Maley 1987; Schmelz
and Mezich 1988). Fallen Australian pines limit access to suitable nest
sites and can entrap nesting females (Reardon and Mansfield 1997). The
shallow root network of these pines can interfere with nest
construction (Schmelz and Mezich 1988). Dense stands of Australian pine
have overtaken many coastal areas throughout central and south Florida.
While non-native vegetation can affect nesting habitat throughout
the range of the DPS, native vegetation can also affect productivity.
For instance, at Sandy Point, St. Croix, changing erosion-accretion
cycles led to native Ipomoea pes-caprae, a creeping vine, extending
into the nesting area in some years. Nesting females at Sandy Point
typically avoided nesting in vegetation, resulting in more nests laid
near the high-tide line (Conrad et al. 2011). As a result, Ipomoea pes-
caprae decreased nest productivity by reducing leatherback hatching and
emergence (percentage of hatchlings that emerge from the nest) success
rates (Conrad et al. 2011).
Mitigations to Habitat Modification
Nesting habitat disruptions are minimized in some areas. Several
areas in the NW Atlantic DPS range are under U.S. Federal ownership as
National Wildlife Refuges in Florida (Archie Carr and Hobe Sound),
Puerto Rico (Culebra and Vieques) and St. Croix (Sandy Point). Beaches
in some Wider Caribbean countries are also protected. In Trinidad,
Matura and Fishing Pond beaches were declared Prohibited Areas in 1990,
and the nesting beach at Grande Riviere in 1997. In 1998, the Amana
Nature Reserve, which includes Awala-Yalimapo beach and a 30 m wide
marine fringe, was established in French Guiana. In Suriname, the Wia
Wia Nature Reserve was implemented in 1961 (amended and enlarged in
1966 to protect sea turtles), and in 1969, the Marowijne beaches were
declared a sanctuary (the Galibi Nature Reserve; Schulz 1971). In
addition, Tortuguero National Park, Costa Rica, was established in 1976
to protect nesting habitat (Bjorndal et al., 1999). Terrestrial habitat
in these areas is therefore protected from the above threats to some
extent. USFWS and NMFS also designated as critical habitat for
leatherback turtles the nesting beaches at Sandy Point, St. Croix (43
FR 43688; September 26, 1978) and surrounding marine waters (44 FR
17710; March 23, 1979), which benefits the turtles in this DPS.
However, if ESA protections did not continue (i.e., if this species
were no longer listed), these protections would be lost.
Marine Habitat Modifications
In the marine environment, habitat threats include anthropogenic
noise and offshore lighting. We discuss other threats to marine habitat
and prey (e.g., marine pollution, oil exploration, and climate change)
in later sections. Anthropogenic noise impacts the marine habitat of
the DPS. Dow Piniak et al. (2012) measured hearing sensitivity of
leatherback hatchlings. They found that hatchlings are able to detect
sounds underwater and in air, responding to stimuli between 50 and 1200
Hz in water and 50 and 1600 Hz in air, with maximum sensitivity between
100 and 400 Hz in water and 50 and 400 Hz in air. This sensitivity
range overlaps with the frequencies and levels produced by many
anthropogenic sources used in the North Atlantic, including seismic
airgun arrays, drilling, low frequency sonar, shipping, pile driving,
and operating wind turbines. These noise sources may affect leatherback
turtles' marine habitat and subsequently impact distribution and
behavior. Offshore artificial lighting occurs in some marine waters of
this DPS (Dow et al. 2007) but is less of a threat than beachfront
lighting throughout the range of the DPS.
Summary
We conclude that nesting females, hatchlings, and eggs are exposed
to the loss and modification of nesting habitat, especially as a result
of coastal development and armoring, erosion, and artificial lighting.
These threats impact the DPS by reducing nesting and hatching success,
thus, lowering the productivity of the DPS. Based on the information
presented above, we conclude that habitat reduction and modification
pose a threat to the NW Atlantic DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization is a threat to the NW Atlantic DPS, mostly due to
poaching of turtles and eggs in certain nations. Legal harvest of
turtles and eggs also occurs in some nations.
While the vast majority of nations within the range of the NW
Atlantic DPS protect leatherback turtles from harvest, it is legal in
some Caribbean and Central American nations (Brautigam and Eckert 2006;
Dow et al. 2007; Richardson et al. 2013; Horrocks et al. 2016). For
example, the harvest of leatherback turtles over 20 pounds is allowed
in Montserrat and Dominica from October 1 to May 31; Saint Lucia allows
leatherback turtles over 65 pounds to be taken from October 2 to
February 27; and St. Kitts and Nevis allows take of leatherback turtles
over 350 pounds from October 2 to February 27 (Montserrat Turtles Act
2002; Br[auml]utigam and Eckert 2006). In some nations, commercial use
is prohibited, but traditional use is allowed, which can still diminish
protection. In Colombia, subsistence fishing of sea turtles is
permitted, and indigenous use is allowed in Honduras. Traditional or
cultural use is permitted in Belize with prior approval (Br[auml]utigam
and Eckert 2006). However, regular leatherback nesting does not occur
in Belize, and its occurrence in surrounding waters is infrequent,
reducing the impact of such mortality. Legal harvest throughout the
range of this DPS is not monitored, and the precise magnitude of this
threat is not clear. However, we conclude that legal harvest of turtles
is significant because, when it occurs, nesting turtles are targeted,
removing the most important individuals from the population. More
often, leatherback eggs, rather than turtle meat, are harvested (TEWG
2007; Pati[ntilde]o-Mart[iacute]nez et al. 2008), reducing productivity
in the DPS.
Poaching of turtles and eggs occurs throughout the NW Atlantic DPS,
and Dow et al. (2007) ranked it as a threat for all turtle species on
the beaches in the Wider Caribbean Region. In Panama, interviews with
locals revealed that the development of a new way for cooking
leatherback turtle meat has resulted in a recent increase of its
consumption in Changuinola, Bocas del Toro Province (CITES Secretariat
2019). Adult turtles are killed in Panama and on remote beaches in
Trinidad and Tobago (Tro[euml]ng et al. 2002; Ordo[ntilde]ez et al.
2007; Trinidad and Tobago Forestry Division et al. 2010). Most
poaching, however, targets eggs, and the level often is determined by
how much monitoring and activity to deter poachers occur on the nesting
beaches. Some of the highest levels of egg poaching occur throughout
Costa Rica (Tro[euml]ng et al. 2004). Tro[euml]ng et al. (2007) found
that, at a minimum, between 13 to 21.5 percent of nests between 2000
and 2005 were illegally
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collected at Tortuguero. Poaching of leatherback nests was higher
outside Tortuguero National Park (minimum 33 percent) than within the
National Park (minimum 9 percent) in 2005 (de Haro and Tro[euml]ng
2006). At Pacuare Playa, Costa Rica, 55 percent of nests were poached
in 2012 (Fonseca and Chac[oacute]n 2012) and 42 percent were poached in
2017, which was the lowest level since Latin American Sea Turtles
(LAST) started to monitor in 2012 (LAST 2017). Poaching at Gandoca
Beach has decreased over time (previously 100 percent of nests were
poached), but rates still averaged 15.5 percent annually from 1990 to
2004 (Chac[oacute]n-Chaverri and Eckert 2007). In the Dominican
Republic, poaching is also high. Revuelta et al. (2012) determined the
poaching of clutches in Jaragua National Park and Saona Island ranged
from 0 to 100 percent from 2006 to 2010, with averages of 19 percent on
western Jaragua National Park beaches, 71 percent on eastern Jaragua
National Park beaches, and 74 percent on Saona. Poaching also occurs at
relatively high levels in Colombia (e.g., 22 to 31 percent of clutches
at Playona in 2006 and 2007; Pati[ntilde]o-Mart[iacute]nez et al. 2008)
and, to some extent, in most other Caribbean nations (e.g., Guyana and
Grenada). Poaching is likely more prevalent, and occurs at higher
levels, on unmonitored or unprotected beaches (Dow et al. 2007; TEWG
2007; Tro[euml]ng et al. 2007; Trinidad and Tobago Forestry Division et
al. 2010; K. Charles, Oceans Spirits Inc., pers. comm., 2018).
Poaching has been significantly reduced at some nesting beaches. In
Suriname, high levels of egg poaching (at least 26 percent of nests)
occurred in the late 1990s, but due to better monitoring and
enforcement, that level has been significantly reduced (Hilterman and
Goverse 2007; M. Hiwat, WWF, pers. comm., 2018). Poaching was also a
major problem in Trinidad, but levels have been reduced with more
people monitoring the beach (Trinidad and Tobago Forestry Division et
al. 2010). The Marine Turtle Conservation Act of 2004 (MTCA) funds
activities in Panama in an attempt to reduce poaching. At Chiriqui
Beach, Panama, intense monitoring efforts have attempted to reduce
poaching. However, of the monitored nests, 29 leatherback nests (0.7
percent) were still poached in 2017 (Sea Turtle Conservancy 2017).
Further, poaching in Panama outside the monitored areas still occurs,
with the clandestine sale of eggs widespread (Brautigam and Eckert
2006). In St. Croix, almost 100 percent of nests were lost to poaching
prior to 1981 (Garner et al. 2017). However, the establishment of the
USFWS Sandy Point National Wildlife Refuge has reduced egg poaching to
0 to 1.8 percent annually as a result of nightly patrols (Garner et al.
2017).
Poaching of eggs is widespread throughout the Caribbean, especially
on beaches of Costa Rica, Dominican Republic, and Colombia. The total
number of individuals affected by poaching cannot be quantified at this
time. However, we conclude that many eggs and some adults are affected
by illegal poaching at nesting beaches. Adults and eggs are also
exposed to legal harvest in some nations. The legal and illegal harvest
of nesting females reduces both abundance (through loss of nesting
females) and productivity (through loss of reproductive potential),
resulting in a high impact to the DPS. Legal and illegal egg harvest
reduces productivity only. Thus, we conclude that overutilization poses
a threat to the DPS.
Disease or Predation
For the NW Atlantic DPS, information on diseases is limited, but
predation is a well-documented threat.
Much of the available information on disease in leatherback turtles
was obtained by necropsy of stranded large juvenile and adult turtles;
the health implications of various conditions reported in this species
are incompletely understood. Solitary large intestinal diverticulitis
of unknown etiology was found in 31 subadult and adult leatherback
turtles stranded in U.S. waters (Stacy et al. 2015). All lesions were
chronic and unrelated to the cause of death in all cases, although risk
of perforation and other complications are possible. Adrenal gland
protozoal parasites were found in 17 leatherback turtles in North
American waters examined from 2001 to 2014; it is not currently known
whether parasitism affects adrenal function (Ferguson et al. 2016). In
addition, leatherback turtles are hosts for several trematode parasites
(flatworms), known species of which also occur in hard-shelled sea
turtles (Manfredi et al. 1996, Greiner et al. 2013). In general,
trematodes are frequently encountered without any apparent clinical
effect on the turtle host but can affect some heavily parasitized
individuals. With regard to other types of potential disease-causing
organisms, there are a small number of reports of bacterial infections
in stranded individuals (Poppi et al. 2012; Donnelly et al. 2016). A
variety of other bacteria have been documented in nesting females on
beaches in Costa Rica (Santoro et al. 2008) and St. Kitts (Dutton et
al. 2013); the majority of identified bacterial species may be
considered as potential or opportunistic pathogens for sea turtles. A
putative case of fibropapilloma, a virus-associated tumor-causing
disease in sea turtles, has been reported in a leatherback; this
disease is considered very rare in the species (Huerta et al. 2002).
An in-water health assessment was performed on 12 turtles directly
caught at-sea and seven turtles bycaught in fishing gear in the NW
Atlantic Ocean (Innis et al. 2010). Most were determined to be in good
health, but several exhibited evidence of past injuries. The blood
chemistry of entangled turtles indicated stress, seawater intake, and
reduced food consumption associated with entanglement. In addition,
Perrault et al. (2012) examined baseline blood chemistry metrics (i.e.,
plasma protein electrophoresis, hematology, and plasma biochemistry) as
indicators of health for nesting females in Florida. They found that
multiple measures of maternal health significantly correlated with
leatherback hatching and emergence success (the percentage of
hatchlings that emerge from the nest).
From these data, we estimate that the exposure of eggs, juveniles,
and adults to disease is low. The impact of disease cannot be
quantified at this time as we have no documentation of any deaths or
reductions in productivity directly related to disease. However,
disease may compound the effects of or have synergistic effects with
other threats to the species and related physiologic derangements. We
conclude that disease, alone or in combination with other threats, is
likely a threat to the DPS.
Throughout the range of the DPS, predation is a threat to
leatherback eggs, hatchlings, and adults. Eckert et al. (2012) provides
an exhaustive list of the documented predators for each life stage and
area. For eggs in the NW Atlantic DPS, predators include ants (Dorylus
spininodis), fly larvae (Diptera spp.), locust larvae (Acrididae spp.),
mole crickets (Scapteriscus didactylus), ghost crabs (Ocypode
quadratus), vultures (Cathartidae), dogs (Canis familiaris), cattle
(Bos taurus; due to trampling), armadillo (Dasypodidae), opossum
(Didelphis marsupialis), coati (Nasua spp.), and raccoons (Procyon
lotor); see Eckert et al. 2012).
In particular, dog predation of eggs occurs in many areas (e.g.,
Colombia, French Guiana, Guyana, Panama, Puerto Rico, and Trinidad and
Tobago). In Trinidad, where the largest nesting aggregation occurs,
feral dogs are
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considered to be the primary threat to eggs, even above poaching and
coastal erosion (Trinidad and Tobago Forestry Division et al. 2010). On
Chiriqui Beach, Panama, 54 percent of the monitored leatherback nests
were depredated by dogs in 2003 and approximately eight percent in 2004
(Ordo[ntilde]ez et al. 2007). Such predation may been reduced as a
result of protection efforts funded by the MTCA. In Playa California,
Maunabo, Puerto Rico, more than 30 percent of the leatherback nests
were depredated by stray dogs in 2012 (Crespo and Diez 2016). A public
outreach project in Puerto Rico was established in 2013 to reduce this
impact. Puerto Rico is a U.S. territory; if ESA protections were
removed, it is likely that predation rates would be higher.
Egg predation by other species is also a notable concern in some
areas. On Gandoca Beach, Costa Rica, dipteran larvae infestation
exceeded 75 percent of nests in 2005 and 2006 (Gautreau et al. 2008).
In French Guiana, on average, mole crickets preyed on 18 percent of all
eggs (Maros et al. 2003). These threats are likely to continue, as no
predator screening typically occurs in Wider Caribbean nations due to
the potential for increased poaching as well as logistical difficulties
in these areas of high density nesting. Nest loss to predators was
found to be the seventh ranked threat to turtles (all species, not
specific to leatherback turtles) on nesting beaches in the Wider
Caribbean Region, and have been noted to frequently occur in Honduras,
Mexico, Panama, Puerto Rico, and Venezuela (Dow et al. 2007).
Hatchlings are preyed upon by a wide variety of species, including
mole crickets, ghost crabs, horse-eye jack fish (Caranx latus), gray
snapper (Lutjanus griseus), tarpon (Megalops atlanticus), vultures,
hawks (Accipitridae), gulls (Larus spp.), night heron (Nyctanassa
violacea), frigate birds (Fregatidae), dogs, mongoose (Atilax
paludinosus), coati, and raccoons (Eckert et al. 2012). Again, dogs are
a serious threat to leatherback hatchlings in some areas, and
especially in Puerto Rico (Crespo and Diez 2016).
There are few documented predators to subadults and adult
leatherback turtles, presumably because of their large size and pelagic
behavior. Predation by sharks (Elasmobranchii) and killer whales
(Orcinus orca) has been reported in Barbados and St. Vincent,
respectively (Caldwell and Caldwell 1969; Horrocks 1989). Sharks have
also been reported to prey on nesting females off St. Croix, USVI
(DeLand 2017; Scarfo et al. 2019). Over the past 6 years, researchers
at Sandy Point have observed an apparent increase in injuries to
leatherback turtles (K. Stewart, NMFS, pers. comm., 2019). These
injuries, many of them consistent with shark predation, affect up to 70
percent of all nesting females at the beach (Scarfo et al. 2019). While
some turtles probably survive these encounters, it is unknown how many
encounters result in mortality or reduced nesting effort. Jaguars
(Panthera onca) prey on nesting females in some areas, including
Suriname, French Guiana, Guyana, and Costa Rica (see Eckert et al.
2012). While three nesting females were killed by jaguars at
Tortuguero, Costa Rica, from 1998 to 2005, this mortality is only
considered to be a minor threat and is therefore unlikely to cause a
population decline on its own (Tro[euml]ng et al. 2007). Archibald and
James (2018) examined 228 leatherback turtles for injuries off Atlantic
Canada and on Matura, Trinidad, and found 15.7 percent of turtles
exhibited injuries of suspected predatory origin.
Predation on early life stages is natural; however, at high rates,
it reduces the viability of the DPS (see the Status Review). Predation
primarily reduces productivity via reduced egg and hatching success and
the loss of hatchlings. Predation on nesting females reduces abundance
and productivity. We conclude that predation is a threat to the NW
Atlantic DPS.
Inadequacy of Existing Regulatory Mechanisms
Many regulatory mechanisms (including state, Federal and
international) have been promulgated to protect leatherback turtles,
eggs, and nesting habitat throughout the range of the NW Atlantic DPS.
We reviewed the objectives of each regulation and to what extent they
adequately address the targeted threat (i.e., the threat that the
regulation was intended to address). The effectiveness of many
international regulations was evaluated by Hykle (2002), who found that
international instruments often do not realize their full potential,
either because they do not include all key countries, do not
specifically address sea turtle conservation, are handicapped by the
lack of a sovereign authority that promotes enforcement, or are not
legally binding.
National regulatory mechanisms are described in full in the Status
Review Report. Although these regulatory mechanisms provide some
protection to the species, most inadequately reduce the threat they
were designed to address, generally as a result of poor implementation
or incomplete enforcement. Specifically, existing regulatory mechanisms
continue to be inadequate to control impacts to nesting beach habitat
and overutilization (harvest of turtles and eggs) for this DPS. In
addition, regulatory mechanisms are inadequate to reduce several other
threats including bycatch in fishing gear, vessel strikes, and marine
debris. Despite existing regulatory mechanisms, bycatch from fisheries
(discussed in detail along with existing regulatory mechanisms in the
Fisheries Bycatch section), incomplete nesting habitat protection, and
poaching remain major threats to the DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat to the NW Atlantic DPS.
Bycatch occurs throughout the range of the DPS, affecting juveniles,
subadults, and adults.
Finkbeiner et al. (2011) analyzed sea turtle bycatch across all
commercial U.S. fisheries from 1990 to 2007. They examined sea turtle
bycatch reduction based on the year a particular fishery implemented
bycatch reduction measures. Prior to implementing bycatch reduction
measures, approximately 3,800 leatherback interactions, of which 2,300
were lethal, occurred in U.S. Atlantic Ocean and GOM commercial
fisheries annually. After bycatch reduction measures were implemented,
1,400 leatherback turtles, 40 of those dead, were estimated to be taken
annually in the Atlantic Ocean. The Atlantic/GOM pelagic longline
fishery was responsible for the most annual interactions (n = 900) and
mortality events (n = 17) in the Atlantic Ocean, followed by the
southeast Atlantic/GOM shrimp trawl fishery (Finkbeiner et al. 2011).
These estimates represent minimum numbers of actual bycatch and
mortality. Because the observer coverage for these fisheries is low (so
some bycatch may not be observed and observed effort may not be a true
representation of actual fleet effort), not all fisheries are observed
and thus some are not included in these estimates. Interactions are
difficult to observe if gear modifications are in place, and so the
methods used are conservative (Finkbeiner et al. 2011).
In the Wider Caribbean Region, reports of leatherback bycatch in
fisheries are common. In a survey of Caribbean nations, Dow et al.
(2007) ranked fisheries bycatch among the highest in-water threat to
sea turtles. Many fisheries in less industrialized nations are coastal
and small-scale, but these fisheries are reported to have significant
ecological impacts due to their high bycatch discards and impacts
[[Page 48349]]
to the marine environment (Shester and Micheli 2011). Of particular
concern are leatherback bycatch in artisanal nearshore and offshore
gillnet, longline and trawl fisheries (Barrios-Garrido and Montiel-
Villalobos 2016). Information on fisheries bycatch is collected mostly
from stranding records but also from fisher surveys (Moncada et al.
2003; Delamare 2005; Madarie 2006, 2010, 2012) and observations of
nesting females. Hilterman and Goverse (2007) recorded fisheries
related injuries on nesting females in Suriname. In 2002, 16.9 percent
of the nesting females had fisheries- related injuries; in 2003, at
least 18.3 percent had such injuries; and in 2005, 9 percent (Hilterman
and Goverse 2007). From 2000 to 2003, an average of 28 leatherback
turtles stranded on the Suriname survey beaches. Although no cause of
death was immediately apparent, Hilterman and Goverse (2007) indicated
that the mortalities were fisheries-related, based upon the fisheries
that occur offshore with high bycatch and documented fisheries-related
injuries on nesting leatherback turtles at the same time. On the
western oceanic nesting beaches of French Guiana, injuries consistent
with fisheries interactions (e.g., scars, wounds) were recorded on 8.4
percent (n = 1,259) of nesting females in 2003 (Morisson et al. 2003).
In Venezuela, 55 percent of strandings from 2001 to 2007 (n = 57)
exhibited evidence of fisheries interactions (Barrios-Garrido and
Montiel-Villalobos 2016). Most recently, an injury assessment of 228
leatherback turtles from two foraging areas off the Atlantic coast of
Canada and Trinidad nesting beaches found 19 percent of turtles
exhibited injuries indicative of entanglement in lines or nets, and 17
percent showed evidence of hooks; 62 percent of turtles assessed
exhibited a minimum of one external injury (Archibald and James 2018).
Fisheries bycatch also occur in the Mediterranean and eastern North
Atlantic Ocean. Casale et al. (2003) analyzed 411 records of
leatherback turtles in the Mediterranean, of which 152 were collected
from Italy. Most of these records were from fishery captures (n = 170)
or found in unknown circumstances (n = 127). Of those reported by
fishermen, set or drift nets had the highest number of interactions
(29.4 percent), followed by unknown fishing equipment (22.9 percent),
longlines (20.6 percent), unspecified nets (12.9 percent), other
fishing equipment (9.4 percent), and trawls (4.7 percent). The main
fisheries affecting turtles in the Mediterranean (all turtle species,
not just leatherback turtles) are Spanish and Italian surface
longlines, North Adriatic Italian trawls, Tunisian trawls, Turkish
trawls, Moroccan driftnets, and Italian driftnets (Cami[ntilde]as
2004). The same types of fishing gear from other nations also affect
turtles, but the bycatch numbers are lower (Cami[ntilde]as 2004).
Stranding records from Portugal from 1978 to 2013 found that 49 of 275
leatherback turtles exhibited evidence of fishery interactions (the
cause of stranding could not be determined in most cases due to
decomposition state; Nicolau et al. 2016). Multifilament nets accounted
for approximately 41 percent of the strandings, followed by
monofilament nets, traps/pots, and longlines. Coastal artisanal
fisheries were recognized as a particular threat in Portugal.
Based upon these summary reports and stranding assessments, it is
clear that fisheries have a large impact on the NW Atlantic DPS. In the
following paragraphs, we review information on specific gear
interactions, including the following fisheries: Gillnet, longline,
trawl, pot/trap, and other.
Gillnet Fisheries
Gillnet fisheries are common throughout the range of this DPS. Due
to the nature of the gear and fishing practices (e.g., relatively long
soak times), bycatch in gillnets is among the highest source of direct
sea turtle mortality (Upite et al. 2013; Wallace et al. 2013; Upite et
al. 2018). Upite et al. (2018) evaluated observed fishery interactions
and post-interaction mortality and determined a 79 percent sea turtle
mortality rate for Northeast and Mid-Atlantic gillnet gear from 2011 to
2015. Wallace et al. (2013) calculated leatherback bycatch in gillnets
throughout the NW Atlantic Ocean of 0.015 turtles/set, with a 21
percent median mortality rate (not considering post-interaction
mortality). This gear was classified as having a relatively high
bycatch impact on the NW Atlantic leatherback population. Small scale
fisheries are of particular concern, given the magnitude of bycatch,
nearshore distribution, and limited monitoring (Lewison et al. 2015).
When nets are used in waters off nesting beaches, where leatherback
turtles mate, nesting females and mature males are often captured and
killed.
The largest documented bycatch of leatherback turtles in gillnet
gear occurs off the coast of Trinidad. Lee Lum (2006) estimated that
more than 3,000 leatherback turtles were captured by coastal surface
gillnets off Trinidad annually, with an approximate 30 percent
mortality rate. These captures involved adult turtles, occurring off
the north and east coasts of Trinidad during January to August, i.e.,
the breeding and nesting season, when nesting females and adult males
occur in the waters off nesting beaches (Lee Lum 2006). Gilman et al.
(2010) extrapolated leatherback bycatch estimates (Lee Lum 2006;
Gearhart and Eckert 2007) to the entire Trinidad Spanish mackerel and
king mackerel surface gillnet fishery, and estimated that almost 7,000
turtles were captured in 2000. Additionally, Eckert et al. (2013)
worked with drift gillnet fishermen to identify leatherback bycatch hot
spots off the north and east coasts of Trinidad (where the nesting
beaches are), with capture probability increasing from March to July
and a secondary peak in October.
Whereas most of the documented leatherback bycatch off Trinidad
occurs in surface drift gillnet fisheries, bottom set gillnet fishing
also captures leatherback turtles (Gass 2006; S. Eckert, WIDECAST,
pers. comm., 2018). The magnitude of effort and turtle bycatch in this
fishery are lower than for surface nets, but mortality rates are higher
(approximately 70 percent; Gass 2006). As such, the bottom set gillnet
fishery is thought to have a comparable level of mortality to the drift
gillnet fishery (approximately 500 to 1,000 leatherback turtles
annually; Gass 2006; S. Eckert, WIDECAST, pers. comm., 2018). The Sea
Turtle Recovery Action Plan for the Republic of Trinidad and Tobago
noted that drowning in gillnets is that nation's most significant cause
of sea turtle mortality (Trinidad and Tobago Forestry Division et al.
2010). Bond and James (2017) tracked a female from Canadian waters to a
nesting beach off Trinidad, but the turtle was confirmed dead,
entangled in coastal fishing gear, just prior to the date of her first
predicted nesting event. Venezuelan fishers have also been seen hauling
leatherback turtles from Trinidad waters into their boats (Brautigam
and Eckert 2006). Together, drift and bottom-set gillnets off the
Trinidad beaches, which host the largest nesting aggregation in the
DPS, are estimated to kill well over 1,000 leatherback turtles
annually, and they thus pose a large threat to the DPS.
High levels of gillnet bycatch occur in other Caribbean and South
American nations, also off major nesting beaches. In French Guiana,
bycatch was confirmed to be high in the Maroni estuary (Chevalier 2001;
Girondot 2015). In 2003, 26 leatherback turtles were caught in coastal
gillnets and released off the Cayenne and Montjoly nesting sites
(Gratiot et al. 2003 in TEWG 2007). Delamare (2005) conducted fishermen
interviews and estimated an average of 1,149 leatherback captures in
2004 and
[[Page 48350]]
2005 by bottom-set or drifting gillnets in French Guiana. No estimate
of mortality was provided, but it is likely similar to Trinidad
fisheries, i.e., 70 and 30 percent, respectively. In Suriname, a World
Wildlife Fund survey of fishermen estimated leatherback bycatch in
drifting gillnets at 584 in 2006, 174 in 2010, and 424 in 2012 (Madarie
2006; Madarie 2010; Madarie 2012). Most of the turtles were captured
alive. In Colombia, 10 to 40 leatherback turtles are killed annually by
gillnets (Pati[ntilde]o-Mart[iacute]nez et al. 2008). Longline and
driftnet gillnet fisheries in Moroccan waters off the northwestern
Africa coast capture approximately 100 leatherback turtles annually
(Benhardouze et al. 2012).
Although not at as high a rate as in the Caribbean (based upon
observed interactions), gillnet bycatch occurs in U.S. and Canadian
waters. Although South Carolina, Georgia, Florida, Louisiana, and Texas
have prohibited gillnets in their State waters, active gillnet
fisheries remain in other states and U.S. Federal waters. No cumulative
estimates of leatherback bycatch in gillnet fisheries in U.S. waters
are available due to the limited observed interactions. However, from
2003 to 2017, fishery observers recorded lethal and non-lethal bycatch
in fixed sink, drift sink, and drift floating gillnets throughout the
U.S. Atlantic Exclusive Economic Zone (EEZ) and GOM (NMFS unpublished
data). From 2012 to 2016, 27 leatherback turtles (coefficient of
variation = 0.71, 95 percent CI over all years: 0-68) were bycaught
with 21 mortalities in sink gillnet gear in the Georges Bank and Mid-
Atlantic regions (Murray 2018). From 1989 to 1998, U.S. drift pelagic
gillnets captured 54 leatherback turtles, but that gear is no longer
used. Hamelin et al. (2017) reviewed leatherback entanglement records
reported by Canada in Atlantic Canadian waters between 1998 and 2014.
Gillnets, mainly targeting groundfish, were involved in 24 of 205
entanglements (11.7 percent), particularly in Newfoundland and Labrador
(n = 15). Often, gillnet entanglements involve the vertical lines
associated with gear (M. James, DFO, pers. comm., 2019).
Gillnet bycatch occurs in the eastern North Atlantic Ocean and in
the Mediterranean Sea. As in other areas, sea turtles have the
potential to interact with set gillnets and drift gillnets. The United
Nations (UN) established a worldwide moratorium on drift gillnet
fishing effective in 1992; the General Fisheries Commission for the
Mediterranean prohibited driftnet fishing in 1997; a total ban on
driftnet fishing by the European Union fleet in the Mediterranean went
into effect in 2002; and the International Commission for the
Conservation of Atlantic Tunas (ICCAT) banned driftnets in 2003.
Nevertheless, unregulated driftnetting continued to occur in some areas
(e.g., the Mediterranean Sea and off Europe; Pierpoint 2000;
Cami[ntilde]as 2004). In the Atlantic Ocean, leatherback bycatch has
been reported from NE Atlantic tuna driftnet fisheries by English,
French and Irish vessels (Pierpoint 2000). Of 20 leatherback turtles
found in nets in British and Irish waters (1980 to 2000), eight were
caught in the NE Atlantic tuna driftnet fishery (with 25 percent
mortality) and one was caught in a hake gillnet (Pierpoint 2000).
Historically, driftnet fishing in the Mediterranean Sea caught
large numbers of sea turtles. And today an estimated 600 illegal
driftnet vessels operate in the Mediterranean, including fleets based
in Algeria, France, Italy, Morocco, and Turkey (Environmental Justice
Foundation 2007). Out of 411 records of leatherback turtles (stranded,
captured, sighted, or found in unknown circumstances) in the
Mediterranean Sea, 170 turtles were captured by fishermen, of which
29.4 percent were caught by set or drift nets (Casale et al. 2003).
Driftnets and gillnets in Greece, Israel, Italy, Tunisia and Turkey
have reported documented leatherback interactions, and occasional
leatherback bycatch occurs in Croatian artisanal gillnet fisheries
(Cami[ntilde]as 2004; Ergene and Ukar 2017). In particular, Karaa et
al. (2013) reviewed 36 leatherback bycatch records from Tunisia
fisheries in the Gulf of Gabes, and found that gillnets are the
dominant threat to leatherback turtles in the region. A similar result
(e.g., gillnets being a high threat to leatherback turtles in the area)
was found in the Adriatic Sea (Lazar et al. 2012). The first
leatherback recorded on the Aegean coast of Turkey was caught in a
gillnet (Taskavak et al. 1998). Further, a review by Casale (2008)
found that leatherback turtles are taken in the drift gillnet fishery
in Spain at a rate of 0.065 turtles/day-boat.
Throughout the range of the NW Atlantic DPS, effective gillnet
bycatch reduction measures have not been required, but measures to
reduce leatherback bycatch have been discussed in some areas (e.g.,
Trinidad; Eckert 2013). If nations have a closed season for fishing, at
least in the nesting season (e.g., Suriname; Madarie 2006), nesting
females are afforded some level of protection from gillnet bycatch.
Some nations have prohibited gillnet gear; St. Barthelemy does not
allow trammel nets in its territorial waters and St. Lucia prohibits
fishing within 100 meters of shore to protect nesting turtles. There
are gillnet and trammel net restrictions in Curacao (Ministry of
Health, Environment, and Nature 2014, UN Environment Programme 2017).
In the United States, gillnets with stretched mesh seven inches and
larger are prohibited at certain times off North Carolina and Virginia
to protect sea turtles (50 CFR 223.206(d)(8); 71 FR 24776, April 26,
2006). While no gear modifications are currently required for U.S.
gillnet fisheries, Federal U.S. fisheries are subject to section 7 of
the ESA, 16 U.S.C. 1536(a)(2), and through formal consultations on
specific fisheries, measures may be required to minimize the impact of
incidental take in gillnets (NMFS 2013). Regardless of some of these
protective measures, gillnet bycatch (especially off nesting beaches)
results in the loss of thousands of mature individuals annually.
Longline Fisheries
Leatherback turtles are known to interact with longline fishing
gear, most commonly pelagic longlines (Lewison et al. 2004; Zollett
2009; Wallace et al. 2010; Wallace et al. 2013). There is significant
concern over the effects of pelagic longline fishing, which extends
globally throughout temperate and tropical waters, including several
high pressure fishing areas in the North Atlantic Ocean (Fossette et
al. 2014; Gray and Diaz 2017). In international waters, numerous flag
states have high seas longline fisheries that frequently catch
leatherback turtles (Lewison et al. 2004). Individuals are found
entangled and hooked in this gear, mostly by the flippers (Witzell and
Cramer 1995; Coelho et al. 2015; Huang 2015). Leatherback bycatch in
longlines throughout the NW Atlantic Ocean was calculated at 0.062
turtles per set, classifying the gear as a relatively low bycatch
impact relative to other sea turtle populations (Wallace et al. 2013;
Lewison et al. 2015). However, because longline fisheries are
widespread across leatherbacks' distribution and use millions of hooks
each year, they pose a large threat to the NW Atlantic DPS and are
estimated to kill thousands of individuals (mature and immature)
annually.
Pelagic longline fishing is widespread throughout the range of the
DPS and involves a number of nations, so an accurate estimate of total
bycatch is difficult to obtain. In the Atlantic Ocean from 2002 to
2013, the largest longline fishing fleets belonged to Taiwan, Japan,
Spain, Belize, and China, with the Taiwanese fleet comprising the
largest distant-water longline effort throughout
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the region (Angel 2014; Huang 2015). In an assessment of the impact of
ICCAT fisheries on sea turtles, Gray and Diaz (2017) estimated
leatherback interactions with pelagic longlines in the ICCAT area from
2012 to 2014 (15 to 16 fleets). Using a combination of published and
assigned sea turtle bycatch rates as a function of estimated fishing
effort submitted to ICCAT by its members, Gray and Diaz (2017) found a
high degree of overlap in the central North Atlantic Ocean and
equatorial waters (some of which are outside this DPS). Within the NW
Atlantic region, an estimated 7,138 leatherback interactions occurred
in 2012, 6,036 in 2013 and 4,991 in 2014 (Gray and Diaz 2017). Applying
a reasonable estimated mortality rate of 21.4 percent, as seen in other
high seas pelagic longline gear (Huang 2015), results in an average
annual estimated mortality of 1,296 leatherback turtles from 2012 to
2014. However, this is likely an underestimate of total mortality, as
the high seas mortality rate in Huang (2015) was based upon the
disposition of the turtle when boarded and therefore did not account
for post-interaction mortality; 240 of 459 leatherback turtles caught
from 2002 to 2013 were alive and 121 were of unknown status (Huang
2015). Angel et al. (2014) conducted a risk assessment of turtles from
the impacts of tuna fishing in the ICCAT region and found the NW
Atlantic RMU (which is comparable to the NW Atlantic DPS; Wallace et
al. 2010) has high-moderate vulnerability to longline gear, with as
many as 270 million longline hooks annually from 2000 to 2009. In
particular, Fossette et al. (2014) analyzed leatherback satellite
tracks (converted to densities) overlaid with longline fishing effort
from 1995 to 2009 in the Atlantic Ocean. In the North Atlantic Ocean, a
total of four seasonal high-susceptibility areas were identified: one
in the central northern Atlantic in international waters, one along the
east coast of the United States, and one each in the Canary and Cape
Verdean basins (Fossette et al. 2014). These areas partly occurred in
the EEZs of eight nations (Cape Verde, Gambia, Guinea Bissau,
Mauritania, Senegal, Spain/Canaries, United States, and Western
Sahara). Given the species' flexible diving behavior, it is reasonable
to expect that turtles are likely to encounter pelagic longlines
throughout the Atlantic Ocean, regardless of whether they are engaged
in foraging or migratory behavior (Fossette et al. 2014).
Bycatch in U.S. Atlantic and GOM pelagic longlines has been
extensively studied in the last decade. Current estimates of
leatherback interactions with the U.S. Atlantic pelagic longline
fishery are lower than previous years. In the late 1990s and early
2000s, estimates of Atlantic U.S. pelagic longline bycatch were around
1000 leatherback turtles annually (NMFS 2001; Yeung 2001; NMFS 2018),
with bycatch rates of about 0.15 to 0.5 turtles per 1000 hooks (Watson
et al. 2005). In 2005, after the United States required pelagic
longline gear modifications (50 CFR 635.21), the fleet was estimated to
have interacted with 351 leatherback turtles outside experimental
fishing operations (Walsh and Garrison 2006). NMFS (2018) estimated 239
leatherback interactions in the U.S. Atlantic pelagic longline fishery
in 2011, 596 in 2012, 363 in 2013, 268 in 2014, 299 in 2015, and 339 in
2016. The majority of interactions occurred in the GOM, Mid-Atlantic
Bight, Northeast Coastal, and Northeast Distant areas (NMFS 2018). The
post-interaction mortality estimate for the most recently available 3-
year period (2013 to 2015) for leatherback turtles is 30.13 percent (L.
Desfosse, NMFS, pers. comm., 2018). Based on the average leatherback
interaction estimate for the entire U.S. pelagic longline fleet from
2011 to 2016 (351), the estimated average annual mortality for the U.S.
pelagic longline fishery is 106 leatherback turtles.
Leatherback interactions also occur in Canadian pelagic longline
fisheries. From summer to fall, primarily on the Scotian Shelf,
encounters with leatherback turtles have been documented in the large
pelagic longline fishery since 2001 (DFO 2012). With observer coverage
ranging from 5 to 30 percent since 2001, there were 102 reported
interactions with pelagic longlines from 2001 to 2005, and 36 from 2006
to 2010 (DFO 2012). Mortality rates are estimated to be in the range of
21 to 49 percent, resulting in an estimated mortality of 13 to 44
leatherback turtles annually. Based on an analysis of Canadian observer
data from 2002 to 2010, the bycatch rate in this fishery is estimated
to have declined from 120-190 leatherback turtles annually from 2002 to
2006 to 60-90 leatherback turtles annually from 2006 to 2010, largely
as a result of gear modifications (Hanke et al. 2012).
In the Mediterranean Sea, longlining is prevalent. Drifting
longlines targeting swordfish (Xiphias gladius), albacore (Thunnus
alalunga), and bluefin tuna (T. thynnus) are considered to be the most
dangerous fishing gear for turtles in the Mediterranean Sea (Lucchetti
and Sala 2010). Drifting longlines (mainly for albacore tuna) in Spain,
Italy, Greece, and Albania have documented leatherback interactions
(Cami[ntilde]as 2004). In the western Mediterranean, swordfish
longlines appeared to be responsible for most of the leatherback
bycatch and entanglements (Cami[ntilde]as 1998; Cami[ntilde]as 2004).
Casale et al. (2003) reviewed bycatch rates for longline fisheries
targeting swordfish and estimated the average Mediterranean longline
bycatch rates at 0.0025 leatherback turtles/1000 hooks, with a maximum
rate of 0.0510 leatherback turtles/1000 hooks in the Tyrrhenian Sea of
Italy (Casale et al. 2003; Casale and Margaritoulis 2010). Of 170
leatherback fishery captures in fisheries from the Mediterranean Sea,
approximately 35 involved longlines (Casale et al. 2003). While
leatherback turtles are encountered in Mediterranean longlines,
loggerheads are the most common species caught; only 0.1 percent of
turtles captured during an observer program in Spain, Italy and Greece
were leatherback turtles (3 out of 2,370 observed turtles; Laurent et
al. 2001). However, given the extensive longline effort in the
Mediterranean Sea (Casale 2008), leatherback bycatch in the
Mediterranean is still a concern. Lewison et al. (2004) estimated a
range of 250 to 10,000 leatherback turtles bycaught in the
Mediterranean in 2000, with 6 percent observer coverage.
Longline bycatch of leatherback turtles in the range of the NW
Atlantic DPS also occurs in waters off Cape Verde (Melo and Melo 2013;
Coelho et al. 2015), Morocco (Benhardouze et al. 2012), and Brazil
(Pacheco et al. 2011). Given the wide distribution of both pelagic
longline gear and leatherback turtles, bycatch of individuals in
longline gear can occur wherever and whenever the gear and sea turtle
distribution overlap.
Large circle hooks (non-offset) have been found to reduce
leatherback bycatch by as much as 55 percent compared to traditional J-
style hooks (Andraka et al. 2013; Coelho et al. 2015). While the
vessels of certain nations may employ large circle hooks, there are no
obligations for international longline fleets to adopt such bycatch
mitigation measures (Richardson et al. 2013). In 2005, an ICCAT
resolution encouraged circle hook research (ICCAT 2005), but no legally
binding measure to require circle hooks exists (Gilman 2011). Without
the widespread use of non-offset circle hooks, it is likely that the
high bycatch rates of leatherback turtles in pelagic longline gear will
continue throughout the North Atlantic high seas fisheries.
Since 2004, the United States has issued regulations that require
[[Page 48352]]
modifications to pelagic longline gear in the U.S. Atlantic and GOM to
reduce the bycatch and post-interaction mortality of sea turtles; these
regulations (50 CFR 635.21(c)(2)) specify hook type and size (18/0 or
16/0 circle hooks depending on the area), bait type, use of turtle
disentangling equipment and handling guidelines. Swimmer et al. (2017)
recently analyzed pelagic longline interactions before (1992 to 2001)
and after (mid-2004 to 2015) these regulations were promulgated.
Throughout the study period, 844 leatherback turtles were captured.
Overall, turtle bycatch was highest in the Northeast Distant
statistical reporting area (0.3 turtles/1000 hooks), followed by the
Northeast Coastal, GOM, and Caribbean areas. Bycatch rates were higher
for years prior to 2004; after the regulations, Atlantic leatherback
bycatch rates declined by 40 percent (0.13 to 0.078 turtles/1000
hooks). Within the Northeast Distant area alone, where additional
restrictions include a large circle hook (18/0) and limited use of
squid bait, rates declined by 64 percent (0.44 to 0.16 turtles/1000
hooks; Swimmer et al. 2017). Gilman and Huang (2017) found similar
results: Fish versus squid bait lowered catch rates of leatherback
turtles, and wider circle hooks reduced leatherback catch rates
relative to narrower J and tuna hooks. Capture probabilities are lowest
when using a combination of circle hook and fish bait.
Efforts have been made to reduce interactions in Canadian waters as
well. Circle hook use has been recommended in the swordfish-directed
Canadian longline fleet since 2003, whereas corrodible circle hooks
have been required in the pelagic longline fishery since 2012 (DFO
2013; C. MacDonald, DFO, pers. comm., 2019). There is no mandatory hook
size restriction for the Canadian longline fleet, but license holders
almost exclusively use 16/0 circle hooks (C. MacDonald, DFO, pers.
comm., 2019). De-hooking and line-cutting kits are required on
swordfish longline fishery vessels (C. MacDonald, DFO, pers. comm.
2019).
Some fishing fleets in the Atlantic Ocean (e.g., U.S., Canadian,
ICCAT vessels) use large circle hooks and modified bait, but these
measures are not required in all areas (Watson et al. 2005; Gilman et
al. 2007; Gilman 2011). Some nations in the Wider Caribbean Region have
implemented circle hook provisions; in Belize, the high seas fishing
fleet adopted the use of circle hooks on 10 percent of the fleet and
are required to report capture of sea turtles by longlines (Belize
Fisheries Department 2017). Because the measures are not widely
required, the number of vessels that do not employ bycatch reduction
measures is likely higher than the number of vessels that do, and so we
conclude on the basis of the best available information that
leatherback bycatch in pelagic longline fisheries is still a
significant threat (Lewison et al. 2015).
Leatherback interactions with bottom longlines also occur. Directed
shark fisheries using bottom longlines in the Atlantic Ocean and GOM
may capture or entangle leatherback turtles (NMFS 2012), and the GOM
reef fishery is also anticipated to take leatherback turtles (NMFS
2011). On February 7, 2007, NMFS published a rule that required
commercial shark bottom longline vessels to carry the same dehooking
equipment as the pelagic longline vessels; this rule was promulgated to
reduce post-interaction mortality (72 FR 5633).
The Canadian east coast groundfish longline fishery targets a wide
variety of groundfish species, including cod, haddock, pollock and
white hake. Observer coverage has ranged from 2 to 30 percent depending
on area, and there have been no reported interactions of leatherback
turtles in the observer database since 2001 (DFO 2012). However, there
have been three reports from Quebec logbooks and 10 reports of
interactions with groundfish longline gear to non-governmental groups
(DFO 2012). This indicates that the risk of interactions in this gear
may be higher than documented through the observer program.
Bottom longlines are also used in the Mediterranean Sea (Casale
2008). While there have not been any documented leatherback captures
from this gear type, loggerheads have been caught at high rates in
Tunisia, Libya, Greece, Turkey, Egypt, Morocco, and Italy (Casale
2008), and interactions with leatherback turtles are possible.
Commercial pelagic longline fisheries do not operate in some
Caribbean nations, such as in Panama where effort is limited to vessels
under six tons (Executive Decree 486, December 28, 2010). However,
other Caribbean nations allow commercial pelagic longline fishing, and
many find leatherback turtles with longline hooks (R[eacute]serve
Naturelle de l'Amana data in Berzins, Office National de la Chasse et
de la Faune Sauvage, pers. comm., 2018 and KWATA data in Berzins 2018).
While no longlines exist in the Caribbean Dutch nations of Bonaire, St.
Eustatius and Saba, there are efforts to introduce circle hooks into
the trolling fishery (Ministry of Economic Affairs 2014). We consider
longline bycatch to be a widespread threat to this DPS, likely
resulting in the loss of thousands of individuals annually.
Trawl Fisheries
Leatherback turtles may interact with bottom and midwater trawl
gear throughout the North Atlantic Ocean. The highest reported trawl
bycatch of leatherback turtles of the NW Atlantic DPS is likely from
the southeastern U.S. shrimp trawl fishery. Epperly et al. (2003)
anticipated an average of 80 leatherback mortalities a year in shrimp
trawl interactions, dropping to an estimate of 26 leatherback
mortalities in 2009 due to reduction in fishing effort (Memo from Dr.
B. Ponwith, SEFSC, to Dr. R. Crabtree, SERO, January 5, 2011). The 2014
NMFS Southeast U.S. Shrimp Fishery Biological Opinion estimated 167
annual leatherback captures (144 mortalities) in the Atlantic Ocean and
GOM shrimp otter trawl fishery, with an additional 34 captures in try
nets (single nets testing for shrimp concentrations; NMFS 2014). The
majority of these interactions were in the GOM. However, a more recent
study of the GOM and southeastern U.S. Atlantic coast shrimp otter
trawl fishery found fewer leatherback captures: From 2007 to 2017, only
3 leatherback turtles were reported in the observer data (with coverage
levels around 2 percent of nominal days at sea; Babcock et al. 2018).
In the mid-Atlantic and northeastern U.S. waters, observers
reported 9 leatherback captures in bottom otter trawl gear and 5
captures in midwater trawls from 1993 to 2017 (NMFS unpublished data
2018). In the Wider Caribbean Region, leatherback turtles are reported
captured in trawls in French Guiana (Ferraroli et al. 2004; TEWG 2007),
Guyana (Reichart et al. 2003), Suriname (Madarie 2010), Trinidad
(Forestry Division et al. 2010), and Venezuela (Alio et al. 2010).
Since 1980, there were eight reports of leatherback turtles
incidentally captured by trawl gear in British and Irish waters
(Pierpoint 2000). In the Mediterranean Sea, leatherback bycatch in
bottom trawls off Tunisia (Caminas 2004) and Egypt (Casale 2008) has
also been reported.
Trawl bycatch reduction measures (e.g., turtle excluder devices
(TEDs) are in place in some nations. The southeastern U.S. shrimp
fishery has required TEDs since the early 1990s. However, TEDs that
were initially required for use in the U.S. Atlantic Ocean and GOM
shrimp fisheries were less effective for leatherback turtles as
compared to smaller, hard-shelled turtle species, because the TED
openings were
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too small to allow leatherback turtles to escape. To address this
problem, NMFS issued a final rule on February 21, 2003, to amend the
TED regulations (68 FR 8456) to require modified TEDs in the
southeastern United States (Atlantic Area and GOM Area) that exclude
leatherback turtles, as well as large benthic immature and sexually
mature loggerhead and green sea turtles. TEDs are also required in
summer flounder trawls operating off Virginia (south of Cape Charles)
and North Carolina (64 FR 55860, October 15, 1999; 67 FR 19933, April
17, 2002).
TEDs are also used outside the United States. Shrimp harvested with
commercial fishing technology that may adversely affect sea turtles
generally cannot be imported into the United States per Public Law 101-
162, Section 609(b), enacted on November 21, 1989 (16 U.S.C. 1537
note). The import ban does not apply to nations that have adopted sea
turtle protection programs comparable to that of the United States
(i.e., require and enforce TED use) or whose fishing activity does not
present a threat to sea turtles (e.g., nations fishing in areas where
sea turtles do not occur). Although most certifications are done on a
national basis, the U.S. State Department guidelines allow some
individual shipments of TED-harvested shrimp from uncertified countries
with appropriate documentation. Approximately 40 nations are currently
certified to import shrimp into the United States, and five fisheries
have been determined as having their products eligible for importation
with proper documentation (83 FR 22739, May 16, 2018). Specifically, on
May 8, 2018, the U.S. State Department certified 13 nations on the
basis that their sea turtle protection programs (e.g., use of TEDs) are
comparable to that of the United States: Colombia, Costa Rica, Ecuador,
El Salvador, Gabon, Guatemala, Guyana, Honduras, Mexico, Nicaragua,
Nigeria, Panama, and Suriname. It also certified 26 shrimp-harvesting
nations and one economy as having fishing environments that do not pose
a danger to sea turtles. In addition, one fishery from a non-certified
nation within the range of the NW Atlantic DPS (the French Guiana
domestic trawl fishery) has been authorized to import shrimp products,
provided certain documentation accompanies the imports. Sixteen nations
have shrimping grounds only in cold waters where the risk of taking sea
turtles is negligible: Argentina, Belgium, Canada, Chile, Denmark,
Finland, Germany, Iceland, Ireland, the Netherlands, New Zealand,
Norway, Russia, Sweden, the United Kingdom, and Uruguay. Ten nations
(Bahamas, Belize, China, the Dominican Republic, Fiji, Jamaica, Oman,
Peru, Sri Lanka, and Venezuela) and Hong Kong only harvest shrimp using
small boats with crews of less than five that use manual rather than
mechanical means to retrieve nets or catch shrimp using other methods
that do not threaten sea turtles. Use of such small scale technology is
not believed to adversely affect sea turtles. For those nations within
the geographical range of the NW Atlantic DPS, the threat of shrimp
trawling is minimized with TED use.
TEDs are also required in trawl fleets in Trinidad, Belize, Brazil,
and Venezuela, but those gear modifications do not currently meet the
U.S. certification protocol. On June 20, 2019, the European Union
passed a regulation (PE-CONS 59/1/19 Rev 1) that requires technical
measures concerning: The taking and landing of marine biological
resources; the operation of fishing gear; and the interaction of
fishing activities with marine ecosystems. Specific to sea turtles, the
regulation requires shrimp trawl fisheries to use a TED in European
Union waters of the Indian and West Atlantic Oceans, consisting of
waters around Guadeloupe, French Guiana, Martinique, Mayotte,
R[eacute]union and Saint Martin.
TEDs are not required in Mediterranean trawls. Some nations, like
Belize, St. Barthelemy, Venezuela (industrial fishing only), and the
Caribbean Netherlands (Bonaire, St. Eustatius, Saba), have banned
trawling (Bolivarian Republic of Venezuela Official Gazette N[deg]
5.877, March 14, 2008; Ministry of Economic Affairs 2016; Belize
Fisheries Department 2017), and Costa Rica does not allow the issuance
of any new permits for shrimp trawling (Costa Rica Ministry of
Environment and Energy 2017). Curacao prohibits fishing in its
territorial waters and inland bays with dragnets (and certain fish
traps). These initiatives reduce the impact of trawling on leatherback
turtles.
Pot/Trap Fisheries
Leatherback turtles are commonly entangled in the vertical lines of
pot and trap gear. Entanglements have been mostly reported from U.S.
and Canadian waters, but line entanglements have occurred in other
areas where similar gear is used (e.g., Britain; Godley et al. 1998).
Due to high numbers of entanglement reports, a Sea Turtle
Disentanglement Network (STDN) was established by NMFS in the
northeastern United States (Maine to Virginia) in 2002. This program
relies primarily on reports from the public and subsequent
documentation and disentanglement by trained responders. From 2008 to
2017, 267 leatherback entanglements were reported in vertical fishing
line (STDN unpublished data). Of those fisheries that could be
identified, 79 were lobster, 21 were fish traps or fish lines, 18 were
conch (or a combination of conch and lobster), and 5 were crab gear;
144 entanglements were from unidentifiable fishing gear. While most
unknown vertical line entanglements likely involve pot/trap gear, this
cannot always be conclusively determined. The majority of the
leatherback turtle reports (67 percent) were from Massachusetts waters.
Of the 267 leatherback entanglements, 221 were released alive and 46
were found dead.
Given the nature of their injuries, it is probable that not all
animals released alive from entanglements survived. Currently there are
limited empirical data on leatherback survival from pot/trap
entanglements. Innis et al. (2010) found that at least some of the
disentangled individuals were able to resume normal behavior and
migratory patterns, but two leatherback turtles were entangled at least
twice, and a third disentangled turtle had significant forelimb skin
and muscle injuries. The effects of entanglement may be sub-lethal
initially, but could result in subsequent mortality. By assessing the
injuries experienced by each turtle that was documented to have been
entangled and using NMFS' post-interaction mortality guidance (NMFS
2017), the resulting mortality rate for northeastern U.S. vertical
fishing line interactions for all sea turtle species combined was
calculated at 55 percent from 2013 to 2017 (NMFS unpublished data).
When the mortality estimate includes those turtles that were not
disentangled and assumed to have died, the rate increases to 61
percent. As a result (and applying the latest 5 year mortality rate to
the last 10 years of entanglement data), 147 to 163 leatherback turtles
died from vertical fishing line gear (most of which were likely pot/
trap gear) in the northeastern U.S. waters from 2008 to 2017, based on
opportunistically reported data. An additional 36 leatherback turtles
were reported entangled in trap/buoy lines from North Carolina to Texas
from 2008 to 2017 (STSSN unpublished data). Of those 36 entanglements,
32 turtles were found alive and 4 dead, but these southeastern U.S.
numbers do not incorporate potential post-interaction mortality so the
total lethal interactions were likely higher. Further, this information
is likely an underestimate of actual
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entanglements and mortality given the opportunistic reporting nature of
the program; therefore, it is clear that leatherback interactions with
vertical fishing lines are a threat to this DPS.
Entanglements in Canadian waters are also frequently reported under
circumstances similar to the U.S. STDN program, i.e., opportunistically
by fishermen or the public. Between 1998 and 2014, 205 leatherback
entanglements were reported in Canada along the Atlantic coast, with
most from Nova Scotia (136) and Newfoundland (40; Hamelin et al. 2017).
Entanglements mostly involved pot fisheries (44 percent; n = 91),
including snow crab (n = 37), inshore lobster (n = 31), rock crab (n =
10), whelk (n = 8), and hagfish (n = 3) fisheries. Trap net fisheries
were involved in 26 percent of the entanglements (n = 53). Of the
overall 205 reports, the majority of turtles were reported alive and
successfully released (n = 174), and the other 15 percent (n = 31) were
reported dead in gear. However, the number of dead turtles is likely an
underestimate of actual entanglement-associated mortality (Hamelin et
al. 2017).
Leatherback turtles are also found entangled in vertical fishing
lines in European waters. Since 1980, 83 leatherback turtles were
bycaught in British and Irish waters, with the method of capture
identified in 58 cases (Piedpoint 2000). The majority of captures (n =
36) were rope entanglements, usually buoy lines used in pot fisheries
for crustaceans or whelk, with a 61 percent recorded mortality
(Pierpoint 2000).
Some types of aquaculture use vertical lines similar to pot/traps
and may pose an entanglement risk (Price et al. 2017). Four leatherback
turtles (two alive, two dead) in Canadian and U.S. waters have been
opportunistically reported in aquaculture gear to date (Price et al.
2017). However, as this industry is anticipated to grow in the near
future, leatherback interactions with aquaculture lines, and subsequent
injury or mortality, may increase.
These data comprise the best available information on pot/trap
fishery interactions with the NW Atlantic DPS. However, due to the high
probability of underreporting leatherback turtle entanglements by
fishers, the ad hoc nature of public reporting, and the uncertainty
about post-release survivorship, the leatherback mortality rate due to
entanglements in vertical lines is likely underestimated (Hamelin et
al. 2017). Estimates indicate that approximately 622,000 vertical lines
are deployed from fishing gear in U.S. waters from Georgia to the Gulf
of Maine (Hayes et al. 2018). There are currently no existing
mitigation measures to reduce leatherback bycatch in vertical fishing
lines, but efforts to reduce the amount of vertical lines in the water
to assist with large whale conservation in the United States may help
reduce the impact to the DPS (https://www.greateratlantic.fisheries.noaa.gov/protected/whaletrp/).
Other Gear Types
Leatherback turtles are also susceptible to bycatch in pound nets,
weirs, and purse seine fisheries. In the United States, pound nets set
in Virginia waters have entangled leatherback turtles. On June 23,
2006, NMFS issued a regulation (71 FR 36024) requiring offshore pound
nets set in a portion of the lower Chesapeake Bay from May 6 through
July 15 of each year to use modified pound net leaders, a gear
modification consisting of vertical hard lay lines spaced at least two
feet apart on the top portion of the leader, and eight inch or smaller
stretched mesh on the bottom portion of the leader. From 2013 to 2017,
16 leatherback turtles have been found entangled in the hard lay lines
of the leaders, of which two were dead (NMFS 2018). While individuals
may continue to be entangled in modified pound net leaders, the impact
of the pound net fishery on the NW Atlantic DPS is likely minor given
the few nets set in the lower Chesapeake Bay using this gear
(approximately four to six) and the frequency of live interactions.
From 2008 to 2017, the STDN also documented leatherback captures in
weirs set off Massachusetts; these turtles were found alive, either
entangled in the netting (n = 2) or free swimming in the weir (n = 4).
Purse seines are used to catch a variety of fish species and are
commonly used in the ICCAT area to catch tuna (Angel et al. 2014).
Leatherback captures have occurred in Atlantic purse seine fisheries,
and this bycatch may have a minor impact on the DPS. In British and
Irish waters, two leatherback turtles were reported to be captured in
purse seine gear between 1980 and 2000 (Pierpont 2000). Clermont et al.
(2012) reported a total capture of 67 leatherback turtles in more than
9000 observed Atlantic purse seine sets between 1995 and 2011, with
only four found dead (representing 10 percent observer coverage). Most
of the interactions were adults (75 percent). However, not all of the
purse seine effort reported by Clermont occurs in the NW Atlantic DPS
range. Thus, purse seine interactions with this DPS may be a fraction
of the total captures reported. For those purse seines in the ICCAT
region using fish aggregating devices and for those setting over free-
swimming tuna schools, the effort (through 2011) was concentrated in
the tropics, off West Africa between Namibia and Mauritania and off
Venezuela (Clermont et al. 2012; Angel et al. 2014). While leatherback
and purse seine interactions may occur where distribution and effort
overlap, the magnitude of the purse seine impacts on the NW Atlantic
DPS is lower than the bycatch values presented in Clermont et al.
(2012). Further, Angel (2014) found that the direct impacts on turtles
from purse seine fishing operations appears to be minor in comparison
to the impacts from longline fishing, especially as most purse seine
captures are released alive.
Summary of Fisheries Bycatch
We conclude that most immature and adult leatherback turtles of
this DPS are exposed to bycatch in multiple fisheries throughout their
range. Bycatch in gillnet fisheries, in particular, is a major threat
with high mortality rates (Lee Lum 2006; Gilman et al. 2010; Girondot
2015), annually killing thousands of NW Atlantic leatherback turtles.
When set off nesting beaches, gillnets result in high mortality of
nesting females and mature males (Lee Lum 2006; Eckert 2013). Longline
bycatch is considered to be a widespread threat throughout the DPS and
a primary source of leatherback mortality (Lewison et al. 2004),
resulting in the death of thousands of leatherback turtles annually. In
general, bycatch mortality reduces abundance by removing individuals
from the population. When nesting females are killed, it also reduces
productivity. We conclude that fisheries bycatch is the primary threat
to the NW Atlantic DPS.
Vessel Strikes
Vessel strikes are a threat to the NW Atlantic DPS. Injuries from
vessel strikes may include blunt force trauma and propeller parallel
slicing wounds affecting the carapace, flippers, head, and/or
underlying organs (Work et al. 2010). Most of what is known about
vessel strikes comes from stranding records; the most extensive
stranding network is found in the United States: The Sea Turtle
Stranding and Salvage Network (STSSN). In the United States (Maine
through Texas), 957 leatherback turtles were reported stranded,
captured, or entangled from 2008 to 2017, and of those, 204 had
probable vessel-related injuries (STSSN unpublished data). For example,
at least 72 leatherback turtles stranded in
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Massachusetts with vessel strike wounds between 2006 and 2018,
including at least three adult females that had previously been
documented nesting in the Caribbean (Dourdeville et al. 2018; Mass
Audubon Wellfleet Bay Wildlife Sanctuary, unpublished data, 2019). It
is sometimes difficult to determine whether the vessel related wounds
occurred before or after the turtle died (Stacy et al. 2015). However,
a recent study estimated that approximately 93 percent of Florida
stranded turtles with vessel strike wounds were killed by those
injuries (Foley et al. 2019). Based on the best available information,
it is reasonable to conclude that approximately 190 leatherback turtles
were killed as a result of vessel strikes in U.S. Atlantic and GOM
waters from 2008 to 2017. This number is likely an underestimate as
strandings represent a small percentage of turtles that are injured or
die at sea, and many vessel strikes are not reported, detected, or
recovered.
Vessel strikes have been documented in other nations as well,
including in Portugal (Nicolau et al. 2016), Britain (Godley et al.
1998), and off the coast of Tunisia in the Strait of Sicily (Karaa et
al. 2013; Caracappa et al. 2017). While there is very limited
observational information on vessel collisions in Atlantic waters of
Canada, there has been at least one recorded vessel strike (DFO 2012).
More recently, an injury assessment of leatherback turtles (n = 228) on
Atlantic Canada foraging grounds and on a Trinidad nesting beach found
only 1.3 percent of turtles exhibited injuries consistent with vessel
strikes (Archibald and James 2018). However, this low injury rate may
indicate that there is low survivorship of vessel strikes. Females with
carapace damage from propellers have been also observed on Costa Rican
nesting beaches (de Haro et al. 2006).
Leatherback behavior data can help predict the potential for vessel
strikes. Based on telemetry data for leatherback turtles (n = 15) on
the northeastern U.S. shelf, leatherback turtles spent over 60 percent
of their time in the top 10 m of the water column and over 70 percent
of their time in the top 15 m (Dodge et al. 2014). Additional turtle-
borne camera and autonomous underwater vehicle research in the waters
off Massachusetts suggests that turtles surface frequently and engage
in subsurface swimming (within the top 2 m) when occupying shallow,
well-mixed, coastal environments, increasing the probability of a
vessel strike (Dodge et al. 2018). Based on 24 free swimming
leatherback turtles tagged in Canadian waters from 2008 to 2013,
Wallace et al. (2015) found these leatherback turtles primarily
occupied the upper 30 m of the water column and had shallow 4 to 6
minute dives. Given most leatherback activity occurs in the top 15 to
30 meters of the water column in temperate shelf waters of the NW
Atlantic Ocean and vessel traffic is high along the U.S. East coast,
the risk of vessel strikes is likely higher than the documented
interactions would suggest (DFO 2012; Hamelin et al. 2014).
While observational data are limited, it is reasonable to conclude
that, based upon the best available information, mortality due to
vessel strikes may occur wherever vessel traffic and leatherback
distribution (juvenile and adult) overlap. The impact is likely
minimized in areas with less frequent vessel traffic (e.g., less
developed areas) and decreased leatherback turtle presence. Nesting
females and mature males may be especially vulnerable to vessel strikes
because they occur in the waters off nesting beaches, which are coastal
areas where vessel traffic is more prevalent. Vessel strikes affect the
NW Atlantic DPS by lowering abundance (if the interaction results in
mortality) and affecting future reproductive potential (productivity)
when nesting females are killed. We conclude that vessel strikes pose a
threat to the NW Atlantic DPS.
Pollution
Pollution includes contaminants, marine debris, and ghost fishing
gear. The detection of pollution impacts on leatherback turtles is
opportunistic and thus likely underestimated. While plastic ingestion
is not always fatal, it can reduce ability to feed, affect swimming
behavior and buoyancy control, potentially lead to chemical
contamination and chronic effects, and weaken physical condition, which
could impair the ability to avoid predators and survive threats (Nelms
et al. 2016). Entanglement in marine debris results in injuries that
can reduce fitness, cause eventual death, reduce ability to avoid
predators, reduce ability to forage and/or swim efficiently due to
drag, and lead to starvation or drowning (Nelms et al. 2016). Pollution
on the beach and in the water occurs throughout the range of the NW
Atlantic DPS.
Dow et al. (2007) defined marine pollution as agriculture,
petroleum, sewage, industrial runoff, vessel discharges, declining
water quality, and marine debris. They found pollution in the marine
environment to be among the greatest threats to all sea turtle species
in the Wider Caribbean Region. Dow et al. (2007) defined beach
pollution as agriculture, petroleum/tar, sewage, industrial runoff, and
beach litter/debris; they found pollution on the beach to be a threat.
Pollution on the beach and in the water occurs throughout the range of
the NW Atlantic DPS.
Leatherback turtles are susceptible to adverse effects from
pollution. Marine pollution, including direct contamination and
structural habitat degradation, can also affect leatherback habitat. In
particular, the Mediterranean is an enclosed sea, so organic and
inorganic wastes, toxic effluents, and other pollutants rapidly affect
the ecosystem (Cami[ntilde]as 2004).
Of particular concern, due to their immune, reproductive, and
endocrine disrupting nature, are persistent organic pollutants (POPs),
such as polychlorinated biphenyls (PCBs), polybrominated diphenyl
ethers (PBDEs), and pesticides (Bergeron et al. 1994; Bishop et al.
1991, 1998; Keller et al. 2004). These chemicals have been identified
in both adults and eggs in several areas occupied by this DPS. Guirlet
et al. (2010) measured maternal transfer of organochlorine contaminants
(OCs) from 38 nesting females in French Guiana. PCBs were found to be
the dominant OC, followed by pesticides, but OC concentrations were
lower than concentrations measured in other marine turtles (potentially
due to the lower trophic level diet and offshore foraging areas). All
OCs detected in nesting adults were detected in eggs, suggesting a
maternal transfer of OCs. In French Guiana, hatching success has been
shown to be low when OCs are present in the sand (most likely
originating from pesticide use in plantations and malaria prophylaxis
(Guirlet 2005). However, a link between OCs and embryonic mortality
could not be determined (Guirlet et al. 2010). Stewart et al. (2011)
also recorded PCB, OC, and PBDE concentrations for nesting and stranded
leatherback turtles in the southeastern United States. Their results
also suggested maternal transfer of POPs in leatherback turtles, but
Stewart et al. (2011) found higher levels of PCBs and pesticides than
those found in French Guiana (Guirlet et al. 2010). While finding that
leatherback contaminant concentrations were substantially lower than
concentrations in other reptile studies that demonstrated toxic
effects, Stewart et al. (2011) suggested that sub-lethal effects
(especially on hatchling body condition and health) may nevertheless be
occurring in this species. De Andres et al. (2016) similarly monitored
PCB and PBDE concentrations in eggs laid in Costa Rica (18 nests). POP
levels were similar to those reported in French
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Guiana nesting females (Guirlet et al. 2010) and slightly lower than
those in Florida (Stewart et al. 2011). Further, De Andres et al.
(2016) found a significant negative relationship between PBDE levels
and hatching success, suggesting potential harmful effects of these
contaminants on leatherback reproduction. OCs (and mercury) have also
been documented in turtles that stranded in the United Kingdom (Godley
et al. 1998). A leatherback that stranded off the coast of Wales, U.K.
was found with PCB levels one-to-three orders of magnitude higher than
the lowest levels reported for fish taken in the North Atlantic, but
similar to the lowest concentrations reported from oceanic cetaceans
(Davenport et al. 1990). Even with the recent restriction of the use of
POPs, due to the widespread persistent nature of these chemicals and
continuing atmospheric deposition (Ross et al. 2009) it is probable
that similar chemical concentrations occur in other areas of this DPS.
Other contaminants have also been documented in leatherback turtles
and their eggs. Heavy metals (e.g., arsenic, cadmium, chromium,
mercury, lead, etc.) enter the environment from a variety of sources
(Guirlet et al. 2008; Perrault 2012). In particular, mercury can affect
a variety of functional processes in wildlife, including the nervous,
excretory and reproductive systems (Wolfe et al. 1998). Mercury,
cadmium, and lead were recorded in nesting females (n = 46) and eggs in
French Guiana (Guirlet et al. 2008). Maternal transfer of all three
elements was documented, and female lead levels increased throughout
the nesting season (Guirlet et al. 2008). This could be explained, in
part, by external contamination via ingestion of contaminated prey or
polluted water during nesting, as the French Guiana coast environment
is exposed to significant environmental pollution via anthropogenic and
natural sources. While mercury concentrations were lower than values
reported for other sea turtle species, cadmium levels documented in
French Guiana were at the same level shown to impact gonadal
development in other turtle species and may impact reproductive
processes and lower fertility (Guirlet et al. 2008). In Massachusetts,
entangled turtles had significantly higher blood lead concentrations
than directly captured turtles (Innis et al. 2010). While similar to
those reported in French Guiana (Guirlet et al. 2008), blood
concentrations of mercury and cadmium were at levels high enough to
induce carcinogenic, teratogenic, and toxic effects in a variety of
species (Innis et al. 2010).
Mercury and selenium have also been recorded in nesting females and
eggs in Florida and St. Croix. Animals persistently exposed to mercury
can experience selenium deficiency, which is of concern because
selenium is important to hatching and emergence success (Perrault et
al. 2011). However, high levels of selenium can be toxic and negatively
impact hatching success (Perrault et al. 2013). Mercury concentrations
in nesting females from Florida were found to be higher than in St.
Croix, which could be a result of different migratory and foraging
areas, whereas hatchling blood mercury values were higher in St. Croix
(Perrault et al. 2011; Perrault et al. 2013). It is interesting to note
that in St. Croix, no correlations were found between mercury or
selenium concentrations and hatching or emergence success, which is
different from results in Florida (Perrault et al. 2011; Perrault et
al. 2013). Hazard quotient results by Perrault et al. (2013, 2014)
imply that mercury and selenium levels could pose a threat to
leatherback turtle reproductive success and/or hatchling health and
survival. Leatherback hatching and emergence success rates are already
low compared to other species of sea turtles (Bell et al. 2004;
Perrault et al. 2011), so the impacts of pollution and contamination on
hatching success is a notable concern. In addition, mercury was found
to be higher in adults than juveniles/sub-adults stranded along the
U.S. Atlantic coast, suggesting potential physiological concerns due to
accumulation and ongoing inputs into the environment (Perrault et al.
2012). It is clear that additional long-term research is needed to
better understand the relationship of non-essential elements in turtle
development and reproduction.
Marine debris (most notably plastic pollution) is a threat
throughout the range of the NW Atlantic DPS (Girondot 2015). Several
global reviews have outlined the persistent and widespread nature of
the issue, both as an ingestion and an entanglement threat (Mrosovsky
et al. 2009; Schuyler et al. 2014; Nelms et al. 2016; Lynch 2018). Law
et al. (2010) assessed plastic content at the surface of the western
North Atlantic Ocean and Caribbean Sea from 1986 to 2008, and found the
highest concentration of plastic debris was observed in subtropical
latitudes and associated with large-scale convergence zones, which
include foraging areas targeted by leatherback turtles.
Ingestion of marine debris is a concern for leatherback turtles,
especially given the similarity of their preferred prey (e.g.,
gelatinous zooplankton) to some plastics. In particular, plastic bags
appear similar to jellyfish in the marine environment, leading to
mistaken and inappropriate triggering of the sensory cue to feed
(Schuyler et al. 2014; Nelms et al. 2016). While plastic ingestion is
not always fatal, it can reduce ability to feed, affect swimming
behavior and buoyancy control, potentially lead to chemical
contamination and chronic effects, and weaken physical condition, which
could impair the ability to avoid predators and survive threats (Nelms
et al. 2016).
Marine debris ingestion can occur in any location, but given the
enclosed nature of the sea and intense human pressure, the
Mediterranean Sea in particular is a hot spot for plastic marine debris
and other pollutants (Cami[ntilde]as 2004; Cozar et al. 2015). Marine
debris ingestion has been documented from leatherback turtles stranded
in Tunisia (Karaa et al. 2013), Israel (Levy et al. 2005), the northern
Adriatic Sea (Poppi et al. 2012), and the Strait of Sicily (Caracappa
et al. 2017). Of particular note, 30 to 73 percent of turtles stranded
in the Bay of Biscay (France) were found to have ingested plastic
annually from 1979 to 1999 (out of 87 leatherback turtles necropsied;
Duguy et al. 2000). The seasonal rate of ingestion was inversely
related to the abundance of jellyfish, leading the authors to propose
that the depletion of jellyfish led to debris ingestion as potential
prey. Cozar et al. (2015) conclude that the effects of plastic
pollution on marine life are anticipated to be frequent in the high
plastic-accumulation region of the Mediterranean Sea.
In U.S. waters, marine debris ingestion has also been documented in
stranded leatherback turtles. However, ingestion does not always cause
mortality and is typically an incidental finding. Of 41 leatherback
turtles necropsied from North Carolina to Texas between 2008 and 2017,
17 had ingested plastics or marine debris (STSSN unpublished data
2018). From Maine to Virginia during that same time period, 10
necropsies detected ingestion, but the total number of necropsied
turtles, out of the 677 strandings in the region, is currently unknown.
It is likely that many more stranded turtles ingested some level of
marine debris (STSSN unpublished data 2018). Out of 33 leatherback
turtles examined in New York Bight (an area with dense population), 30
percent had
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synthetic material ingestion, mostly consisting of thin, clear plastic
(Sadove et al. 1989). Of two leatherback turtles stranded in North
Carolina during 2017 whose gastrointestinal tracts were analyzed,
microplastics were present in both (Duncan et al. 2018).
Marine debris ingestion is not limited to microplastics or plastic
bags. Off the northeastern U.S. coast, necropsies of disentangled
leatherback turtles that have died post-release have documented
considerably large pieces of plastic (e.g., 83 by 35 cm) in their
stomachs (Innis et al. 2010). These numbers likely underestimate the
true marine debris ingestion rate because many turtles likely ingest
marine debris and do not strand.
Leatherback turtles can also become entangled in marine debris.
From 2008 to 2017, the Northeast U.S. STDN documented 24 entanglements
from miscellaneous sources not attributed to obvious fisheries
entanglements, as described above (STDN unpublished data). These
unknown entanglements could involve a myriad of sources but are
considered as entangling marine debris. The Sea Turtle Recovery Action
Plan for the Republic of Trinidad and Tobago noted that entanglement in
lost or abandoned fishing gear (primarily nets) poses a threat to
leatherback turtles in the marine and terrestrial environment (Forestry
Division et al. 2010).
Marine debris is also a problem on nesting beaches and can reduce
nesting success. Pollution and debris often are deposited on high
energy beaches, which are also the preferred nesting habitat of
leatherback turtles (TEWG 2007). Coastal and inland littering (which
can ultimately reach the sea) is a problem throughout Trinidad and
Tobago, and ocean borne debris is particularly prevalent on the east
and north coasts, which host the main leatherback nesting beaches
(Trinidad and Tobago Forestry Division et al. 2010). Extensive debris
on nesting beaches is not uncommon throughout the Caribbean, often
carried by rivers to the sea and later washed ashore (e.g., in Costa
Rica; Chac[oacute]n-Chaverri and Eckert 2007). Debris on nesting
beaches may impede females during the nest-site selection stage, limit
and degrade the amount of habitat available, and/or result in aborted
nesting attempts (Chac[oacute]n-Chaverri and Eckert 2007). If line or
netting is encountered on nesting beaches, entanglement of nesting
females and hatchlings is also a risk.
The majority of the NW Atlantic DPS is exposed to pollution
throughout all life stages. These threats are a result of the developed
nature of many of the nations within the range of the DPS. The degree
of impact is difficult to quantify, especially given the widespread
nature of pollution and the diverse types of impacts. Contaminants may
affect this DPS by reducing productivity, if hatching success is
lowered, and by lowering abundance, if contamination results in
mortality. Marine debris affects the DPS by lowering abundance, when it
causes death through ingestion or entanglement, and reducing
productivity, when hatchlings and nesting females are affected. While,
we do not have quantitative estimates of the number of individuals that
are killed or injured as a result of pollution, we conclude that it is
prevalent throughout the range of the DPS and constitutes a threat to
the NW Atlantic DPS.
Oil and Gas Exploration
Oil and gas activities have the potential to impact the NW Atlantic
DPS directly (e.g., exposure to oil following oil spills) and
indirectly (e.g., increased probability of vessel strikes and habitat
degradation/destruction). In addition to lethal effects, sublethal
effects may occur and include displacement from primary foraging areas
with accompanying energy costs (TEWG 2007).
Several areas within the range of the NW Atlantic DPS have intense
oil and gas development and exploration close to major nesting beaches.
The potential for oil spills is of particular concern in the Wider
Caribbean Region due to its effect on all life stages in the marine
environment. The biggest oil producing nations in South America are
Brazil, Mexico, Venezuela, and Colombia. Although only three Caribbean
nations currently have exportable oil and natural gas reserves
(Barbados, Cuba, and Trinidad and Tobago, with Trinidad and Tobago the
only significant exporter), in 2017, a major oil field was discovered
off Guyana, which will likely lead to extensive new development and
extraction. As a result, marine traffic is likely to increase in the
area as well as the possibility for oil spills. In Panama,
contamination from oil spills, primarily in area of the Trans-Isthmus
oil pipeline and the Panama Canal, is of particular concern
(Br[auml]utigam and Eckert 2006; Ruiz et al. 2006). Some Caribbean
nations (e.g., Belize, French Guiana) have permanent moratoria on oil
and gas exploration in offshore waters.
In the United States, oil and gas extraction primarily occurs in
the GOM (BOEM 2016; BOEM 2017), an area with leatherback foraging and
migratory habitat (Aleksa et al. 2018). Increased shipping traffic and
marine noise due to oil and gas explorations in the GOM pose a direct
threat for leatherback turtles in foraging grounds and migratory
routes, due to the potential for vessel strikes and harassment (Wallace
et al. 2017; Ward 2017). Oil spills regularly occur in the GOM, from
small amounts of varying types of oil product to large catastrophic
spills. In 2010, a major oil spill occurred in the north-central GOM,
affecting important foraging habitat used by leatherback turtles
(Deepwater Horizon NRDA Trustees 2016). Evans et al. (2012) tracked a
post-nesting leatherback from Chiriqui Beach, Panama, into the GOM
during the Deepwater Horizon oil spill. The track followed similar
tracks from turtles in previous years and did not seem to change once
entering areas with visible oil slicks (on two occasions). Injuries to
leatherback turtles caused by the GOM Deepwater Horizon oil spill could
not be quantified (Deepwater Horizon NRDA Trustees 2016). However,
given that the GOM is important habitat for leatherback turtles (Aleksa
et al. 2018) and leatherback turtles were documented in the Deepwater
Horizon oil spill zone during the oil spill period, the Deepwater
Horizon NRDA Trustees (2016) concluded that leatherback turtles were
exposed to Deepwater Horizon oil, and some portion of those exposed
likely died.
In Atlantic Canada, impacts from oil and gas may also occur.
Several petroleum production projects occur offshore of Nova Scotia
(https://www.cnsopb.ns.ca/offshore-activity/offshore-projects). Howard
(2012) determined that oil pollution from coastal refineries, ships,
small engine vessels, and oil and gas exploration and production is a
risk to leatherback survival in Canada. There are also offshore oil and
gas platforms in the North (United Kingdom, Denmark) and Mediterranean
Seas, where similar impacts to leatherback turtles may also occur (EU
Offshore Authorities Group 2018; https://euoag.jrc.ec.europa.eu/node/63). In particular, the Mediterranean Sea has been declared a ``special
area'' by the International Convention for the Prevention of Pollution
from Ships (MARPOL), in which deliberate petroleum discharges from
vessels are banned, but numerous repeated offenses are still thought to
occur (Pavlakis et al. 1996). Some estimates of the amount of oil
released into the region is as high as 1,200,000 metric tons (Alpers
1993). Direct oil spill events also occur, as in Lebanon in 2006 when
10,000 to 15,000 tons of heavy fuel oil spilled into the eastern
[[Page 48358]]
Mediterranean (UN Environment Programme 2007).
In summary, oil and gas activities are prevalent in foraging,
migratory, and offshore nesting habitats of the NW Atlantic DPS,
potentially exposing all life stages to oil associated threats, such as
direct miring in oil, oil ingestion, vessel strikes, and nesting beach
contamination. Oil and gas activities have the potential to affect this
DPS by reducing productivity (e.g., if hatching success is reduced by
oil spills) and potentially lowering abundance (e.g., if oil exposure
results in mortality). As such, oil and gas activities are a threat to
the NW Atlantic DPS.
Natural Disasters
Natural disasters, such as hurricanes and other storms, and natural
phenomena, such as Sargassum events on or near nesting beaches, pose a
threat to the NW Atlantic DPS.
Hurricanes are common in the Caribbean and southeastern United
States. Hurricanes and tropical storms impact nesting beaches by
increasing erosion and sand loss and depositing large amounts of
debris. In 2017, Hurricane Maria devastated the islands of Dominica,
St. Croix, and Puerto Rico, and even though the nesting season was
nearly over, many beaches were impacted, including Maunabo, Puerto Rico
(one of the most abundant nesting beaches on the island; R. Espinoza,
Conservaci[oacute]n ConCiencia, pers. comm., 2017). Dewald and Pike
(2014) found that a lower level of leatherback nesting attempts
occurred on sites that were more likely to be impacted by hurricanes.
These types of storm events may ultimately affect the amount of
suitable nesting beach habitat, potentially resulting in reduced
productivity, especially as leatherback turtles typically nest on high
energy beaches (TEWG 2007).
Hurricanes may also result in egg loss by destroying and inundating
nests. However, hurricanes are usually aperiodic so the impacts are
expected to be infrequent. Hurricanes also typically occur after the
peak of the leatherback hatching season and would not be expected to
affect the majority of incubating nests (USFWS 1999). That said,
according to the Intergovernmental Panel on Climate Change (IPCC),
climate change may be increasing the frequency and patterns of
hurricanes (IPCC 2014) potentially causing such impacts to nests to
become more common in the future.
Sargassum is a genus of macroalgae found in temperate and tropical
waters. When large amounts of Sargassum wash ashore, they form thick
mats that have the potential to disrupt females' nesting activities and
impede hatchlings' access to the ocean (Maurer et al. 2015). In 2011
and 2015, large amounts of Sargassum were present in the Caribbean
(mainly Trinidad and Tobago and Grenada) and frequently washed ashore,
covering large expanses of sandy shoreline on nesting beaches. While
females still nested in these areas, hatchlings needed intervention to
reach the ocean (Wang and Hu 2016; Audroing, TVT, pers. comm., 2018; K.
Charles, Ocean Spirits Inc., pers. comm., 2018). Most recently, large
amounts of Sargassum were found in 2018 on Caribbean beaches, causing
Barbados to declare a national emergency in June 2018. Such widespread
blanketing of Sargassum on leatherback nesting beaches throughout the
Caribbean has the potential to impact future hatching success and
survival.
In summary, natural disasters and phenomena have the potential to
impact the NW Atlantic DPS. However, given the infrequent and temporary
nature of the occurrences, only a small proportion of eggs, hatchlings,
and nesting females are exposed to these threats. Impacts include egg
and hatchling mortality that affect productivity of the DPS. Seasonal
losses at individual beaches may be large, but we do not expect such
impacts to be spatially or temporally widespread. However, we conclude
that natural disasters pose a threat to the DPS.
Climate Change
Climate change is a threat to the NW Atlantic DPS. The impacts of
climate change include increases in temperatures (air, sand, and sea
surface); sea level rise; increased coastal erosion; more frequent and
intense storm events; and changes in ocean currents. These impacts may
affect leatherbacks through alterations of the incubation environment,
reduction of nesting habitat, and changes in prey as described in the
following subsections.
Modeling results show that global warming (rise in average surface
temperature) poses a ``slight risk'' to females nesting in French
Guiana and Suriname relative to those nesting in Gabon, Congo, and West
Papua (Dudley et al. 2016). As global temperatures continue to
increase, some beaches will experience changes in sand temperatures,
which in turn will alter the thermal regime of incubating nests.
Changing sand temperatures at nesting beaches may result in changing
sex ratios of hatchling cohorts and reduced hatching output (Hawkes et
al. 2009). Leatherback turtles exhibit temperature-dependent sex
determination (Binckley and Spotila 2015) and warmer temperatures
produce more female embryos (Mrosovsky et al. 1984; Hawkes et al.
2007). In the NW Atlantic DPS, the pivotal temperature (the temperature
at which a sex ratio of 1:1 is produced) is estimated to be between
29.25 [deg]C and 30.5 [deg]C (Eckert et al. 2012) but there are
variations in measurements (Girondot et al. 2018), over time, and among
locations. An increase over that temperature would result in more
female hatchlings. Such increases in female hatchling output have
already been documented (Pati[ntilde]o-Mart[iacute]nez et al. 2012),
and with an increase in temperatures from climate change, these trends
are likely to continue if other nesting factors remain constant. For
example, Pati[ntilde]o-Mart[iacute]nez et al. (2012) developed a model
to relate measured incubation temperature to sex ratio and estimated
that females nesting at Caribbean Colombian beaches currently produce
approximately 92 percent female hatchlings. Under all future climate
change scenarios, complete feminization could occur as soon as 2021
(Pati[ntilde]o-Mart[iacute]nez et al. 2012). In St. Eustatius,
leatherback hatchling production was female biased from 2002 to 2012,
with less than approximately 24 percent of males produced every year
(Lalo[euml] et al. 2016). Future warming air temperatures will
exacerbate this female bias, and female leatherback sex ratios are
projected to consistently reach 95 percent after 2028 on that island,
which has dark and light sand beaches (Lalo[euml] et al. 2016). Warming
trends in Costa Rica are expected to be higher than the global average
and resulting female-biased sex ratios are also expected (Gledhill
2007). While the assumption is that most nesting beaches will become
female-biased due to increased sand temperatures, this may not be the
case in all areas. In Grenada, increased rainfall (another effect of
climate change) was found to have a cooling influence on nests, so that
more male producing temperatures (less than 29.75 [deg]C) were found
within the clutches (Houghton et al. 2007). Further, due to the
tendency of nesting females to deposit some clutches in the cooler
intertidal zone of beaches, the effects of long-term climate on sex
ratios may be mitigated (Kamel and Mrosovsky 2004; Pati[ntilde]o-
Mart[iacute]nez et al. 2012).
Hatching success is affected by warming temperatures. Extremely
high sand/nest temperatures are anticipated to result in embryonic
mortality (Gledhill 2007, Santidri[aacute]n Tomillo et al. 2012,
Valentin-Gamazo et al. 2018). In Costa Rica, warmer conditions can
[[Page 48359]]
exacerbate the effects of biotic contamination and mold infestations of
developing embryos (Gledhill 2007), resulting in reduced hatching
success.
Temperature increases are likely to be associated with more extreme
precipitation and faster evaporation of water, leading to greater
frequency of both very wet and very dry conditions that reduce
productivity (Pati[ntilde]o-Mart[iacute]nez et al. 2014;
Santidri[aacute]n Tomillo et al. 2015). These impacts may affect nests
in different ways, but the result (e.g., reduced hatching output) is
similar. Very wet conditions may inundate nests or increase fungal and
mold growth, reducing hatching success (Pati[ntilde]o-Mart[iacute]nez
et al. 2014). Very dry conditions may affect embryonic development and
decrease hatchling output. Under climate change scenarios, very dry
conditions are expected for St. Croix, an area already showing
decreased productivity and reduced first time nesting female abundance
(Santidri[aacute]n Tomillo et al. 2015; Garner et al. 2017).
Santidri[aacute]n Tomillo et al. (2015) assessed climatic conditions on
hatchling output at four nesting sites (Sandy Point, St. Croix;
Pacuare, Caribbean Costa Rica; Playa Grande, Pacific Costa Rica;
Maputaland, South Africa), and found that St. Croix had the highest
projected warming rate (+ 5.4 [deg]C), highest absolute temperature and
lowest precipitation levels. With these further increases in dryness
and air temperatures, hatchling productivity is expected to be
compromised by the end of the 21st century in this area
(Santidri[aacute]n Tomillo et al. 2015). Santidri[aacute]n Tomillo et
al. (2015) suggested that the lack of rain is what reduces
developmental success and hatchling emergence. However, Rafferty et al.
(2017) evaluated long-term climate data for St. Croix, using climate
data collected from a nearby weather station, and found no significant
trend in incubation temperatures or precipitation that could be
associated with observed decreases in productivity at this location.
Finally, incubation temperatures can also influence hatchling
morphology and locomotion (Mickelson and Downie 2010). Leatherback
hatchlings originating from nests incubated at lower temperatures
exhibited carapace and front flipper length-width ratios that
significantly improved their crawling speeds relative to those
hatchlings incubated at high temperatures (Mickelson and Downie 2010).
Sea level rise is another threat to leatherback turtles. Thornalley
et al. (2018) found that the Labrador Sea deep convection and the
Atlantic Meridional Overturning Circulation, a system of ocean currents
in the North Atlantic, have been unusually weak over the past 150 years
or so, and this weakened state may have modified northward ocean heat
transport, as well as atmospheric warming by altering ocean-atmosphere
heat transfer. Further, the documented weakening of this system is
related to above-average sea level rise along the U.S. East Coast
(Caesar et al. 2018). Sea level rise may result in intensified erosion
and loss of nesting beach habitat (Fish et al. 2005; Fuentes et al.
2010; Fonseca et al. 2013). In Bonaire, up to 32 percent of the current
beach area could be lost with a 0.5 m rise in sea level, with lower,
narrower beaches being the most vulnerable (Fish et al. 2005). Ussa
(2013) predicted a 20 to 25 percent loss in beach areas due to sea
level rise by the year 2100 within the Archie Carr National Wildlife
Refuge, Florida, as well as areas adjacent to the Refuge. With the
threat of increasing sea level rise, protection of developed coastlines
often involves shoreline armoring that reduces the amount of beach
available, thus creating a smaller amount of space for turtles to nest
(Hawkes et al. 2009). Along such developed coastlines, rising sea
levels may cause severe effects on eggs, because nesting females are
forced to deposit eggs seaward of shoreline armoring, potentially
subjecting them to repeated tidal inundation and/or egg exposure from
exacerbated wave action near the base of these structures.
Sea level rise is expected to result in more nests being inundated,
reducing hatching success. On Playona Beach, Colombia, Pati[ntilde]o-
Mart[iacute]nez et al. (2014) found that nests in wet sand suffered
higher mortality (emergence success of zero percent for wettest nests
to 64 percent for the driest nests), suggesting that nesting success
should be expected to decrease under future climate change sea level
rise scenarios. Inundation is likely to reduce hatching success
(Pati[ntilde]o-Mart[iacute]nez et al. 2008; Caut et al. 2010) and will
continue to occur (or worsen) with sea level rise.
However, leatherback turtles may be less susceptible than other
species of sea turtles to loss of nesting habitat, because they exhibit
lower nest-site fidelity (Dutton et al. 1999). Nesting beaches in the
Guianas are already highly dynamic and interseasonally variable, and
leatherback nesting females have been successful in those areas despite
the fact that some beaches disappear between nesting years (Plaziat and
Augustinus 2004; Kelle et al. 2007; Caut et al. 2010). If global
temperatures increase and there is a range shift northwards, beaches
not currently used for nesting could in the future become used by
leatherback turtles, potentially offsetting some loss of accessibility
to beaches in southern portions of the range. Leatherbacks' behavioral
flexibility may allow for opportunities to colonize new beaches, but
whether turtles can colonize nesting areas that become available,
either thermally or geographically, by climate change, and whether
these colonized areas provide incubation regimes that will lead to
successful nesting, emergence success, and hatchling fitness cannot be
known at this time (Hawkes et al. 2009).
Observed changes in marine systems are associated with other
aspects of climate change, including rising water temperatures, as well
as related changes in ice cover, salinity, oxygen levels, and
circulation. Ocean temperatures of the U.S. northeastern continental
shelf and surrounding NW Atlantic waters have warmed faster than the
global average over the last decade (Pershing et al. 2015). New
projections for the U.S. northeastern shelf and NW Atlantic Ocean
suggest that this region will warm two to three times faster than the
global average and existing projections from the IPCC may be too
conservative (Saba et al. 2015). This increase in northeastern shelf
waters is relevant for NW Atlantic leatherback turtles, as they rely on
U.S. and Canadian waters to forage during the warmer months (James
2005a, 2006b, 2007; Dodge 2014, 2015).
Global warming is expected to expand leatherback foraging habitats
into, and increase residency time in, higher latitude waters (James et
al. 2006a; McMahon and Hays 2006; Robinson et al. 2009). For example,
leatherback turtles have extended their range in the Atlantic north by
around 200 km per decade over the last two decades as warming has
caused the northerly migration of the 15 [deg]C sea surface temperature
(SST) isotherm, the lower limit of thermal tolerance for leatherback
turtles (McMahon and Hays 2006). Documented weakening of the Meridional
Overturning Circulation is related to above-average warming in the Gulf
Stream region and an associated northward shift of the Gulf Stream
(Caesar et al. 2018). This weakening of the deep, cold-water
circulation in the North Atlantic is likely to continue to occur with
global warming. Migratory routes may be altered by climate change as
increasing ocean temperatures shift range-limiting isotherms north
(Robinson et al. 2009). Post-nesting females from French Guiana were
found to migrate northward toward the Gulf Stream north wall, targeting
similar habitats in terms of physical characteristics, i.e., strong
gradients of
[[Page 48360]]
SST, sea surface height, and a deep mixed layer (Chambault et al.
2017). Hatchling dispersal may also be affected by changes in surface
current and thermohaline circulation patterns (Hawkes et al. 2009; Pike
2013).
The effects of global warming are difficult to predict, but changes
in reproductive behavior (e.g., remigration intervals, timing and
length of nesting season) could occur (Hawkes et al. 2009; Hamann et
al. 2013). Robinson et al. (2014) found that the median nesting date at
Sandy Point (St. Croix) occurred on average 0.17 days earlier per year,
between 1982 and 2010. However, Neeman et al. (2015) found that
increased temperatures at the foraging grounds tend to delay
leatherback nesting. Temperatures at the nesting beaches (Playa Grande,
Costa Rica; Tortuguero, Costa Rica; and St. Croix) did not affect the
timing of leatherback nesting (Neeman et al. 2015). Because the
relation between temperatures (local sea surface and the foraging
grounds) and timing of nesting is complex, Neeman et al. (2015)
indicated that further study is needed at the nesting beaches to
determine how environmental conditions change within the season and how
these changes affect nesting success. Robinson et al. (2014) suggests
that shifts in the nesting phenology may make the Atlantic populations
more resilient to climate change.
Extreme precipitation events over most of the mid-latitude and
tropical regions will very likely become more intense and more frequent
(IPCC 2014). Changes in the frequency and timing of storms or changes
in prevailing currents could lead to increased beach loss via erosion
(Van Houtan and Bass 2007; Fuentes and Abbs 2010). More frequent and
intense storm events will have the same effect on leatherback nesting
success as previously described for natural disasters.
In summary, climate change is likely to affect multiple life stages
of turtles in the NW Atlantic DPS. Likely impacts include altering sex
ratios and reducing nest success, reducing nesting beach habitat and
nests due to sea level rise and storms, and potentially changing
distribution. Climate change therefore has the potential to alter
productivity and diversity. These impacts could be more severe in
certain areas with more dynamic beach environments, or could be
widespread throughout the DPS. Impacts are likely to range from small,
temporal changes in nesting season to large losses of productivity.
That said, leatherback turtles are considered to be the best able to
cope with climate change of all sea turtle species due to their wide
geographic distribution and relatively weak nesting site fidelity.
Overall, we conclude that climate change is a threat to the NW Atlantic
DPS.
Conservation Efforts
Next we consider ``conservation efforts'' under Section 4(b)(1)(A)
(16 U.S.C. 1533(b)(1)(A)).\1\ There are numerous efforts to conserve
the leatherback turtle. The following conservation efforts apply to the
NW Atlantic DPS (for a description of each effort, please see the
section on conservation efforts for the taxonomic species): African
Convention on the Conservation of Nature and Natural Resources (Algiers
Convention); Central American Regional Network; Convention on the
Conservation of Migratory Species of Wild Animals; Convention on
Biological Diversity; Convention on International Trade in Endangered
Species of Wild Fauna and Flora; Convention Concerning the Protection
of the World Cultural and Natural Heritage (World Heritage Convention);
Convention for the Protection and Development of the Marine Environment
of the Wider Caribbean Region, Specially Protected Areas and Wildlife
(SPAW); Convention on the Conservation of European Wildlife and Natural
Habitats; Convention for the Co-operation in the Protection and
Development of the Marine and Coastal Environment of the West and
Central African Region (Abidjan Convention); Memorandum of
Understanding Concerning Conservation Measures for Marine Turtles of
the Atlantic Coast of Africa (Abidjan Memorandum); Convention for the
Protection and Development of the Marine Environment of the North East
Atlantic; Convention on Nature Protection and Wildlife Preservation in
the Western Hemisphere (Washington or Western Hemisphere Convention);
Convention for the Protection and Development of the Marine Environment
of the Wider Caribbean Region (Cartagena Convention); Cooperative
Agreement for the Conservation of Sea Turtles of the Caribbean Coast of
Costa Rica, Nicaragua, and Panama (Tri-Partite Agreement); Council
Regulation (EC) No. 1239/98 of 8 June 1998 Amending Regulation (EC) No.
894/97 Laying Down Certain Technical Measures for the Conservation of
Fishery Measures (Council of the European Union); Council Directive 92/
43/EEC on the Conservation of Natural Habitats and of Wild Fauna and
Flora (EC Habitats Directive); Food and Agricultural Organization (FAO)
Technical Consultation on Sea Turtle-Fishery Interactions; Inter-
American Convention for the Protection and Conservation of Sea Turtles
(IAC); MARPOL; Inter-American Tropical Tuna Convention (IATTC); IUCN;
North American Agreement for Environmental Cooperation; Protocol
Concerning Specially Protected Areas and Biological Diversity in the
Mediterranean; Ramsar Convention on Wetlands; Regional Fishery
Management Organizations (RFMOs); UN Convention on the Law of the Sea
(UNCLOS); and UN Resolution 44/225 on Large-Scale Pelagic Driftnet
Fishing. Although numerous conservation efforts apply to the turtles of
this DPS, they do not adequately reduce its risk of extinction.
---------------------------------------------------------------------------
\1\ For a related discussion of existing regulatory mechanisms
to protect turtles, which are considered separately under Section
4(a)(1)(D), see the discussion above at ``Inadequacy of Existing
Regulatory Mechanisms.''
---------------------------------------------------------------------------
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the NW Atlantic DPS is at high risk of extinction. The total index
of nesting female abundance is 20,659 females at consistently monitored
beaches, and the most recent annual rate of decline is estimated to be
approximately nine percent (NW Atlantic Leatherback Working Group
2018). The best available nest data reflect a steady decline for more
than a decade, becoming more pronounced since 2008 (Eckert and Mitchell
2018; NW Atlantic Leatherback Working Group 2018). This decreasing
trend is observed when all available nest data are combined and at most
nesting beaches (NW Atlantic Leatherback Working Group 2018), including
the largest nesting aggregation in Trinidad (i.e., Grande Riviere,
which is declining at 6.9 percent annually). In terms of productivity,
the DPS exhibits low hatching success, while other key parameters such
as clutch size, remigration interval, and clutch frequency are similar
to species' averages. There are also indications of decreased
productivity within the DPS at one of the most intensively monitored
nesting beaches (i.e., Sandy Point, St. Croix; Garner et al. 2017). The
declining region-wide nest trend and potential changes in productivity
make the DPS highly vulnerable to threats.
However, the DPS exhibits broad spatial distribution and some
diversity. Based upon genetic data, as well as flipper tagging and
satellite telemetry data, this DPS shows significant spatial structure
with some connectivity among nesting and foraging areas. Further,
nesting occurs in a variety of habitats,
[[Page 48361]]
including islands and mainland, as well as muddy, sandy, and shelly
beaches. The DPS uses multiple, distant, and diverse foraging areas,
including oceanic and coastal waters throughout the North Atlantic
Ocean, Mediterranean Sea, and GOM, providing some resilience against
reduced prey availability. While the numerous and diverse nesting and
foraging locations, along with moderate levels of genetic diversity,
provide some level of buffer to the DPS, the highest concentrations of
nesting occur in Trinidad, French Guiana, and Panama, where a
catastrophic event could have a disproportionate impact on the DPS.
The primary threat to the NW Atlantic DPS is bycatch in commercial
and artisanal, pelagic and coastal fisheries. Gillnet fisheries, in
particular those off nesting beaches, are the greatest concern given
the high mortality rate. In particular, the coastal surface drift
gillnet fishery off Trinidad kills an estimated 1,000 adult leatherback
turtles annually (Lee Lum 2006; Eckert et al. 2008; Eckert 2013).
Bycatch, and subsequent mortality, in Trinidad bottom set gillnets and
surface gillnets in Suriname and French Guiana are major threats to the
NW Atlantic DPS. Trinidad and French Guiana host the highest number of
nesting females in this DPS, so the continued mortality of adults in
that area is of significant concern. Further, no adequate regulatory
mechanism is currently in place (e.g., no gear modifications or
closures) to address this incidental bycatch. These fisheries and the
related mortality rates have been occurring for years (Lee Lum 2006;
Eckert 2013). Longline fisheries are the most widespread threat,
occurring throughout the Atlantic Ocean by fisheries from multiple
nations, incidentally capturing thousands of leatherback turtles
annually based on the best available data. Longline gear modifications
(e.g., circle hooks) are sometimes, but not consistently, used. Fishery
bycatch in pot/trap gear, especially off the northeastern U.S. coast
and in Canadian waters, and trawls are also significant threats.
Fisheries bycatch reduces abundance by removing individuals from the
population; when those individuals are nesting females, it reduces
productivity as well. Given the lack of observer coverage and
reporting, cumulative mortality due to fisheries bycatch is likely
higher than available estimates.
Additional threats to the DPS include habitat loss, the legal and
illegal harvest of turtles and eggs, predation, vessel strikes,
pollution, climate change, oil and gas activities, and natural
disasters. Coastal development and armoring, erosion (natural and
anthropogenic), and artificial lighting are some of the most
significant stressors on nesting beach habitat, reducing nesting and
hatching success (i.e., productivity). Habitat loss and modification is
also anticipated to increase over time with additional development and
climate change. Legal and illegal harvest of turtles and eggs reduces
abundance and productivity. Illegal egg poaching occurs in several
nations, particularly Costa Rica, Dominican Republic, and Colombia.
While reduced in some nations, illegal poaching still occurs on
unmonitored beaches throughout most of the Caribbean, including
Suriname and Trinidad. While leatherback eggs and hatchlings are preyed
upon by many species, the biggest threat is from feral dogs. Egg
predation by dogs occurs in many nations, but it is a particular
concern in Colombia, French Guiana, Guyana, Panama, Puerto Rico, and
Trinidad and Tobago. Intervention (e.g., screening) to reduce predation
is not used in most places, partially due to the concern of attracting
poachers as well as the infeasibility of implementing effective
measures at high-density or remote beaches. Egg predation reduces
productivity.
Vessel strikes are also a threat, killing numerous leatherback
turtles each year. While exposure to vessel strikes may be most severe
in developed areas, the total impacts are high, affecting both
abundance and productivity. Pollution, ingestion of plastics, and
entanglement in marine debris are threats to all leatherback turtles,
most likely resulting in injury and compromised health, and sometimes
mortality. Exposure to pollution is widespread in the NW Atlantic
Ocean, but effect data are limited. Oil and gas activities are threats
with the potential to grow in some Caribbean areas. Natural disasters
(hurricanes) and phenomenon (large Sargassum events) have an
intermittent impact to the NW Atlantic DPS. Climate change is likely to
result in reduced productivity due to greater rates of coastal erosion
and sea level rise and subsequent nest inundation and habitat loss,
reduced hatching success, changing sex ratios, and distributional
changes. Although many international, national, and local regulatory
mechanisms are in place, they do not adequately reduce the impact of
these threats.
The cumulative impact of these multiple threats is potentially
large (Andersen et al. 2017). Innis et al. (2010) reported that many
individuals are simultaneously exposed to multiple threats, including:
entanglement, injury, plastic ingestion, adrenal gland parasitism,
diverticulitis, and burdens of environmental toxins (Innis et al.
2010). Such cumulative pressures affect individual survival and
productivity. In some cases, it is possible to directly link individual
threats to demographic reductions (e.g., high mortality in gillnets off
nesting beaches reduces nesting female abundance). More often, however,
several threats contribute synergistically to demographic reductions.
For example, reductions in hatching success may be caused by one or
more of the following threats alone or in combination: erosion,
poaching, predation, climate change, and pollution.
We find that the NW Atlantic DPS is affected by numerous severe
threats. These present, ongoing threats injure or kill turtles and
contribute to the declining nest trend. The Team evaluated whether the
DPS is at risk of extinction currently or would become so within the
foreseeable future. To answer this question, they asked how long it
would take for the total index of nesting female abundance to be
reduced by 50 percent, a drastic decline that would reduce abundance to
a level where demographic risks would present an independent threat to
the DPS's continued existence, and whether this time period places the
DPS at risk currently or within the foreseeable future. Using estimates
of the mean time to maturation for the population (approximately 19
years; Avens et al. in review) and mean nesting longevity
(approximately 11 years; Avens et al. in review) of the species, they
estimated a generation time of approximately 30 years. They then
considered three different scenarios. First, they calculated the time
until 50 percent reduction in the total index of nesting female
abundance using data on a significant and influential, well-documented,
threat: Gillnet bycatch mortality of 1,000 adult turtles annually off
the largest nesting aggregation, i.e., Trinidad. Assuming that half of
the turtles at Trinidad killed are female, total index of nesting
female abundance would decrease by 50 percent in 28 years, which is
approximately one generation.
Second, the Team used regional nest trend data from the NW Atlantic
Leatherback Working Group (2018). Using the most recent trends as is
typical for population projections (i.e., -9.32 percent per year from
2008 to 2017), they found that the total index of nesting female
abundance would fall by 50 percent within 8 years (95 percent CI: 6 to
13 years). Using trends from the longer data set (-4.21 percent per
year
[[Page 48362]]
from 1990 to 2017), the total index of nesting female abundance would
fall by 50 percent within 17 years (95 percent CI: 11 to 31 years).
Finally, using their calculation of nest trend for the highest
abundance nesting area in the DPS, Trinidad (-7.3 percent per year, 95
percent CI: -34 to 18 percent), the Team found that the total index of
nesting female abundance would decrease by 50 percent within 10 years
(95 percent CI: 3 years to ``never;'' however, ``never'' is highly
unlikely, given that there is a 75 percent likelihood that the true
value of the nest trend in Trinidad is negative (f = 0.754)). There are
several caveats with using nest trend data: Adult females typically
account for, at most, a small percentage of the population; trends in
nesting female abundance may not be an index of the remainder of
population; stable age distribution is assumed; and time series of
available data do not always span one generation (let alone multiple
generations required to reach stable age distribution). Despite these
caveats, all scenarios resulted in a 50 percent reduction in the total
index of nesting female abundance in less than one generation. While
the first scenario did not involve the use of nest trend data, it did
result in a 50 percent reduction within one generation when considering
only one threat (albeit the most severe), and we know that the DPS
faces many large-impact threats, (suggesting that the first scenario
understates the DPS's degree of risk).
For the purpose of the extinction risk analysis, the Team discussed
whether the resulting range of time periods (8 to 28 years) suggests a
present risk of extinction or a risk of extinction within the
foreseeable future. The Team did not have a unanimous view. All but one
Team member were present to vote on the level of extinction risk. Eight
Team members concluded with moderate confidence that the DPS is at high
extinction risk due to threats and the declining trend that has
accelerated in recent years. Their confidence was moderate rather than
high due to some resilience provided by the abundance, spatial
distribution, and diversity for this DPS. Two Team members concluded
with low confidence that the DPS is at moderate extinction risk. Their
confidence in this conclusion was low due to the declining trend that
has accelerated in recent years. The Terms of Reference called for a
simple majority, and after voting, the Team concluded that the DPS met
the definition for high risk of extinction. We agree with the Team's
overall conclusion that a 50 percent decline in less than one
generation equates to a current, high risk of extinction. We find
support for this conclusion in well documented examples of other
leatherback populations that have quickly declined despite larger
abundances (e.g., the Mexico nesting aggregation declined from 70,000
nesting females in 1982 to under 1,000 nesting females by 1994; Spotila
et al. 2000).
We conclude that the NW Atlantic DPS is presently in danger of
extinction due to the number and magnitude of threats, of which
fisheries bycatch is the greatest concern. These present and ongoing
threats have resulted in imminent and substantial demographic risks
(i.e., declining trends and reduced abundance). Although numerous
conservation efforts apply to the turtles of this DPS, they do not
adequately reduce the risk of extinction. We conclude that the NW
Atlantic DPS is in danger of extinction throughout its range and
therefore meets the definition of an endangered species. The threatened
species definition does not apply because the DPS is currently at risk
of extinction (i.e., at present), rather than on a trajectory to become
so within the foreseeable future.
SW Atlantic DPS
The Team defined the SW Atlantic DPS as leatherback turtles
originating from the SW Atlantic Ocean, north of 47[deg] S, east of
South America, and west of 20[deg] W; the northern boundary is a
diagonal line between 5.377[deg] S, 35.321[deg] W and 12.084620[deg] N,
20[deg] W. The southern boundary is based on the Antarctic circumpolar
current which prevents sea turtles from nesting further south. The
western end of the northern boundary is based at the ``elbow'' of the
Brazilian coast, where the Brazilian Current begins and likely
restricts the northern nesting range of this DPS. We placed the eastern
boundary at the 20[deg] W meridian as an approximate midpoint between
SW Atlantic and SE Atlantic (i.e., turtles that nest in western Africa)
nesting beaches and to reflect both DPS's wide foraging range
throughout the South Atlantic Ocean. However, due to its low abundance,
the SW Atlantic DPS is less likely to be encountered compared to the
more abundant SE Atlantic DPS.
The SW Atlantic DPS only nests on the southeastern coast of Brazil,
primarily in the state of Esp[iacute]rito Santo, on a continuous
stretch of beach, less than 100 km in length, with concentrated nesting
in Povoa[ccedil][atilde]o and Comboios. While there is occasional,
limited nesting south of these primary nesting beaches, the sand
becomes coarser further south, and the excavation of nests becomes more
difficult because the sand falls back into the holes (Thom[eacute] et
al. 2007).
While nesting is limited geographically, the overall range of this
DPS (i.e., all areas of occurrence) is extensive, as demonstrated by
individuals tracked to numerous foraging areas. Leatherback turtles of
this DPS use coastal waters off South America from the ``elbow'' of
Brazil southwards to Uruguay and Argentina, where quality foraging
areas allow for coastal foraging in addition to open-ocean foraging
(Almeida et al. 2011). Individuals of this DPS are also known to
migrate to the waters off western Africa and forage in the oceanic
habitat in between South America and Africa (Almeida et al. 2011).
Likewise, Prosdocimi et al. (2014) found 84 to 86 percent of
leatherback turtles sampled from the foraging grounds off Argentina and
Eleva[ccedil][atilde]o do Rio Grande (an elevated offshore area across
from Brazil) to originate from western African beaches.
Abundance
The total index of nesting female abundance for the SW Atlantic DPS
is 27 females. We based this index on nest monitoring data from Projeto
TAMAR, the Brazilian Sea Turtle Conservation Program, which has
established an index nesting survey area along 47 km of beach (10 km
along Povoa[ccedil][atilde]o and 37 km along Comboios; IAC Brazil
Annual Report 2018), where complete daily surveys have been conducted
during the primary nesting season from September through March, since
the 1986/1987 nesting season. Some nesting occurs along the non-index
stretches of Povoa[ccedil][atilde]o and the beaches to the northern
part of the area, but it is minor relative to nesting on the index
survey area (Thom[eacute] et al. 2007). To calculate the index of
nesting female abundance (i.e., 27 nesting females) for the
Esp[iacute]rito Santo index area, we divided the total number of nests
between the 2014/2015 and 2016/2017 nesting seasons (i.e., a 3-year
remigration interval; Thom[eacute] et al. 2007) by the clutch frequency
(5 clutches/season; Thom[eacute] et al. 2007; Tiwari et al. 2013).
Minimal, scattered nesting has been reported on beaches outside
Esp[iacute]rito Santo (Barata and Fabiano 2002; Thom[eacute] et al.
2007; Bezerra et al. 2014), but these beaches exhibit suboptimal sand
characteristics for nesting, limiting the possibility of substantial
nesting expansion into those areas (Thom[eacute] et al. 2007).
Therefore, while the nest counts from the index beach surveys do not
provide a full estimate of all nesting for
[[Page 48363]]
the DPS, they provide a high-quality dataset, account for the majority
of the nests (approximately 80 percent; Colman et al. 2019), and are
used for determining our index of nesting female abundance and the nest
trend in the next section.
Our total index of nesting female abundance is similar to the IUCN
Red List assessment's estimate of 35 mature individuals (female and
male, assuming a 3:1 sex ratio) based on nesting data through 2010
(Tiwari et al. 2013).
The total index of nesting female abundance (i.e., 27 nesting
females at the index beach) places the DPS at risk for environmental
variation, genetic complications, demographic stochasticity, negative
ecological feedback, and catastrophes (McElhany et al. 2000; NMFS
2017). These processes, working alone or in concert, place small
populations at a greater extinction risk than large populations, which
are better able to absorb losses in individuals. Due to its small size,
the DPS has limited capacity to buffer such losses. Given the intrinsic
problems of small population size, we conclude that the nesting female
abundance is a major factor in the extinction risk of the SW Atlantic
DPS.
Productivity
The SW Atlantic DPS exhibits an increasing, although variable nest
trend. Long-term monitoring data for this small DPS are limited to the
index nesting survey area in southeastern Brazil, where data has been
collected between the 1986/1987 and 2016/2017 nesting seasons. Over the
31-year data collection period, the mean annual number of nests for
these beaches was 35. While this is below the criterion of 50 annual
nests for conducting a trend analysis, we determined that this site
should nevertheless be included due to the high quality and consistency
of the data, and the fact that these data accurately represent the low
level of nesting of this DPS. The median increase in nest counts was
4.8 percent annually (sd = 5.8 percent; 95 percent CI = -8.4 to 15.5
percent; f = 0.832; mean annual nests = 35). As the index area hosts
the majority of known nesting activity, these data are representative
of the entire DPS. We conclude that nesting has increased from 1986 to
2017. Our trend estimate is similar to that of the IUCN Red List
assessment, which characterizes the population as increasing (Tiwari et
al. 2013). It is also in agreement with the recent study by Colman et
al. (2019), which describes the trend as increasing but variable, with
the mean annual number of nests increasing from 25.6 nests in the first
5 years to 89.8 nests in the last 5 years of monitoring (between 1988
and 2017).
While the long term trend indicates an increase in nesting, the
most recent 3 years of data (i.e., 30, 64, and 38 nests from 2014 to
2016) show a marked reduction in nests compared to the previous 3 years
(i.e., 78, 124, and 102 nests from 2011 to 2013). The reason for this
reduction is unknown. It could reflect declining nesting female
abundance or changes in productivity metrics (i.e., a longer
remigration interval or reduced clutch frequency) related to
environmental shifts or prey availability. Therefore, there is
uncertainty regarding whether the increasing trend will continue.
The productivity parameters for this DPS are fairly typical for the
species. In Brazil, the average clutch size appears to be on the lower
end of the range for Atlantic populations; conversely, Brazilian nests
tend to have a higher average number and percentage of eggs per clutch
(Thom[eacute] et al. 2007). Therefore, the egg production of this DPS
appears to be weighed more towards production of viable, hatchling-
producing eggs compared to other Atlantic populations (Thom[eacute] et
al. 2007). Nesting females produced an average of 3,496 hatchlings
annually over the past 10 years of nesting, which was calculated by
multiplying 60.4 nests annually, 87.7 eggs per nest, and 66.0 percent
hatching success (Colman et al. 2019). This estimate does not include
the limited nesting outside the index area. The mean size of nesting
females (CCL) has changed from 159.8 cm, with a range of 139 to 182 cm
(Thom[eacute] et al. 2007) to 152.9 cm 10.0 SD, with a
range of 124.7 to 182.0 cm; the decrease was statistically significant
and may indicate recruitment (Colman et al. 2019). Hatching success has
increased from a mean of 65.1 percent (with a range of 53.3 to 78
percent; Thom[eacute] et al. 2007) to a mean of 66 percent (with a
range of 38.8 to 82.4 percent; Colman et al. 2019).
While the overall nest trend for this DPS is increasing, there is
uncertainty regarding the continuation of this trend, given the data
for the past 3 years. The population remains extremely small, and thus
overall productivity is limited. Additionally, the potential for
population growth is not clear, given the limited suitable nesting
habitat available. We conclude that limited productivity places the DPS
at risk of extinction.
Spatial Distribution
The SW Atlantic DPS comprises a single, small nesting aggregation
concentrated on the beaches of one state in Brazil (Esp[iacute]rito
Santo). A tagging study has shown internesting movements along 300 km
of the coast, including over 100 km on either side of known nesting
beaches (Almeida et al. 2011), indicating connectivity throughout this
area. The nesting spatial distribution is extremely restricted, with
nesting constrained to a small area, with little suitable nesting
habitat into which it can expand. Conversely, the DPS exhibits a broad
foraging range, extending south to waters off Uruguay and Argentina,
throughout the pelagic waters of the South Atlantic, and across to
western Africa (Almeida et al. 2011).
The wide distribution of foraging areas likely provides some level
of buffer for the DPS against local catastrophes or environmental
changes that could limit prey availability. However, the limited
nesting range, and apparent lack of suitable nesting beaches into which
to expand, renders the DPS highly susceptible to detrimental
environmental impacts, both acute (e.g., storms and singular events)
and chronic (e.g., sea level rise and temperature changes). Any such
change would impact the entire extent of the DPS's nesting habitat.
With no metapopulation structure, the DPS has reduced capacity to
withstand other catastrophic events. Thus, despite widely distributed
foraging areas, the extremely narrow nesting distribution and lack of
population structure increases the extinction risk of the SW Atlantic
DPS.
Diversity
Despite its extremely low nesting female abundance, the Brazilian
nesting aggregation has the second-highest haplotype diversity among
all Atlantic populations (h = 0.498-0.532; Dutton et al. 2013; Vargas
et al. 2017). According to Thom[eacute] et al. (2007), while most
nesting occurs from September through March, sporadic nesting has been
recorded throughout the year, which may provide temporal resilience if
environmental conditions limit nesting during the primary nesting
season. The use of estuarine waters (of the Rio de la Plata) as a year-
round foraging ground is an unusual characteristic shared with the SE
Atlantic DPS (Lopez-Mendilaharsu et al. 2009; Prosdocimi et al. 2014).
Despite genetic and foraging diversity, the limited size and range of
the nesting aggregation reduces the resilience of this DPS.
[[Page 48364]]
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
Within the limited nesting range of the SW Atlantic DPS, habitat
modification is a threat. The 2015 collapse of a tailings dam at an ore
mine upstream of the index nesting survey area had an undetermined, but
potentially long-term, impact on the nesting beach of the DPS. Tens of
millions of cubic meters of heavy metal-laden mining waste entered the
Doce River and ultimately passed through the mouth of the river, in the
middle of the index nesting area. Nests laid near the river mouth were
relocated to prevent hatchlings from entering polluted waters. Hatching
success was not significantly different between years in the period of
2012 to 2017, which include three seasons before (2012-2014) and three
seasons after (2015-2017) the mining event (Colman et al. 2019). While
no difference was noted in the distribution of nests following the dam
breach, non-lethal impacts to individuals encountering the polluted
waters, especially hatchlings, could not be measured. Such impacts may
have occurred but may not be evident for decades following the spill.
Projeto TAMAR is monitoring for heavy metals in eggs and nesting
females and is closely watching for changes in fitness and reproductive
parameters (Thom[eacute] et al. 2017). As a result of the dam's
collapse, the Brazilian Federal government is implementing a marine
protected area (APA-Area de Protecao Ambiental da Foz do Rio Doce),
including about 100 kilometers of coastline, which should encompass the
entire extension of the index nesting beaches, with both coastline and
surrounding marine areas. Such a measure is an environmental
compensation for the dam's collapse, and should be implemented with
specific resources in the coming years (ICMBio, MMA, Brazil; J.
Thom[eacute], Projeto TAMAR, pers. comm., 2019).
Beach erosion and tidal flooding are also threats to this DPS.
According to Thom[eacute] et al. (2007), occasional relocation of nests
and nest protection occur when inundation or predation risk is
considered high. The majority of nests are relocated when in danger of
beach erosion or tidal flooding (J. Thom[eacute], Projeto TAMAR, pers.
comm., 2019).
Although coastal light pollution has been documented to be
increasing in Brazil, nesting has not been notably impacted thus far
(Colman et al. 2018). The lack of impact may be attributable to
conservation strategies including the creation of protected areas and
minimization of direct lighting on the nesting beaches. Nests are
relocated from heavily lit areas. All light sources with a light
intensity greater than 0 lux (lux = lumen per m\2\) on these beaches
are prohibited by a Federal ordinance (Portaria IBAMA 11/1995).
Construction, lighting, and poaching were not considered a significant
problem at the leatherback nesting beaches by Thom[eacute] et al.
(2007). However, such problems persist in several other turtle nesting
beaches in Brazil (Mascarenhas et al. 2004; Lara et al. 2016). More
recently, coastal development and artificial lighting have been
identified as potential threats for leatherback turtles on the beaches
of Esp[iacute]rito Santo (TAMAR/Unpublished data) and further research
is needed to better understand these threats. Nests are relocated from
heavily lit areas. Colman et al. (2018) found a negative relationship
between nest density and light levels. Additionally, as oil industry
and other economic developments are explored, the potential threat to
the nesting habitat may increase (Thom[eacute] et al. 2007).
A significant portion of the nesting beach is protected as a
Federal reserve under Brazilian Decree no. 90222 (September, 25 1984),
which covers 15 km of Comboios Beach, south of the mouth of the Doce
River. An additional 22 km, south of the reserve, falls within
indigenous land that has restricted access under Federal law. No
Federally protected areas exist north of the Doce River mouth, where
Povoa[ccedil][atilde]o Beach occurs. However, local, state, and Federal
regulations provide some coastal zone protections in that area.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization poses a threat to the SW Atlantic DPS. Though
specific information on leatherback turtle harvests is not available,
there was historically traditional harvest of sea turtles and eggs in
Esp[iacute]rito Santo (Hartt 1941; Medeiros 1983). This harvest,
however, has been largely curtailed through the work of Projeto TAMAR,
which promoted other economic activities and hired ex-turtle hunters to
protect nests (Marcovaldi et al. 2005; Almeida and Mendes 2007). The
capture of leatherback turtles was banned in Brazil in 1968, and full
protection for all sea turtles was enacted in 1986 (Marcovaldi and
Marcovaldi 1999). At present, egg poaching has been reduced, and there
is no known subsistence hunting for sea turtles of any species
(Thom[eacute] et al. 2007). As previously noted, there is protection
for or limited access to much of the nesting habitat south of the Doce
River. However, this protection does not extend north of the river,
where additional nesting occurs. Because of the very small size of the
population, even very low levels of egg poaching have the potential to
impact the population. Therefore, we conclude that overutilization
poses a threat to the SW Atlantic DPS.
Disease or Predation
While we could not find any information on disease for this DPS,
predation is a threat to the SW Atlantic DPS. Invertebrates, reptiles,
and mammals prey on eggs, while hatchlings fall prey to land, air, and
marine predators. According to Thom[eacute] et al. (2007), relocation
and protection of nests may be undertaken when inundation (primarily)
or predation (secondarily) risk is considered high (J. Thom[eacute],
Projeto TAMAR, pers. comm., 2019). Predators include foxes (Cerdocyon
thous), raccoons (Procyon cancrivorus), and domestic dogs, although
there are no quantitative estimates of predation for this DPS (J.
Thom[eacute], Projeto TAMAR, pers. comm., 2019). Some predation of
large juveniles and adults occurs in the marine environment, especially
by sharks (Bornatowski et al. 2012), but the frequency and impact on
those populations is not well understood. For this DPS, predation
primarily impacts productivity (i.e., reduced egg and hatching
success). We conclude that predation is a threat to the SW Atlantic
DPS, but that there is insufficient information to classify disease as
a threat.
Inadequacy of Existing Regulatory Mechanisms
The SW Atlantic DPS is protected by several regulatory mechanisms.
For each, the Team reviewed the objectives of the regulation and to
what extent it adequately addresses the targeted threat.
Beach habitat is protected throughout much of the nesting range of
this DPS. The vast majority of nesting occurs in Esp[iacute]rito Santo,
where beaches have been protected since 1982. All light sources with a
light intensity greater than 0 lux (lux = lumen per m\2\) on these
beaches are prohibited by a Federal ordinance (Portaria IBAMA 11/1995).
The take of leatherback turtles is illegal throughout the SW
Atlantic Ocean. Regional regulations include: Brazil Portaria, Manter
proibida a captura de tartarugas marinhas das esp[eacute]cies Caretta,
Dermochelys coriacea, Eretmochelys imbricata e Lepidochelys
[[Page 48365]]
olivacea \2\ No.27/1982; Uruguay Presidential Decree 144 and additional
legislation to reduce bycatch and prevent habitat alteration, and to
prevent the removal of individuals from their natural environment;
Argentina National Decree 666 from 1997; and various laws prohibiting
hunting and selling sea turtles. Harvest and consumption of sea turtles
are illegal under Brazilian law (Law on Environmental Crimes N[deg]
9605/1998). While these protections are mostly effective, very low
levels of egg poaching still exist (Thom[eacute] et al. 2007).
---------------------------------------------------------------------------
\2\ Prohibition of the capture of sea turtles of the species
Caretta caretta, Dermochelys coriacea, Eretmochelys imbricata, and
Lepidochelys olivacea.
---------------------------------------------------------------------------
Fisheries bycatch is the primary threat to the SW Atlantic DPS.
Although regulations address this issue to some extent, they do not do
so adequately and it continues to be a threat. In 2001, Brazil
established the National Plan for the Reduction of Incidental Capture
of Sea Turtles in Fishing Activities (Marcovaldi et al. 2005). However,
bycatch continues to be a major problem. In Brazil, the use of TEDs in
trawl fisheries is mandatory (Instru[ccedil][atilde]o Normativa MMA No.
31; December 13, 2004), but most fishermen nevertheless do not use such
gear, and there is little or no enforcement by authorities (IAC Brazil
Annual Report 2018). The UN established a worldwide moratorium on drift
gillnet fishing effective in 1992, the General Fisheries Commission for
the Mediterranean prohibited driftnet fishing in 1997, and the
International Commission for the Conservation of Atlantic Tunas (ICCAT)
banned driftnets in 2003. Despite these and other numerous regulations
and international instruments to protect sea turtles, significant
bycatch still occurs in artisanal and commercial fisheries operating in
the territorial waters of Argentina, Uruguay, and Brazil and on the
high seas (Gonz[aacute]lez et al. 2012).
In summary, while numerous regulatory mechanisms have been enacted
to provide some protections to leatherback turtles, their eggs, and
nesting habitat throughout the range of this DPS, they have been
inadequate. Many do not effectively reduce the threat that they were
designed to address, generally as a result of limited implementation or
enforcement. Fisheries bycatch, in particular, remains a major threat
to the DPS despite regulatory mechanisms. We conclude that the failure
to implement and enforce effective regulations is a threat to the DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat to the SW Atlantic DPS.
Leatherback turtles are captured as bycatch in commercial and artisanal
fisheries, along coastal foraging and breeding areas, and on the high
seas. The extensive foraging range of this DPS makes it vulnerable to
interactions with fisheries off the coasts of Brazil, Uruguay, and
Argentina, in the pelagic waters of the South Atlantic Ocean, and along
the coastal waters off western Africa. Recoveries of females tagged in
Esp[iacute]rito Santo are scarce, however. Three were found dead on the
Brazilian coast (incidentally captured in fisheries around the Doce
River mouth (TAMAR, unpublished data)), one in Argentina (Alvarez et
al. 2009), and one in Namibia, West Africa (Almeida et al. 2014).
Fisheries interaction information specific to this DPS is limited,
because the data do not differentiate among individuals from this DPS
and SE Atlantic individuals that forage within the same range. Because
the SE Atlantic DPS is much more abundant than the SW Atlantic DPS,
most fishery interactions likely involve SE Atlantic individuals.
However, data about bycatch involving the SE Atlantic DPS is
informative because the impact to the SW Atlantic DPS individuals is
likely to be proportional to their relative presence in the area.
Bycatch in gillnets; surface, deep-water longline hooks; and trawls are
the principal causes of sea turtle deaths, with not only higher
interaction numbers, but higher mortality rates than other fishery
interactions (Kotas et al. 2004; Pinedo and Polacheck 2004; Tudela et
al. 2005; Giffoni et al. 2013).
Coastal gillnet fisheries interactions are one of the largest
threats to the survival of the SW Atlantic DPS. In an analysis of
Brazilian fishery data from 1990 to 2012, Giffoni et al. (2013)
documented 237 leatherback turtle interactions, and 31 percent
mortality, in coastal set, fixed, encircling, and pelagic drift
gillnets. The actual number of interactions is likely substantially
higher, as many interactions go unreported.
Smaller scale artisanal gillnet fisheries occur in coastal waters
that are used by SW Atlantic individuals for mating, access to nesting
beaches, and foraging. Thom[eacute] et al. (2007) described the
occurrence of artisanal gillnet fisheries close to the nesting beach
but indicated that Brazil was investing resources in developing lower-
impact fishing techniques. However, such fisheries occur throughout
other important coastal foraging areas off South America. Additionally,
coastal artisanal gillnet fishery interactions with leatherback turtles
are known to occur off the western coast of Africa, where some
individuals from the SW Atlantic DPS forage (Riskas and Tiwari 2013).
The Rio de la Plata estuary, an important foraging area off Uruguay,
has numerous documented instances of leatherback turtle entanglements,
including mortalities from coastal bottom-set gillnet fisheries
(Fallabrino et al. 2006; Lopez-Mendilaharsu et al. 2009; Velez-Rubio et
al. 2013).
Larger-scale commercial ocean gillnet fisheries are also a
significant threat for the SW Atlantic DPS, with high bycatch rates
reported off Brazil in drift and set gillnets (Fiedler et al. 2012;
Ramos and Vasconcellos 2013). Drift gillnet fishing off Brazil started
in 1986, targeting hammerhead sharks (Domingo et al. 2006). Marcovaldi
et al. (2006) reported that leatherback turtles comprised about 70
percent of all sea turtles captured in Brazilian driftnet shark
fisheries. From 2002 to 2008, 351 sea turtles were incidentally caught
in 41 fishing trips and 371 sets. Leatherback turtles accounted for
77.3 percent of the take (n = 252 turtles, capture rate = 0.1405
turtles/km of net) with 22.2 to 29.4 percent of turtles dead upon
retrieval and no estimate of post-release mortality for those released
alive. The annual catch by this fishery ranged from 1,212 to 6,160
leatherback turtles, as estimated based on bootstrap procedures under
different fishing effort scenarios in the 1990s (Fiedler et al. 2012).
In 1998, a Brazilian Federal ordinance limited the use and transport of
bottom and drift gillnets over 2.5 km long. Such regulations were
difficult to enforce, and vessels from the ports of Itaja[iacute],
Navegantes and Porto Belo, Santa Catarina, Brazil, deployed nets up to
7,846 m long between 2005 and 2006 (Kotas et al. 2008). In 2010 the
ordinance was suspended, permitting unrestricted fishing with driftnets
(Fiedler 2012). The shark drift gillnet fishery declined steeply in
later years, with no vessels operating in 2009 (UNIVALI/CTTMar 2010)
likely because of target species reduction, reduced profitability, and
IBAMA Normative Instruction N166/2007 which temporarily stopped the
issuance of new driftnet fishing licenses and established a 2-year
deadline by which vessels were to replace driftnets with other gear.
Various other gillnet fisheries, such as bottom gillnets for sharks and
mollusks, have reported leatherback mortalities as well, such as that
occurring off Uruguay (Fallabrino et al. 2006; Laporta et al.
[[Page 48366]]
2006; Eckert et al. 2009) and the western coast of Africa (Riskas and
Tiwari 2013).
Longline fisheries pose a significant threat to the SW Atlantic
DPS, as the spatio-temporal distribution of leatherback turtles
overlaps with longline fishing effort (Fossette et al. 2014). In a
review of reported, observed takes in hook and line fishery (primarily
longline) interactions with leatherback turtles in all of Brazil from
1990 to 2012, 1061 takes were documented, with 3 percent of the taken
turtles found dead on the line and another 37.5 percent of unknown
condition after release (Giffoni et al. 2013). High frequencies of
leatherback deaths from bycatch have been documented on longline
fishing vessels from southern Brazil and Uruguay (Kotas et al. 2004;
Pinedo and Polacheck 2004; Domingo et al. 2006; Giffoni et al. 2008;
Monteiro 2008). Between 2004 and 2005, in a study off southern Brazil,
eight leatherback turtles were captured, with a mean capture rate of
0.03 turtles per 1,000 hooks (Monteiro 2008). In 1999, there were 70
longliners in the Brazilian fleet, with 33 vessels operating out of
southern Brazil and fishing a total of 13,598,260 hooks (ICCAT 2001).
However, the overall effort in the area was much higher, as longliners
from Uruguay, Chile, Japan, Taiwan, and Spain fish in this area (Folsom
1997; Weidner and Arocha 1999; Weidner et al. 1999). Scientific
observers documenting 10 trips by longline vessels from the Uruguayan
fleet operating in the SW Atlantic Ocean between 26[deg] and 37[deg] S
between April 1998 and November 2000 observed 27 incidentally caught
leatherback turtles (Balestre et al. 2003). The prevalence of
leatherback interactions in pelagic longline fisheries is likely a
result of the longline fleet fishing the productive areas in the
convergence zone of the Brazilian Current and the cold waters from the
Falklands Current (Kotas et al. 2004), which coincides with important
sea turtle foraging and developmental habitat. As with gillnets, the
scope of the longline threat to the SW Atlantic DPS spans across the
South Atlantic Ocean in both coastal and oceanic waters. In addition to
exposure to longline fisheries off South America, coastal longline
fisheries off Cameroon, Angola, and Namibia, and pelagic longlines in
the Gulf of Guinea and the eastern portion of the South Atlantic Ocean
have also been documented to take leatherback turtles (Honig et al.
2007; Riskas and Tiwari 2013; Angel et al. 2014; Huang 2015; Gray and
Diaz 2017). Additional evidence of longline interactions comes from the
stranding data, where flipper injuries on some of the stranded
leatherback turtles could have been caused by interactions with pelagic
longlines. Onboard observers in longline fisheries off Brazil have
reported that leatherback turtles tend to be foul-hooked in the flipper
rather than the mouth (Kotas et al. 2004; Pinedo and Polacheck 2004;
Lima 2007). In 2017, Brazil enacted a law (PORTARIA INTERMINISTERIAL No
74, DE 1o- DE NOVEMBRO DE 2017) requiring the use of circle hooks in
the pelagic longline fisheries as well as keeping specified dehooking
and gear removal equipment on board any Brazilian longline vessel.
Specifically, the Brazilian government required the use of 14/0 or
larger circle hooks for all longline vessels targeting swordfish or
tuna (https://www.jusbrasil.com.br/diarios/166677996/dou-secao-1-06-11-2017-pg-81).
Trawl fisheries also impact the SW Atlantic DPS, mainly along
coastal waters off southern Brazil, Argentina, and Uruguay (Gonzalez
Carman et al. 2011; Velez Rubio et al. 2013; Monteiro et al. 2016).
Although there are fewer interactions with trawl fisheries relative to
other fisheries (i.e., gillnet and longline fisheries), mortality rates
in trawl fisheries are far higher (Miller et al. 2006; Laporta et al.
2013). Observation of the Uruguayan bottom trawl fishery, during a
tagging and data collection program designed to increase the
understanding of the fishery impacts on sea turtles, documented 17
leatherback interactions from April 2002 to June 2005 (Laporta et al.
2013). Coastal bottom trawl and artisanal gillnet fisheries were the
main causes of death of leatherbacks found stranded in Uruguay (Velez
Rubio et al. 2013). Recorded interactions in coastal trawl fisheries
are also known from Gabon, Congo, and Namibia (Riskas and Tiwari 2013).
Other fisheries such as corrals, pound nets, and pots appear to
present a much lower risk for leatherback turtles than to other sea
turtle species. From 1990 to 2012, Giffoni et al. (2013) documented
only two leatherback turtles (both alive) of the 8,367 total sea
turtles taken in those fisheries.
While specific information is not available to permit calculating
an estimate of overall bycatch and mortality rates of SW Atlantic
leatherback turtles, it is clear that fisheries bycatch, especially in
gillnets and longlines, is a major threat to the DPS. Immature and
adult individuals are exposed to high fishing effort throughout their
foraging range and in coastal waters near nesting beaches. Bycatch
mortality is also high, with reported rates of up to 31 percent
(Giffoni et al. 2013). Mortality reduces abundance, by removing
individuals from the population; it also reduces productivity, when
nesting females are incidentally captured and killed. Given the small
size of the DPS, the loss of even a small number of individuals from
fishery interactions has the potential to reduce abundance and
productivity. Therefore, we conclude that fisheries bycatch is the
primary threat to the SW Atlantic DPS.
Vessel Strikes
There is little information regarding vessel strikes for the SW
Atlantic DPS. Many of the primary foraging areas for this DPS off the
coasts of Argentina, Uruguay, and Brazil are experiencing increased
vessel traffic from fishing vessels, cargo transport, and tourism
(L[oacute]pez-Mendilaharsu et al. 2009; Fossette et al. 2014), so
leatherback turtle interactions with vessels may occur. Affected
individuals likely include immature and mature turtles. Impacts range
from injury to mortality. We conclude from the best available
information that vessel strikes are likely a threat to the DPS.
Pollution
As with all leatherback turtles, entanglement in and ingestion of
marine debris and plastics is a threat that likely kills several
individuals a year. Multiple studies have implicated the ingestion of
marine debris and/or entanglement in cases of injury or death of
turtles found in waters occupied by the SW Atlantic DPS (Bugoni et al.
2001; Eckert et al. 2009; Schulyer et al. 2013; Scherer et al. 2014).
However, no individuals were assigned to a particular population and
could have been members of the more abundant SE Atlantic DPS, which is
known to occupy the same waters.
While there is no specific information on effects to leatherback
turtles of this DPS, pollution from various economic activities
including maritime transport, tourism, and domestic and industrial
waste discharges that are known to occur within their range, may also
have an impact (L[oacute]pez-Mendilaharsu et al. 2009; Fossette et al.
2014). Events such as the failure of a mining tailings dam in 2015 that
resulted in the discharge of tons of mining mud contaminated with heavy
metals into the Doce River, and subsequently into the waters off
Esp[iacute]rito Santo nesting beaches, are also a concern, though no
specific impacts to leatherback turtles have so far been noted from
that event (Garcia et al. 2017). There is also concern about the
potential for increased oil and gas exploration activities
(Thom[eacute] et al. 2007). The petroleum industry in Brazil
[[Page 48367]]
has implemented a beach monitoring program, along large stretches of
the Brazilian coast, including Esp[iacute]rito Santo, to monitor for
potential impacts to sea turtles and their nesting beaches from
industry activities (Werneck et al. 2018)
Assigning impacts of pollution specifically to individuals within
the SW Atlantic DPS is difficult, and the best available information
does not quantify such impacts. However, given its prevalence, we
conclude that pollution poses a threat to the DPS.
Climate Change
Climate change poses a threat to the SW Atlantic DPS. The impacts
of climate change include: Increases in temperatures (air, sand, and
sea surface); sea level rise; increased coastal erosion; more frequent
and intense storm events; and changes in ocean currents.
Because leatherback turtles nest lower on the beach than other sea
turtles, their eggs are more at risk of being exposed and destroyed by
increases in sea level and coastal erosion (Boyes et al. 2010).
Additionally, given the limited availability of suitable nesting
habitat, the loss of the current nesting habitat with no buffer area to
move into would pose a major problem for the DPS. Thus, rising sea
level and beach erosion are potential threats to the DPS.
While we do not have specific information on pivotal temperatures
and temperature thresholds for egg mortality for this DPS, sand
temperatures influence egg viability and sex determination. Given the
potential lack of suitable nesting habitat outside the area currently
being utilized, there is little opportunity for a spatial shift in
nesting in response to changing temperatures. This DPS exhibits some
year-round nesting, which provides a small measure of resilience to
counteract increasing temperatures. However, it is not likely to be
sufficient to make up for the loss of nesting habitat and opportunity
resulting from sea level rise and temperature increases. The impacts on
productivity and survivorship for such shifts in nesting are unknown.
The threat of climate change is likely to modify the nesting
conditions for the DPS. Adverse impacts on turtles of the SW Atlantic
DPS would be inescapable because the entire DPS is confined to a
limited nesting area. Impacts are likely to range from small, temporal
changes in nesting season to large losses of productivity. Therefore,
we conclude that climate change is a threat to the DPS.
Channel Dredging
There is evidence of interactions with hopper dredges associated
with channel dredging and maintenance. Between 2008 and 2014, four
leatherback turtles were killed by hopper dredges in Rio de Janeiro
(Goldberg et al. 2015).
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply turtles of the SW Atlantic DPS
(for a description of each effort, please see the section on
conservation efforts for the overall species): Southwest Atlantic Sea
Turtle Network, Convention on the Conservation of Migratory Species of
Wild Animals, Convention on Biological Diversity, Convention on
International Trade in Endangered Species of Wild Fauna and Flora,
Convention Concerning the Protection of the World Cultural and Natural
Heritage (World Heritage Convention), FAO Technical Consultation on Sea
Turtle-Fishery Interactions, IAC, MARPOL, IUCN, Ramsar Convention on
Wetlands, RFMOs, South Atlantic Association, UNCLOS, and UN Resolution
44/225 on Large-Scale Pelagic Driftnet Fishing. Although numerous
conservation efforts apply to the turtles of this DPS, they do not
adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the SW Atlantic DPS is at ``high'' risk of extinction. The DPS
exhibits a total index of nesting female abundance of 27 females at the
index beach. Such a nesting population size places this DPS at risk of
stochastic or catastrophic events that increase its extinction risk.
Although there has been substantial variability in nesting at the index
nesting beach since 1986, the nest trend shows a strong, nearly five
percent annual increase through 2017, with the largest increase
occurring in the past decade. However, nesting has declined in the past
3 years. There is only one nesting aggregation, limited to a relatively
small stretch (47 km) of beach along a single coast. Some nesting also
occurs outside that area, but is mostly sporadic and limited by sand
and temperatures unsuited for nesting. Thus, stochastic events have the
potential to have catastrophic effects on the entire DPS, with no
distant subpopulations serving as a buffer or source of additional
individuals or diversity. Based on these factors, we find the DPS to be
at risk of extinction as a result of its limited abundance, spatial
structure, and resilience.
Current threats place this DPS at further risk of extinction. The
primary threat to this DPS is bycatch in commercial and artisanal,
pelagic and coastal fisheries, especially gillnet and longline
fisheries. Fisheries bycatch reduces abundance by removing individuals
from the population. Because several fisheries operate near nesting
beaches, productivity is also reduced when nesting females are
prevented from returning to nesting beaches. Exposure to and impact of
this threat are high. Additional threats include: Habitat modification,
overutilization, predation, pollution, vessel strikes, and climate
change. Habitat modification includes incidents such as the mining dam
breach upstream of the Doce River, which flows into the ocean through
the middle of the primary nesting beach. Overutilization and predation
are threats for this DPS as well, though some protective measures
exist. While many laws are in place to protect sea turtles from fishery
impacts, the continued impact of bycatch indicates that regulatory
mechanisms are inadequate to sufficiently address the threat. Pollution
and vessel strikes are potentially increasing threats to the DPS.
Climate change is another threat that is likely to increase, resulting
in reduced productivity due to greater rates of coastal erosion and
nest inundation, and in some areas, nest failure or skewed sex ratios
due to increased sand temperatures.
We conclude, consistent with the Team's findings, that the SW
Atlantic DPS is currently in danger of extinction. The total index of
nesting female abundance make the DPS highly vulnerable to threats
despite the apparent increasing nesting trend. In addition, this DPS
consists of only one small nesting aggregation with limited potential
nesting beaches to the north and south for expansion. The limited
nesting range and small size makes the DPS highly vulnerable to
stochastic impacts in the natural environment as well as singular,
large-scale, anthropogenic events such as oil spills. Some degree of
resilience is provided by the use of multiple foraging areas across a
vast geographic area. However, that expansive foraging range also
exposes the DPS to numerous fisheries (which are coastal and on the
high seas, artisanal and commercial, off both South America and western
Africa), making fisheries bycatch by far the biggest threat to the DPS.
Although numerous conservation efforts apply to the turtles of this
DPS, they do not adequately reduce the risk of extinction.
[[Page 48368]]
We conclude that the SW Atlantic DPS is currently in danger of
extinction throughout its range and thus meets the definition of an
endangered species. The threatened species definition does not apply
because the DPS is at risk of extinction now (i.e., at present), rather
than on a trajectory to become so within the foreseeable future.
SE Atlantic DPS
The Team defined the SE Atlantic DPS as leatherback turtles
originating from the SE Atlantic Ocean, north of 47[deg] S, east of
20[deg] W, and west of 20[deg] E; the NW boundary is a diagonal line
between 12.084620[deg] N, 20[deg] W and 16.063[deg] N, 16.51[deg] W.
The eastern boundary occurs at the southern tip of Africa, where the
Agulhas and Benguela Currents meet. Along with the cold waters of the
Antarctic Circumpolar Current, these currents likely restrict the
nesting range of this DPS. We placed the western boundary at the
20[deg] W meridian as an approximate midpoint between SE Atlantic and
SW Atlantic (i.e., turtles that nest in Brazil) nesting beaches and to
reflect the DPS's wide foraging range throughout the South Atlantic
Ocean; this DPS is more likely to be encountered in these waters
compared to individuals from the less abundant SW Atlantic DPS. The
northern boundary is a diagonal line between the elbow of Brazil and
the northern boundary of Senegal because the SE Atlantic DPS does not
appear to nest above this boundary (Fretey et al. 2007).
The range of the SE Atlantic DPS is extensive, mirroring that of
the SW Atlantic DPS. While nesting occurs along the western coast of
Africa, data indicate that foraging areas and migratory paths stretch
along the Atlantic coast of Africa from Senegal to South Africa, across
the South Atlantic Ocean, and into the coastal waters of Brazil,
Uruguay, and Argentina. As with the SW Atlantic DPS, this DPS does not
appear to forage in northern latitudes.
All nesting for the SE Atlantic DPS occurs along the Atlantic coast
of western Africa, from Senegal to Angola, a nesting range of over
7,500 km. However, the vast majority of nesting occurs in Gabon,
Equatorial Guinea (including Bioko Island), and the Republic of Congo
(TEWG 2007; Fretey et al. 2007, Witt et al. 2009; Tiwari et al. 2013).
Gabon may have once hosted the largest nesting aggregation in the world
when it was discovered in the early 2000s (Witt et al. 2009), but
current data indicate much lower levels of nesting (Formia et al. in
prep) compared to those described in Witt et al. (2009).
While nesting occurs along the western coast of Africa, foraging
grounds and migratory paths stretch across the South Atlantic Ocean to
the coastal waters of Brazil, Uruguay, and Argentina. Because of the
greater abundance of this DPS, most individuals found in the western
South Atlantic along the coast of South America, and on the high seas,
belong to the SE Atlantic DPS. Prosdocimi et al. (2014) found 84 to 86
percent of leatherback turtles sampled from the foraging grounds off
Argentina and Eleva[ccedil][atilde]o do Rio Grande (an elevated
offshore area across from Brazil) to originate from western African
beaches.
Abundance
The total index of nesting female abundance for the SE Atlantic DPS
is 9,198 females. We based this total index on nine nesting
aggregations in Gabon (n = 8,495 nesting females), Equatorial Guinea (n
= 457), Republic of Congo (n = 69), Sierra Leone (n = 39), Liberia (n =
45), Ivory Coast (n = 40), Ghana (n = 4), Cameroon (n = 3), and Sao
Tome and Principe (n = 46). Our total index does not include 10
unquantified nesting aggregations in Guinea-Bissau, Angola, and other
nations. For more information on data sources and calculations, please
see the Status Review Report.
Our total index of nesting female abundance is an index because we
do not have consistent data from much of the nesting range of the DPS,
which extends from Senegal to Angola. However, the largest nesting
aggregations occur in Gabon, Equatorial Guinea (including Bioko
Island), and the Republic of Congo (TEWG 2007; Fretey et al. 2007; Witt
et al. 2009; Tiwari et al. 2013), which are represented in our total
index. The IUCN Red List assessment did not provide an estimate of
population size but instead concluded that the subpopulation was ``data
deficient'' (Tiwari et al. 2013).
To calculate the index of nesting female abundance in Gabon, where
annual aerial surveys of 600 km of nesting beaches gather emergence
data, we used a remigration interval of 3 years, a clutch frequency of
7.8 clutches per season per female, and estimated that 95 percent of
emergences resulted in nesting (Formia et al. in prep). Our index of
nesting female abundance for Gabon (i.e., 8,495 nesting females) is
lower than previous estimates. According to Witt et al. (2009), Gabon
once hosted the largest leatherback nesting aggregation in the world,
with an estimated 36,185 to 126,480 clutches per year (approximately
15,730 to 41,373 nesting females). These estimates were based on a
combination of aerial surveys and ground-truthing surveys, conducted
during the 2002/2003, 2005/2006, and 2006/2007 nesting seasons. More
recent aerial surveys indicate a steep decline in nesting since the
early 2000s, with a high of 108,588 estimated nests in 2002/03, a low
of 4,275 estimated nests in 2009/10, and fewer than 25,000 nests in the
final year of available data (2015/16; Formia et al. in prep).
Nesting is scattered on continental Equatorial Guinea (Fretey
2001), but it occurs on several beaches of Bioko Island and is
monitored at the Gran Caldera Scientific Reserve (n = 457 nesting
females, based on body pit data from the 2000/2001 through 2017/2018
nesting seasons; D. Venditti et al., Drexel University, pers. comm.,
2018). Rader et al. (2006) documented an average of 3,896 nests
annually between the 2000/2001 to 2004/2005 nesting seasons, which
equates to approximately 2,338 nesting females (i.e., using a 3-year
remigration interval and a clutch frequency of 5 nests annually). Based
on the data available on nesting in the Republic of Congo from the
2003/2004 to 2016/2017 nesting seasons (N. Breheret, SWOT, pers. comm.,
2018), we estimated 69 nesting females. In an analysis of older data
(1999 to 2008), Girard et al. (2016) estimated 933 nests per year on
the monitored beaches, which equates to approximately 560 nesting
females.
In Guinea-Bissau, only one beach is monitored regularly, in Orango
National Park, Bijagos Archipelago, where occasional leatherback
nesting tracks are recorded. Each season, a few nests are reported
elsewhere throughout the nation (Barbosa et al. 1998; Fretey et al.
2007).
In the Ivory Coast (n = 40 nesting females), Gomez (2005) counted
218 nests over 41 km of beach in February 2001. Pe[ntilde]ate et al.
(2007) reported 189 nests reported from non-exhaustive surveys of 27 km
of coastline during the 2001/2002 nesting season.
In Ghana, nest monitoring occurs on three beaches: Mankoadze (n = 4
nesting females), Ada, and Keta. We were unable to calculate the index
for Ada and Keta beaches because we only received information on nest
averages. From 2000 to 2017, an annual average of 34 nests were
observed on Ada Beach (D. Agyeman, pers. comm., 2018). During the 2006/
2007 nesting season, 481 leatherback nests were counted on Ada Beach
(Allman and Armah 2010). Over an unspecified time frame, an annual
average of 80 nests were observed on Keta Beach (A. Fuseini, pers.
comm., 2018).
[[Page 48369]]
In Cameroon (n = 3 nesting females; Fretey and Nibam unpublished
data 2018), Girard et al. (2016) estimated an average of 43 leatherback
nests annually, which would equate to 26 nesting females, from 1999 to
2008. In S[atilde]o Tom[eacute] and Principe (n = 46 nesting females),
Girard et al. (2016) estimated an average of 78 nests annually from
1999 to 2008, which is similar to our estimate.
Nesting occurs on other beaches throughout western Africa. However,
recent consistent and standardized monitoring data are not available.
Sporadic nesting occurs in Senegal (Maigret 1978; Dupuy 1986), Republic
of The Gambia (Barnett et al. 2004, Hawkes et al. 2006), Togo
(Segniagbeto 2004), Nigeria (Fretey 2001; Mojisola et al. 2015),
Democratic Republic of Congo, (OCPE-ONG 2006), and Angola (Carr and
Carr 1991; Weir et al. 2007).
The total index of nesting female abundance of the SE Atlantic DPS
(9,198 females) does not reduce the risk for environmental variation,
genetic complications, demographic stochasticity, negative ecological
feedback, and catastrophes (McElhany et al. 2000; NMFS 2017). Such
abundance provides little resilience to buffer losses of individuals.
We conclude that the nesting female abundance, as estimated, does not
reduce the extinction risk of this DPS.
Productivity
Based on data collected from the largest nesting aggregation (i.e.,
Gabon), the SE Atlantic DPS exhibits a declining nesting trend. Data
collected between the 2002/2003 and 2015/2016 nesting seasons (with two
years of missing data) indicated a median trend in nesting activity of
-8.6 percent annually (sd = 21.9 percent; 95 percent CI = -52.6 to 36.9
percent; f = 0.676; mean annual nesting activities = 35,204). The trend
in Gabon is likely representative of the entire DPS, because the
majority of nesting occurs there. Additional nest trend data are
available from the Gran Caldera Scientific Reserve of Bioko Island,
where the number of body pits increased 2.8 percent annually (sd = 15.6
percent; 95 percent CI = -27.2 to 36.0 percent) from 1996/1997 to 2017/
2018.
Regarding productivity parameters, available information is often
from a limited area and may not be representative of the entire DPS.
However, based on available data, the size of nesting females, clutch
size, hatching success, and incubation period appear to be similar to
the species' averages. We conclude that the declining nesting trend
contributes to the extinction risk of this DPS.
Spatial Distribution
The SE Atlantic DPS has a broad spatial distribution. The nesting
range is centered on Gabon, with nesting occurring from Senegal to
Angola. Genetic data available for Gabon and Ghana indicate significant
genetic differentiation based on mtDNA data, but weak differentiation
based on analysis of nuclear DNA, likely indicating demographically
independent subpopulations connected by limited gene flow (Dutton et
al. 2013).
In addition to the extensive nesting range, this DPS also has an
expansive foraging and migratory range, from the coastal waters of
Atlantic Africa, across the pelagic waters of the South Atlantic, and
along the South American coast from Brazil to Argentina. While nesting
along the coast of Africa extends only to Angola, recent tag returns
and satellite telemetry indicate that turtles utilize the waters in
Namibia as well (Almeida et al. 2014). Transatlantic movements were
first recorded from tag returns of four leatherback turtles tagged on
the nesting beaches of Gabon and recaptured in the waters of Argentina
and Brazil (Billes et al. 2006). Satellite telemetry confirmed that
nesting females from Gabon follow three different post-nesting movement
trajectories towards the equatorial Atlantic Ocean, South America, or
southern Africa (Witt et al. 2011). For combined foraging areas off
Argentina and Eleva[ccedil][atilde]o do Rio Grande (an elevated
offshore area across from Brazil), the mean estimate from western
Africa was 84 to 86 percent (45 percent Gabon, 41 percent Ghana;
Prosdocimi et al. 2014).
The wide distribution of foraging areas likely buffers the DPS
against local catastrophes or environmental changes that could limit
prey availability. The expansive nesting range may buffer the DPS from
acute environmental impacts (e.g., storms and singular events) and to
some degree, chronic impacts (e.g., sea level rise and temperature
changes). Thus, the combination of extensive nesting range, widely
distributed foraging areas, and population structure reduces the
extinction risk of the SE Atlantic DPS.
Diversity
Genetic analyses for the SE Atlantic DPS are limited, but Dutton et
al. (2013) found moderate genetic diversity in samples from Gabon and
Ghana, including four new haplotypes unique to western African nesting
females. Nesting occurs on continental and insular beaches. There are
multiple foraging strategies, including pelagic and coastal, along
either side of the Atlantic Ocean. The genetic diversity, along with
multiple and diverse foraging sites (i.e., coastal and pelagic), and
combination of insular and mainland nesting provide diversity and
resilience that may reduce the extinction risk of this DPS.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
Modification and loss of habitat is a threat to the SE Atlantic
DPS. Present threats include obstructions, erosion, and light pollution
at nesting beaches. Future threats include coastal construction and
development in the region.
Nesting beach obstruction due to logs is a problem in Gabon,
Equatorial Guinea, and Cameroon (Formia et al. 2003). Logs that have
broken loose from timber rafts of industrial logging operations wash up
on the beaches of Gabon at densities of up to 247 logs/km; logs blocked
30.5 percent of the beach in Pongara, Gabon, resulting in an estimated
2,111 disrupted or aborted nesting attempts (Laurance et al. 2008). In
addition, several leatherback turtles have died as result of being
trapped by logs (Laurance et al. 2008). Pikesley et al. (2013)
determined that between 1.6 percent and 4.4 percent of nesting females
could be trapped at beaches with high log- and turtle-densities.
However, Gabon has since banned the export of whole logs. The Gabon Sea
Turtle Partnership has carried out log removal efforts for at least one
high-density nesting beach in Pongara National Park (Kingere Beach),
and a 3 km stretch of nesting beach is now virtually free of logs; at
the other main monitored beaches in Gabon, such as Mayumba and Gamba,
logs are not a major threat (A. Formia, WCS, pers. comm. 2019).
Habitat loss from coastal erosion due to sand mining, harbor
building, and irregular current flows has compromised the suitability
of long stretches of coastal areas as nesting sites. This issue is
especially prevalent between Ghana and Nigeria (Formia et al. 2003).
Ikaran (2010) found low hatching/emergence success rates at three
nesting sites in Gabon: Pointe Denis (17/16 percent), Mayumba (43/40
percent), and Kingere (16/16 percent).In addition to predation, the
main identified sources of egg mortality were beach erosion and
inundation (Ikaran 2010).
Light pollution modifies nesting beach habitat, deterring nesting
females
[[Page 48370]]
and disorienting both hatchlings and nesting females. Bourgeois (2009)
found that artificial lighting disoriented leatherback hatchlings in
Pongara National Park, Gabon: Hatchlings in 27 of the 41 nests (66
percent) studied crawled towards artificial lights. Deem et al. (2007)
documented 71 disoriented females that crawled directly into the
savannah behind the beach and towards the artificial lights. Bourgeois
et al. (2009) concluded that light pollution from Libreville and Pointe
Denis, Gabon is a major threat to nesting females and hatchlings, which
become disoriented and die in the surrounding savannah.
Urbanization and coastal development are rapidly growing threats at
some nesting beaches (Girard and Honarvar 2017). There is a high
potential for coastal development in Gabon, including the beaches near
Pointe Denis, an important and growing tourist area (Ikaran 2010).
Along with direct habitat loss from coastal development and
urbanization, impacts from pollution and litter are expected to
increase.
In Gabon, a network of marine protected areas was created by decree
00161/PR in 2017, covering 26 percent of Gabon's territorial seas,
including a vast area in front of the most important nesting beach in
Gabon (Mayumba National Park) that stretches to the outer limits of the
EEZ.
We conclude that a large portion of nesting females, hatchlings,
and eggs are exposed to the reduction and modification of nesting
habitat, as a result of logging, erosion, coastal development, and
artificial lighting. These threats impact the DPS by reducing nesting
and hatching success, thus lowering the productivity of the DPS.
Logging also results in the death of nesting females, reducing the
abundance of the population by removing its most reproductively
important individuals. Based on the information presented above, we
conclude that habitat loss and modification are major and increasing
threats to the DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization is a threat to the SE Atlantic DPS. Although
receiving some legal protections, eggs and turtles nevertheless are
poached for consumption, traditional medicine, and religious practices.
In Gabon, poaching is limited because 78 percent of nesting occurs
within national parks and human population density along the coast is
low (A. Formia, Gabon Sea Turtle Partnership, pers. comm., 2018).
However, elsewhere in the region, poaching occurs at a high rate, or
would be reasonably expected to return to high levels, if not limited
by activities funded through the USFWS' Marine Turtle Conservation Fund
enacted under the MTCA. These activities reduce poaching through
increased presence on nesting beaches, beach monitoring, hiring of
local citizens for participation in the projects, and raising awareness
and providing education to local communities (M. Tiwari, NMFS, pers.
comm. 2018).
Conflicting beliefs about sea turtles exist throughout the region.
In some communities sea turtles are considered divinely provided food,
while in others they have been historically protected by indigenous
custom, often based on stories passed down by ancestors (Barbosa and
Regalla 2016; Alexander et al. 2017). In general, however, poaching is
a significant problem throughout the region. Catry et al. (2009)
concluded that, in addition to fisheries bycatch, poaching of eggs and
nesting females is the main threat to sea turtles, including
leatherback turtles, in Guinea-Bissau. In many cases ``few if any
turtles or nests are left alone when found by locals'' (Catry et al.
2009). The fat of leatherback turtles is often used for various
purported medicinal applications, including: Treatment of convulsions
and malaria (Togo), fever, fainting spells, liver problems, tetanus
(Benin), and to induce vomiting (Togo, Benin). In one community in the
Ivory Coast and parts of Cameroon, leatherback turtle fat is applied to
wounds in the mouth and is used to massage into painful joints. In
northwestern and southern Cameroon, it is applied to bruises (Fretey et
al. 1999). In Togo, some mothers add turtle bones daily to the baby's
bath water; some believe that the power of the turtle (especially the
leatherback) will be transmitted to the child through this practice
(Segniagbeto 2004).
Turtles and eggs are poached throughout the nesting range of the
DPS. Though most nesting females and eggs are protected in Gabon,
poaching is widespread in other areas. Poaching of nesting females
reduces both abundance (through loss of nesting females) and
productivity (through loss of reproductive potential). Such impacts are
high because they directly remove the most productive individuals from
DPS, reducing current and/or future reproductive potential. Egg
poaching reduces productivity. Given the moderate exposure and high
impact, we conclude that the poaching of turtles and eggs poses a
threat to the DPS.
Disease or Predation
Information on diseases among leatherback turtles originating in
the SE Atlantic is minimal, but an analysis of samples from nesting
females in Gabon indicated normal blood chemistry parameters (Deem et
al. 2006). Predation may occur at high rates in some areas, but
information is limited.
Predation of leatherback eggs and/or hatchlings has been documented
for a variety of predators, including: Various ants, ghost crabs,
monitor lizards (Varanus niloticius), crows (Corvus albus), mongoose,
porcupine (Atherurus africanus), domestic dogs, African civet cat
(Civettictis civetta and Viverra civetta), and drills (Mandrillus
leucophaeus) (summarized from Eckert et al. 2012). In Kingere, Gabon,
Ikaran (2010) noted high predation rates of eggs by crabs, lizards,
mongooses, small cat species, and ants. Predation was the main source
of egg mortality at three nesting sites in Gabon: Pointe Denis (43
percent), Mayumba (44 percent), and Kingere (51 to 56 percent; Ikaran
2010).
As is common for all sea turtle species, leatherback hatchlings
likely experience predation from various fish species as they enter the
water and swim towards the open ocean. In-water predation of juveniles
and adults is not well-documented, but there is evidence of shark and
killer whale predation. Shark predation was determined to be the cause
of one leatherback stranding reported from Central Africa (Parnell et
al. 2007), while interactions between killer whales and leatherback
turtles resulting in possible predation has been observed in Namibian
waters (Elwen and Leeney 2011).
While all eggs and hatchlings have some exposure to predation, the
species compensates for a certain level of natural predation by
producing a large number of eggs and hatchlings. For this DPS, the
primary impact is to productivity (i.e., reduced egg and hatching
success). We conclude that predation poses a threat to the SE Atlantic
DPS.
Inadequacy of Existing Regulatory Mechanisms
The SE Atlantic DPS is protected by various regulatory mechanisms.
For each, the Team reviewed the objectives of the regulation and to
what extent it adequately addresses the targeted threat.
The harvest of turtles and eggs is illegal in most of the nations
where the DPS nests. In some cases, however, these protective
mechanisms are inadequate. In addition, many nesting beaches are not
protected.
In Gabon, the harvest of turtles and eggs is illegal (2011 decree
0164/PR/
[[Page 48371]]
MEF) and much of the nesting beach habitat (and turtles utilizing that
habitat) is protected because of inclusion in parks as well as being
far from any city or town. However, low levels of poaching occurs, and
the threats from encroaching development and associated impacts are
increasing.
In Congo, wildlife laws prohibit the hunting and collection of
wildlife and their products, including eggs, between November 1 and
April 31. Turtles are also protected in the Conkaouati-Douli National
Park. However, in areas without permanent beach monitoring, almost all
eggs and nesting individuals are collected and eaten (Bal et al. 2007).
In the Democratic Republic of Congo, leatherback turtles are cited
under the 1982 Hunting Act for protection. However, there is no post-
independence legislation protecting sea turtles, and there is little
commitment to the legislated protections (Fretey 2001).
Since 1988, Equatorial Guinea has protected all sea turtles under
Law 8/1988 and Decree 183/87 on fishing (Tom[aacute]s et al. 2010).
However, the poaching of eggs and females for local consumption and
sale has occurred (Castroviejo et al. 1994).
In Ghana, the Wildlife Regulations Act of 1974 prohibits all
harvest of eggs and turtles. However, poverty is prevalent, and eggs
and sea turtles are poached at nesting beaches (Tanner 2013).
Enforcement is likely inadequate because of funding issues, the
remoteness of some nesting beaches, and cultural practices.
Fishery bycatch is the primary threat to this DPS. While most
nations in the region have some form of legal protection for sea
turtles, many leatherback turtles die from fisheries bycatch throughout
the range of the DPS. Examples of fisheries legislation include
Brazil's gear restrictions and Nigeria's requirement to use TEDs in
bottom trawls.
In summary, numerous regulatory mechanisms provide some protection
to leatherback turtles, their eggs, and nesting habitat throughout the
range of this DPS. Though the regulatory mechanisms provide some
protection to the turtles, many do not adequately reduce the threat
that they were designed to address, generally as a result of limited
implementation or enforcement. Fisheries bycatch, poaching, and habitat
loss remain major threats to the DPS despite regulatory mechanisms. We
conclude that inadequacy of the regulatory mechanisms are a threat to
the SE Atlantic DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat to the SE Atlantic DPS.
Leatherback turtles are captured as bycatch in commercial and artisanal
fisheries along coastal foraging and breeding areas as well as on the
high seas. Because of the overlapping range with the SW Atlantic DPS,
this DPS is vulnerable to interactions with fisheries off the coasts of
Brazil, Uruguay, and Argentina, in the pelagic waters of the South
Atlantic Ocean, and along the coastal waters off western Africa.
Therefore, the information presented on the fisheries bycatch for the
SW Atlantic is applicable to this DPS.
One of the biggest threats for leatherback turtles in Atlantic
waters is bycatch in artisanal and commercial fisheries (Wallace et al.
2010; Riskas and Tiwari 2013;). Lewison et al. (2004) estimated that
30,000 to 60,000 leatherback turtles were taken as longline fisheries
bycatch in the entire Atlantic Ocean in 2000. Stewart et al. (2010)
estimated that in West Africa, Benin, Togo, and Cameroon had the
highest average fishing densities, ranging from 11.1 to 6.5 boat-
meters/km\2\, and gillnet densities ranked among the highest on a
global scale. Despite very active artisanal and industrial fisheries in
the region, overall bycatch data are quite sparse or qualitative
(rather than quantitative) in nature, and Africa still represents a
significant gap in bycatch evaluation studies (Wallace et al. 2010,
2013). Accurate and reliable bycatch data are difficult to achieve, as
direct observation rates are low (<1 percent of total fleets) and
statistics from the region's many small-scale fisheries are largely
incomplete (Kelleher 2005; Moore et al. 2010; Wallace et al. 2010).
However, several studies have concluded that bycatch rates in the
region are high, given the degree of fishing activity near nesting and
foraging areas (Lewison et al. 2004; Moore et al. 2010; Wallace et al.
2010).
Along the coasts of Angola, Namibia, and South Africa, Honig et al.
(2007) evaluated turtle bycatch by longline fisheries in the Benguela
Large Marine Ecosystem by using data from observer reports, surveys,
and specialized trips from the coastal nations of South Africa, Namibia
and Angola. They estimated bycatch at 672 leatherback turtles annually
(based on an annual bycatch estimate of 4,200 turtles, of which
approximately 16 percent are leatherback turtles) in the southern and
central regions and as many as 5,600 leatherback turtles (based on an
annual bycatch estimate of 35,000 turtles) for the entire Benguela
Large Marine Ecosystem (Honig et al. 2007). Mortality rates were not
provided in this study but may range from 25 to 75 percent (Aguilar et
al. 1995). The estimates mostly include turtles from the SE Atlantic
DPS, but telemetry studies indicate that the turtles of the much
smaller SW Indian DPS also use this foraging area (Luschi et al. 2006;
Robinson et al. 2016). Evaluating ICCAT data, Angel et al. (2014)
confirm exposure to high longline fishing effort and some purse seine
effort for the population originating from the SE Atlantic Ocean.
The limited bycatch data available for waters of the western coast
of Africa show that other fisheries interact with leatherback turtles.
Between 2005 and 2015, artisanal fishing nets in Loango Bay in the
Republic of Congo killed a total of 45 leatherback turtles; 0 to 628
leatherback turtles were captured or recaptured annually over that time
period (Br[eacute]heret et al. 2017). An assessment of bycatch in the
trawling fisheries in Gabon found that leatherback turtles represented
only 2 percent of the bycatch despite being the most abundant sea
turtle species in Gabonese waters; the low rate is possibly because
leatherback turtles do not occur in the section of the water column
where the trawl net is towed (Casale et al. 2017). Trawl bycatch in the
waters around S[atilde]o Tom[eacute] and Principe included 4 juvenile
leatherback turtles (17 to 21 cm in carapace length) in March 1994
(Fretey et al. 1999).
While specific information to estimate overall capture and
mortality rates of SE Atlantic leatherback turtles in fisheries is not
available, it is clear that bycatch in fisheries, especially gillnets
and longlines, are a threat to the DPS across its range. Immature and
mature individuals are exposed to high fishing effort throughout their
foraging range and in coastal waters near nesting beaches. Mortality is
also high. Mortality reduces abundance, by removing individuals from
the population; it also reduces productivity, when nesting females are
incidentally captured and killed. We conclude that fisheries bycatch is
a major, and the primary, threat to the SE Atlantic DPS.
Vessel Strikes
There is little information regarding vessel strikes for the SE
Atlantic DPS, but such interactions are a potential, and possibly
increasing, threat across at least a portion of this DPS's range. In
the western South Atlantic foraging grounds off Brazil, Uruguay, and
Argentina, increasing vessel traffic from fishing vessels, cargo
transport, and tourism has been noted (L[oacute]pez-Mendilaharsu et al.
[[Page 48372]]
2009; Fossette et al. 2014), potentially increasing the likelihood of
vessel strikes on leatherback turtles. Although no specific information
is available for the waters off western Africa, any economic
development along the coast is likely to result in an increase in
vessel traffic. We conclude that vessel strikes are a threat to the SE
Atlantic DPS.
Pollution
The SE Atlantic DPS faces the threat of pollution across its
extensive range throughout the South Atlantic Ocean, from Africa to
South America. As the ranges of the SW Atlantic and SE Atlantic DPSs
overlap, they are exposed to the same pollutants, which include
contaminants, marine debris, and ghost fishing gear. Throughout Africa,
marine and coastal pollution is widespread in industrial and urban
areas, and garbage litters many developed beaches (Formia et al. 2003;
Agyekumhene et al. 2017). Off the coast of South America, the Argentine
and Brazilian coastal waters are increasingly impacted by economic
activities, such as maritime cargo transport, tourism, and the
discharge of domestic and industrial waste (L[oacute]pez-Mendilaharsu
et al. 2009; Fossette et al. 2014).
The Gulf of Guinea has increasingly been the focus of extensive oil
exploitation activities, following the discovery of large oil reserves.
Drilling activities by large oil corporations, with associated
pollution and habitat destruction, are threats to nesting aggregations
in the area (Formia et al. 2003; Agyekumhene et al. 2017). In 2012/
2013, oil spills following the dredging of the Port of Pointe-Noire in
the Republic of Congo significantly degraded the fauna and flora of
Loango Bay, where leatherback turtles occur. However, the ecosystem is
believed to be slowly recovering (Br[eacute]heret et al. 2017). In
2005, a moderate slick of oil on the beaches of Mayumba National Park
in Gabon was observed, although its impacts on turtles are unknown
(Parnell et al. 2007).
In Nigeria, the main sources of pollution include industrial waste,
raw/untreated sewage, and pesticides. Oil exploration, exploitation,
and transportation have a significant effect on the environment. Spills
of crude and refined oil are frequent in the coastal and marine
environment, especially during periods of very strong ocean currents,
when they can spread to cover the entire 853 km coastline of Nigeria.
It is clear that individuals from the SE Atlantic DPS have a high
probability of encountering pollution across their range and throughout
their lifecycle. Although the best available information does not
quantify such impacts, ample information demonstrates that these
threats are ongoing. We conclude that pollution is a threat to the DPS.
Climate Change
Climate change is a threat to the SE Atlantic DPS. The impacts of
climate change include: Increases in temperatures (air, sand, and sea
surface); sea level rise; increased coastal erosion; more frequent and
intense storm events; and changes in ocean currents.
Sea level rise resulting from climate change negatively impacts sea
turtle nesting. Erosion of important nesting beaches in Gabon may be at
least partially attributable to sea level rise. From 1983 through the
2000s, some areas have lost up to 100 m of beach width, reducing the
availability of suitable nesting beach (Gabon Sea Turtle Partnership
2018; https://www.seaturtle.org/groups/gabon/erosion.html). Because
leatherback turtles nest lower on the beach than other sea turtles,
their eggs are more at risk of being inundated and destroyed by
increases in sea level and coastal erosion (Boyes et al. 2010).
Changes in sand temperatures are likely to impact egg viability and
sex determination. Ikaran (2010) found the thermal range of sand over
the nesting season to be adequate for hatchling sex ratios to be mixed
or even male dominated. In Gabon, the early rainy months tend to
produce males, while the later, warmer months produce females, with a
tendency towards a net higher production of males. Ikaran (2010)
considered the nesting beaches of Gabon to be an important male
producing area. However, based on predictions of warming trends, he
found that within two decades the ratio could skew towards 100 percent
female.
The threat of climate change is likely to modify the nesting
conditions for turtles of the DPS, and it is unclear whether they have
or can develop the ability to nest in different locations along
existing beaches, or on new beaches. Impacts from climate change are
likely to range from small, temporal changes in nesting season to large
losses of productivity. Therefore, we conclude that climate change is a
threat to the DPS.
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply within the range of the SE
Atlantic DPS (for a description of each effort, please see the section
on conservation efforts for the overall species): Convention on the
Conservation of Migratory Species of Wild Animals, Convention on
Biological Diversity, Convention on International Trade in Endangered
Species of Wild Fauna and Flora, Convention Concerning the Protection
of the World Cultural and Natural Heritage (World Heritage Convention),
FAO Technical Consultation on Sea Turtle-Fishery Interactions, IAC,
MARPOL, IUCN, Memorandum of Understanding Concerning Conservation
Measures for Marine Turtles of the Atlantic Coast of Africa, Ramsar
Convention on Wetlands, South-East Atlantic Fisheries Organization,
UNCLOS, and UN Resolution 44/225 on Large-Scale Pelagic Driftnet
Fishing. Although numerous conservation efforts apply to the turtles of
this DPS, they do not adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
overall that the SE Atlantic DPS is at high risk of extinction. The
total index of nesting female abundance is 9,198 females. Since 2002,
the first year that aerial survey data was collected, nesting activity
has declined by -8.6 percent annually in Gabon, the largest nesting
aggregation of the DPS, and what was, in 2002, the largest nesting
aggregation in the world. This declining trend has the potential to
further lower abundance and increase the risk of extinction. Nesting
and foraging is broadly distributed; thus, the population is somewhat
buffered from stochastic events that could otherwise have catastrophic
effects on the entire DPS. There is a metapopulation structure within
this DPS, with fine-scale genetic differentiation between Gabon and
Ghana. Genetic diversity also appears to be moderate. Based on the
reduced nesting female abundance and declining nest trend, we find the
DPS to be at risk of extinction, likely as a result of past threats.
Current threats place the DPS at further risk of extinction. The
primary threat to this DPS is bycatch in commercial and artisanal,
pelagic and coastal, fisheries, especially coastal gillnet and pelagic
longline fisheries. Fisheries bycatch reduces abundance by removing
individuals from the population. Because several fisheries operate near
nesting beaches, productivity is also reduced when nesting females are
prevented from returning to nesting beaches. Thus, exposure and impact
of this threat are high. Habitat loss or modification is a threat that
reduces abundance and productivity and includes the impacts of logs,
which block access to the
[[Page 48373]]
beaches or trap nesting females and hatchlings. Poaching of turtles and
eggs is also a threat to this DPS, although most nesting beaches in
Gabon are somewhat protected because they occur in parks or are far
from any towns. Many of the beaches outside Gabon (e.g., Guinea-Bissau)
have limited or no protection. The degree of overutilization is highly
varied across locations, but quite extensive in some areas. Funding
from the MTCA has resulted in some reduction of this threat as
conservation activities, research, and community involvement results in
lower poaching on those beaches. However, poaching continues at high
levels in other areas. Additional threats include: predation and
disease, inadequate regulatory mechanisms, pollution, and climate
change. Predation can be extensive at some specific beaches, but
overall it does not occur at a high level. Pollution is a persistent
and potentially increasing threat. Ingestion of plastics and
entanglement in marine debris result in injury and reduced health, and
sometimes mortality. Climate change is likely to result in reduced
productivity due to greater rates of coastal erosion and nest
inundation, and in some areas, nest failure or skewed sex ratios due to
increased sand temperatures. Vessel strikes are a threat that is likely
to increase over time as recreational and commercial vessel activity
increases, resulting in more opportunity for interactions. Though many
regulatory mechanisms are in place, they do not adequately reduce the
impact of logs, poaching, and fisheries. Additionally, many areas in
the region have little or no enforcement of laws protecting turtles or
nests on the beach.
The DPS is relatively data-poor, reducing our ability to quantify
threats for more than a small portion of the population. For this
reason, the Status Review Team did not come to consensus regarding the
extinction risk analysis for the SE Atlantic DPS. All Team members were
present to vote on the level of extinction risk. Nine Team members
concluded with moderate confidence that the DPS is at high extinction
risk due to threats and loss of abundance; their confidence was
moderate due to the lack of data on this DPS. Two team members
concluded with low confidence that the DPS is at moderate extinction
risk; their confidence in this conclusion is low due to the lack of
data on this DPS.
We conclude, consistent with the Team's overall conclusion, that
the SE Atlantic DPS is currently in danger of extinction. The
decreasing nesting trend (i.e., 8.6 percent annually since 2002) is at
or near a level that make the DPS highly vulnerable to threats, given
the total index of nesting female abundance of 9,198 females. It faces
present, ongoing threats that are likely to create imminent and
substantial demographic risks (i.e., declining trends and reduced
abundance). Though numerous conservation efforts apply within the range
of this DPS, they do not adequately reduce the risk of extinction. We
conclude that the SE Atlantic DPS is currently in danger of extinction
throughout its range and therefore meets the definition of an
endangered species. The threatened species definition does not apply
because the DPS is at risk of extinction currently (i.e., at present),
rather than on a trajectory to become so in the foreseeable future.
SW Indian DPS
The Team defined the SW Indian DPS as leatherback turtles
originating from the SW Indian Ocean, north of 47[deg] S, east of
20[deg] E, and west of 61.577[deg] E. The western boundary occurs at
the southern tip of Africa, approximately where the Agulhas and
Benguela Currents meet. The eastern boundary occurs at the border
between Iran and Pakistan, where the Somali Current begins. These
currents, and the cold waters of the Antarctic Circumpolar Current,
likely restrict the nesting range of this DPS.
The range of the DPS (i.e., all documented areas of occurrence)
extends into the SE Atlantic Ocean, where leatherback turtles forage in
the highly productive Benguela Current Large Marine Ecosystem, which
occurs along the western coast of Africa, from Angola to South Africa.
Leatherback turtles also range throughout the waters of eastern Africa
(Ross 1985) and possibly into the Red Sea (Gasparetti et al. 1993).
Records indicate that the species has been observed in the waters of
the following nations: Djibouti; Eritrea; French Territories (Reunion
Island, Mayotte, and Iles Eparses); Kenya; Madagascar; Mozambique;
Seychelles; Somalia; South Africa; Tanzania; and Yemen (Hamann et al.
2006). Leatherback turtles may occur in the waters of the following
nations: Bahrain, Kuwait; United Arab Emirates; Oman; and Sudan (Hamann
et al. 2006).
Leatherback turtles of the SW Indian DPS nest over a distance of
approximately 900 km, from Cape Vidal, South Africa to Bazaruto
Islands, Mozambique (Videira et al. 2011; Nel et al. 2015). The vast
majority of nesting (80 to 90 percent) occurs in South Africa, between
Bhanga Nek and Leifeld's Rock (Nel et al. 2015). In Mozambique, most
nesting occurs from the southern border to Inhaca Island, Mozambique,
with low levels of nesting farther north at Bilene Beach and Bazaruto
Islands (Nel et al. 2015). This DPS nests at the highest latitude (and
southernmost location) of all leatherback turtles (Saba et al. 2015).
Nesting occurs on long (5 to 15 km), broad (50 to 100 m), silica
sand beaches with little vegetation (Botha 2010; Nel et al. 2015;
Robinson et al. 2017). The beaches are characterized by pristine,
intact dunes that rise up to 100 m above sea level, interspersed with a
few dynamic dunes and small, primary dunes (Nel et al. 2015). The
beaches are separated by short rocky headlands (Robinson et al. 2017).
Subtidal rock formations are dispersed throughout the high energy
coastline. Nesting females approach the beach using strong rip-currents
through obstruction-free areas (Hughes 1974; Hughes 1996; Botha 2010;
Nel et al. 2015).
Foraging areas of the SW Indian DPS include coastal and pelagic
waters of the SW Indian Ocean and the SE Atlantic Ocean. The DPS is
somewhat unique in that turtles forage in two ocean basins and do not
need to undergo long migrations between nesting and foraging areas
because highly productive foraging areas are available adjacent to
nesting beaches or connected to nesting beaches via fast-moving
currents. For example, the warm, fast-flowing Agulhas Current
(Lutjeharms 2001; Nel et al. 2015) results in high productivity
foraging areas near nesting beaches and provides a migratory corridor
to distant foraging areas. As a result, the SW Indian turtles have the
largest body size, largest clutch size, and highest reproductive output
of all leatherback turtles (Saba et al. 2015).
Satellite tracking of post-nesting females (n = 27) reveals the use
of one of three post-nesting migratory corridors: north into the nearby
coastal waters of the Mozambique channel; south and west (via the
Agulhas and Benguela Currents) into the pelagic waters of the South
Atlantic Ocean; or south and east (via the Agulhas Current and
Retroflection) into the oceanic eddies in the SW Indian Ocean (Luschi
et al. 2006; Robinson et al. 2016; Harris et al. 2018). Luschi et al.
(2006) reviewed satellite telemetry data of 11 post-nesting females
tagged between 1996 and 2003 (Hughes et al. 1998; Luschi et al. 2003;
Sale et al. 2006); and Robinson et al. (2016) satellite tracked 16
post-nesting females tagged between 2011 and 2013. Evaluating tracking
data for 14 post-nesting females between 2006 and 2014, Harris et al.
(2018) found that leatherback turtles equally used all three migration
corridors. In the other studies, a total of 11 post-nesting
[[Page 48374]]
females migrated a relatively short distance (approximately 500 km) to
the shallow (less than 50 m depth), coastal waters of the Sofala Banks
(i.e., the Mozambique Channel), where net primary productivity and sea
surface temperatures remain elevated year-round (n = 4, Sale et al.
2006; n = 7, Robinson et al. 2016). One post-nesting female migrated to
the similarly hospitable coastal waters of Madagascar (Robinson et al.
2016). Ten post-nesting females tracked to pelagic waters of the
Atlantic Ocean (n = 6, Sale et al. 2006; n = 4, Robinson et al. 2016).
These waters are among the most productive in the world, as a result of
strong upwelling (caused by the southeast trade winds) and the area's
unique bathymetry, hydrography, chemistry, and trophodynamics (Honig et
al. 2007). Five post-nesting females appeared to track oceanic eddies
into the SW Indian Ocean (n = 1, Sale et al. 2006; n = 4, Robinson et
al. 2016). Luschi et al. (2003 and 2006) characterized leatherback
turtles using this latter strategy as ``wanderers, ranging over vast
oceanic areas while searching for their planktonic prey.''
Opportunistically encountered and highly productive eddies likely
shaped the circuitous routes of these foraging turtles, which resemble
drifters more than active swimmers (Luschi et al. 2006; Robinson et al.
2016; Harris et al. 2018). Thus, this DPS benefits from the use of
three migratory corridors that all provide highly productive foraging
opportunities, with minimal energetic cost required to return to waters
off nesting beaches.
Abundance
The total index of nesting female abundance of the SW Indian DPS is
149 females. We based this index on two nesting aggregations: South
Africa (Ezemvelo KwaZulu-Natal Wildlife (Ezemvelo), unpublished data,
2018) and Mozambique (Centro Terra Viva Estudos e Advocacia Ambiental
(CTV), unpublished data, 2018). Our total index does not include two
unquantified nesting aggregations in Mozambique. To calculate the index
of nesting female abundance (i.e., 134 females) for the South Africa
``monitoring area'' (i.e., a 52.8 km stretch of beach that has been
monitored for decades), we divided the total number of nests between
the 2014/2015 and 2016/2017 nesting seasons (i.e., a 3-year remigration
interval; Hughes 1996; Lambardi et al. 2008; Nel et al. 2013; Saba et
al. 2015) by the clutch frequency (7 clutches/season; Nel et al. 2013;
Saba et al. 2015). To calculate the index of nesting female abundance
in Mozambique (i.e., 15 females), we divided the total number of nests
between the 2015/2016 and 2017/2018 nesting seasons (i.e., a 3-year
remigration interval) by the clutch frequency for South Africa (7
clutches/season; Nel et al. 2013; Saba et al. 2015).
This is an index for the DPS because it only includes available
data from recently and consistently monitored nesting beaches. While
nesting occurs on beaches that stretch across 900 km of South Africa
and Mozambique, consistent and standardized monitoring occurs only
across approximately 300 km of beaches across the two nations (Nel et
al. 2013; Nel et al. 2015). Furthermore, while nesting is known to
occur at low levels at Inhaca Island and Bazaruto Archipelago in
Mozambique, we did not include these sites because we did not have data
from the most recent 3 years.
Other estimates of total or annual nesting female abundance have
been published. The IUCN Red List assessment estimated the total number
of mature individuals (males and females) at 148 individuals, based on
an average of 259 annual nests (Nel et al. 2013), a 3-year remigration
interval (Nel et al. 2013), and a 3:1 sex ratio (Wallace et al. 2013).
Their estimates are based on nesting surveys conducted in South Africa,
which hosts approximately 80 to 90 percent of nesting, and Mozambique
(Wallace et al. 2013; Nel et al. 2015). Their estimate is less than our
index, despite including mature males and females. The reason for this
difference is because they used an average annual number of nests that
was lower than recent nest counts over the 3-year remigration interval.
Nel et al. (2015) estimated the size of the total nesting population at
approximately 100 females per season (Nel et al. 2015), based on 2010
data: 375 emergences and 336 nests in South Africa; and 61 emergences
in Mozambique (Videira et al. 2011). This estimate (n = 300, based on a
3 year remigration interval) is greater than our index because there
were more nests in 2010 compared to more recent years (2014 to 2016).
Hamann et al. (2006) estimated approximately 20 to 40 nesting females
annually in South Africa and approximately 10 nesting females annually
in southern Mozambique. This estimate (n = 90 to 150, based on a 3 year
remigration interval) is less than our index, likely as a result of
using data collected over a different time-frame. The difference in
estimates likely results from using different methods of calculation
and different time frames and reflects some uncertainty in the precise
number of nesting females. Our total index of nesting female abundance
falls within the range of other estimates and is based on the best
available data for the DPS at this time.
There are additional published estimates for the South Africa
monitoring area. Nel et al. (2013) identified 2,578 nesting females
over 45 years (1965 to 2009), with a mean of 69.4 38.1
nesting females per season (or 209 total nesting females) in the
monitoring area. Hughes (1996) reported an annual average of 24 nesting
females in the first decade (1976 to 1985) and an annual average of 86
nesting females in the second decade (1986 to 1995) in the monitoring
area. Hughes (1996) also reported an annual average of 113 nesting
females from 1986 to 1995 in an extended protected area that includes
the monitoring area plus another 93 km in the St. Lucia Marine Reserve,
which is surveyed periodically. The difference between these two
averages reflects that most estimates of nesting female abundance in
South Africa are minimum estimates because nesting occurs outside the
monitoring area. Thorson et al. (2012) found that annual resightings
for leatherback turtles decreased from the 1960s to 2009, and their
modeling indicated that this decline was due to decreased detection
probabilities (i.e., decreased probability of returning to the
monitored portion of the KwaZulu-Natal nesting beach), rather than
decreased survival. Based on satellite tracking of 17 post-nesting
females, Harris et al. (2015) estimates that approximately 66 percent
of leatherback nesting activity occurs outside the monitoring area.
However, considerable inter-annual variability exists, ranging from
less than 30 percent to over 80 percent, with a median of approximately
49 percent (Harris et al. 2015). Thus, incomplete beach monitoring is a
source of uncertainty for this DPS and for our total index of nesting
female abundance.
For Mozambique, our index of nesting females is similar to other
published estimates, which are generally less than 20 nesting females
(Hamann et al. 2006; Louro 2014; Pereira et al. 2014; Fernandes et al.
2018). If we use the clutch frequency for Ponta Malongane (2.25
clutches per season; Louro et al. 2006), which is low for the species,
our index of nesting female abundance is 45 females. This clutch
frequency may be underestimated due to females nesting in distant areas
where monitoring does not regularly occur. If we use the clutch
frequency for South Africa, (7 clutches/season; Nel et al. 2013; Saba
et al. 2015), the resulting index of nesting female abundance for
Mozambique (i.e., 15
[[Page 48375]]
nesting females) is closer to published estimates.
The total index of nesting female abundance of 149 females places
the DPS at risk for environmental variation, genetic complications,
demographic stochasticity, negative ecological feedback, and
catastrophes (McElhany et al. 2000; NMFS 2017). These processes,
working alone or in concert, place small populations at a greater
extinction risk than large populations, which are better able to absorb
losses in individuals. Due to its small size, the DPS has restricted
capacity to buffer such losses. Given the intrinsic problems of small
population size, we conclude that the limited nesting female abundance
is a major factor in the extinction risk of this DPS.
Productivity
The SW Indian DPS exhibits a slightly decreasing nesting trend. We
base our conclusion on data consistently collected in a standardized
approach in the 56 km South African monitoring area (Ezemvelo,
unpublished data, 2018), where nest counts decreased by -0.3 percent
annually (sd = 2.1 percent; 95 percent CI = -4.5 to 4.1 percent; f =
0.557; mean annual nests = 301) between the 1973/1974 and 2016/2017
nesting seasons. The trend in South Africa is likely representative of
the entire DPS, as 80 to 90 percent of nesting is estimated to occur
there (Wallace et al. 2013; Nel et al. 2015) and the 44-year time
series is quite robust.
Our trend estimates yield similar results to other published
findings for the population. The IUCN concluded that this population
has declined slightly, by 5.6 percent over the past three generations,
with an annual decline of -0.1 percent in South Africa and -0.7 percent
in Mozambique (Wallace et al. 2013). Hamann et al. (2006) also
identified a declining trend in the nesting population of the SW Indian
Ocean. Studies focused on the South African monitoring area (i.e., the
source of data for our trend analysis), however, disagree on the
whether the trend has declined recently (Hamann et al. 2006; Nel et al.
2013) or is stable (Nel et al. 2015; Saba et al. 2015). The nest trend
may be stable if nesting in unmonitored areas has increased over time
(Thorson et al. 2012; Harris et al. 2015). Different datasets lead to
different conclusions due to different methods of calculation,
different time frames, incomplete monitoring of all nesting areas, and
therefore uncertainty in the precise number of nesting females. We find
that Nel et al. (2013) provide the best available published data, which
are based on the most recent, primary data, and we agree with their
characterization of the trend as declining or recently declining.
Despite the recent decline in nesting, productivity parameters
remain relatively high for the SW Indian DPS, which has the largest
body size, largest clutch size, and highest reproductive output of all
leatherback turtles, likely due to the close proximity between their
nesting beaches and highly productive foraging areas (Saba et al.
2015). Nel et al. (2015) reports that most metrics (i.e., female size,
egg size, incubation time, and hatching success) are above average for
this DPS. Nesting females produced 1,171 to 53,139 hatchlings each
season in the South Africa monitoring area between 1965 and 2009, with
an average of 36,583 to 51,610 hatchlings per season, which was
calculated by multiplying 480 hatchlings per nesting female by 69.4
38.1 nesting females per season (Nel et al. 2013).
The recent nesting decline may reflect the effects of past and
current threats that overwhelm the population's high productivity
metrics. We conclude that the slightly declining nest trend places the
DPS at risk of extinction, which is further exacerbated by the limited
nesting female abundance.
Spatial Distribution
The SW Indian DPS comprises, in essence, a single nesting
aggregation, with nesting females moving freely between South African
and Mozambican beaches (Hughes 1996; Luschi et al. 2006; Nel et al.
2015). Nesting is limited to a total distance of approximately 900 km
along South African and Mozambican coasts (Nel et al. 2015). While 80
to 90 percent of nesting is concentrated in South Africa, nesting is
somewhat concentrated in the southern section of the South African
monitoring area, although most characterize nesting as low density
throughout South Africa (Hughes 1974; Lambardi et al. 2008; Botha 2010;
Nel et al. 2013; Harris et al. 2015; Nel et al. 2015).
The DPS exhibits a broad foraging range that extends into coastal
and pelagic waters of the eastern Atlantic and western Indian Oceans
(Luschi et al. 2006; Lambardi et al. 2008; Girondot 2015). There is
limited evidence that leatherback turtles may remain in South African
waters throughout the year, as suggested by year-round fisheries
bycatch records (Luschi et al. 2003, 2006; Petersen et al. 2009). Some
forage off the coast of Madagascar (Robinson et al. 2016; Harris et al.
2018). Some turtles follow the Agulhas and Benguela Currents into
foraging areas in the southeast Atlantic Ocean, off the coasts of
Angola and Namibia (Girondot 2015; Robinson et al. 2016; Harris et al.
2018). Others follow the Agulhas Retroflection and deep-sea eddies into
the SW Indian Ocean (Luschi et al. 2006; Lambardi et al. 2008; Robinson
et al. 2016; Harris et al. 2018). Leatherback turtles, possibly from
this DPS, have also been observed in the Red Sea, presumably foraging
(Hamann et al. 2006). The use of various foraging areas may be
influenced by the prevalent currents encountered off the nesting
beaches (Luschi et al. 2006; Lambardi et al. 2008; Robinson et al.
2016).
The wide distribution of foraging areas likely buffers the DPS
somewhat against local catastrophes or environmental changes that would
limit prey availability. Nesting occurs along one coastline, which is
3,000 km in length and may be similarly affected by environmental
variation and directional changes (e.g., sea level rise). Because the
DPS is essentially a single nesting aggregation, it has limited
capacity to withstand other catastrophic events. Thus, spatial
distribution likely has little net effect on the extinction risk of the
SW Indian DPS.
Diversity
Within the SW Indian DPS, genetic diversity is low, with only two
mtDNA haplotypes found in 41 nesting females in South Africa (haplotype
diversity = 0.298 0.078 and nucleotide diversity = 0.0004
0.0004; Dutton et al. 2013). Nesting habitat is mainly
restricted to beaches along the same coast, with a few nests on
Mozambican islands. The DPS does not exhibit temporal or seasonal
nesting diversity, with most nesting occurring between October and
March. The foraging strategies are diverse, however, with turtles using
coastal and pelagic waters in the Atlantic and Indian Oceans. Diverse
foraging strategies may provide some resilience against local
reductions in prey availability or catastrophic events, such as oil
spills, by limiting exposure. Low genetic diversity indicates the DPS
may lack the raw material necessary for adapting to long-term
environmental changes, such as cyclic or directional changes in ocean
environments due to natural and human causes (McElhany et al. 2000;
NMFS 2017). We conclude that limited overall diversity increases the
extinction risk of this DPS by reducing its resilience to threats.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
Coastal erosion, foot and vehicle traffic, and artificial lighting
modify the available, suitable nesting habitat and
[[Page 48376]]
thus are threats to the SW Indian DPS. Angel et al. (2014) identifies
coastal erosion as the main beach-based threat to this population and
one that is likely to increase with climate change.
Coastal erosion removes sand from nesting beaches, inundating nests
and destroying eggs. Because leatherback turtles nest lower on the
beach than other sea turtles, they have greater exposure to tidal
erosion and deposition (Boyes et al. 2010). At South African nesting
beaches over a duration of 70 days, Boyes et al. (2010) found an
average of 0.62 m deposition (S.D. 0.15 m; range 0.34-0.85 m) and 0.42
m erosion (S.D. 0.17 m; range 0.14- 0.71 m). Because the average depth
of leatherback nests was 0.66 m (S.D. 0.19 m; range 0.15-1.07 m), eggs
are at some risk of being exposed and destroyed (Boyes et al. 2010).
Nel et al. (2006) concludes that coastal erosion is a threat in South
Africa, where the high-energy coastline varies seasonally. During two
nesting seasons (2009/2010 and 2010/2011), de Wet (2012) found that 6.3
percent of nests in the South African monitoring area were destroyed by
erosion. In Bazaruto Archipelago, Mozambique, coastal erosion and
rising sea levels destroyed approximately 12 percent of nests over 10
seasons of monitoring (Videira and Louro 2005; Louro 2006). Despite
nest loss due to erosion, hatching success remains high in South Africa
(70 to 80 percent; Nel et al. 2015; Santidri[aacute]n Tomillo et al.
2015). Though the introduction of Casuarina trees do not necessarily
increase the risk of erosion, they obstruct nesting females' access to
and from beaches and alter nest incubation environments (de Vos et al.
2019). Evolving in a high-energy coastline environment with seasonal
variation has likely provided the DPS with some resilience to nesting
losses due to coastal erosion. Sea level rise as a result of climate
change, however, is likely to increase the rate and magnitude of this
natural process.
In Mozambique, Louro (2006) describes beach driving as a ``very
serious problem.'' Tourism and beach driving are increasing in Ponta
Malongane and Bazaruto Island, nesting areas in Mozambique, where there
is no legislation regarding beach driving (Louro 2006). Foot and
vehicular traffic, for tourism and recreational purposes, have been
found to impact nesting beach habitat and turtles in several ways.
Beach activities can deter females from using a nesting beach. Beach
driving causes sand compaction, which may lower nest success. It also
creates ruts that slow hatchlings' crawl to the surf, increasing their
vulnerability to predators. Beach driving occurs to a lesser extent in
South Africa. Recreational beach driving is allowed on a 1.5 km stretch
of beach, and tourism driving (for concession, management, and media)
involves a maximum of 10 vehicles per night across 40 km of beach (Nel
2006).
Artificial lighting modifies the quality of nesting beaches because
lights over land disorient nesting females and hatchlings. Instead of
crawling toward the surf and their marine habitat, they crawl further
inland, where they may become dehydrated and die or become susceptible
to predation. Within the 280 km of coastline within the iSimangaliso
Wetland Park, South Africa, there are only four areas of less than 100
m each that contain artificial lighting (Nel 2006). We were unable to
find data on artificial lighting in Mozambique.
The majority of nesting habitat occurs within the 280 km coastline
of the iSimangaliso Wetland Park in South Africa, which has been a
World Heritage Site since 1999 (UN Educational, Scientific and Cultural
Organization 1999; Hughes 2010; Robinson et al. 2016). From 1979 to
1999, much of the nesting habitat and nearshore marine habitat was
protected, first as the St. Lucia Marine Reserve, then the Maputaland
Marine Reserve (Hughes 1996). Such protections contributed to the
prevention of dredging a deep water harbor through turtle nesting
beaches and mining heavy minerals in the adjacent dunes (Hughes 2009,
2010). In Mozambique, the Ponta do Ouro Partial Marine Reserve has
provided beach and marine habitat protection since 2009. Additional
protection is provided to Mozambican nesting beaches in: The Ponto du
Ouro--Kosi Bay Transfrontier Marine Conservation Area; the Maputo
Special Reserve; the Bazaruto Archipelago National Park; and the
Quirimbas Archipelago National Park. However, nest protection only
occurs over nine percent of the Mozambique coastline (Videira et al.
2008; Garnier et al. 2012). Such protections have minimized vehicular
traffic at nesting beaches in South Africa, but beach driving remains a
threat in Mozambique. Erosion is a threat to nesting beaches in both
South Africa and Mozambique. Thus, we conclude that the present
modification of nesting habitat is a threat to the SW Indian DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization is a threat to the SW Indian DPS (Bourjea 2015;
Williams et al. 2016; Williams 2017). Two of nine leatherback turtles
equipped with satellite tags between 1996 and 2006 were incidentally or
intentionally captured in Mozambique and Madagascar and likely retained
for food or sale (de Wet 2012). In Mozambique, eggs and turtles were
once legally harvested and are now illegally poached for consumption
(Nel 2012; Wallace et al. 2013; Fernandes et al. 2018). Turtle poaching
includes turtles taken on the beaches and at sea (Williams et al. 2016;
Williams 2017). We do not have recent, quantitative estimates of egg or
turtle poaching in Mozambique. However, significant usage has been
documented at various points in time. Hughes (1995) reported that
nearly every nesting female was killed during the civil war (1977 to
1992). An estimated 32 loggerhead and leatherback turtles were killed
at Ponta Malongane in 11 years (Louro 2006). Recent egg and turtle
poaching rates in Mozambique have been qualitatively described as
``alarming,'' ``significant,'' ``widespread,'' ``prominent,'' and
``prevalent'' (Fernandes et al. 2015; Williams et al. 2016; Williams
2017; Pereira and Louro 2017; Fernandes et al. 2017; Fernandes et al.
2018). Nest monitoring programs in Mozambique have provided some
protection since the 1990s (Garnier et al. 2012). Pereira et al. (2014)
reports that as a result of the monitoring program at the Ponta do Ouro
Partial Marine Reserve, where the majority of nesting in Mozambique
occurs, turtle mortalities are very rare. Egg poaching has been reduced
in the Bazaruto Archipelago, where it was previously prevalent (Louro
2006). National legislation in Mozambique include: Diploma Legislativo
2627 (7 August 1965), Forest and Wildlife Regulation (Decree 12/2002 of
6 June 2002) and Conservation Law (Law 5/2017 of 11 May). These laws
protect turtles and eggs and impose fines for poaching or possession.
However, the laws are poorly implemented and enforced (Costa et al.
2007; Louro 2006; Williams et al. 2016; Fernandes et al. 2018). We
conclude that the poaching of turtles and eggs remains a significant
threat in Mozambique.
Poaching of turtles is also a threat in Madagascar, where
leatherback turtles caught in gillnets are taken back to local villages
and consumed, which is documented to have occurred twice in 2016
(Williams 2017). Leatherback turtles were caught and consumed or sold
in five of eight Malagasy villages surveyed between October 2004 and
March 2004. Fishers reported that leatherback turtles were uncommon but
large, possibly indicative of mature individuals (Walker and Roberts
2005).
[[Page 48377]]
No leatherback turtles were reported caught during a 2007 Malagasy
village survey (Humber et al. 2010). Although protected by Presidential
Decree (2006-400), fishers target turtles at sea for consumption
(Ratsimbazafy 2003; Epps 2006; Humber et al. 2010). Humber et al.
(2010) report that the Malagasy law is not adequately implemented due
to lack of enforcement, a reluctance to manage the local, cultural
fishery, and the size of the coastline (Rakotonirina and Cooke 1994;
Okemwa et al. 2005). We conclude that the poaching of turtles remains a
significant threat in Madagascar.
Egg and turtle poaching does not appear to be a significant threat
in South Africa. Prior to the ban on egg harvest in 1963, substantial
numbers of leatherback eggs in South Africa were harvested, likely
contributing to the critically low number of nesting females at that
time (Nel et al. 2015). Hughes et al. (1996) concluded that nesting
females were not harvested. As a result of the ban, and with a
lucrative tourism industry centered on the nesting turtles, egg and
turtle harvest has been nearly eliminated (Hughes et al. 1996). Nesting
females and hatchlings receive ``intensive and effective'' protection,
as most nesting beaches fall within the iSimangaliso Wetland Park (Nel
et al. 2015). Such beach protections have been key to recovering the
number of nesting females to current levels (Hughes et al. 1996; Saba
et al. 2015; Nel et al. 2015). We conclude that the poaching of turtles
and eggs is not a significant threat in South Africa.
Exposure to poaching is low in South Africa, where the majority of
females nest. Few females nest in Mozambique, reducing the DPS's
overall exposure to egg and nesting female poaching during nesting.
However, turtles regularly forage in the Mozambique Channel, where they
may be poached along the coasts of Mozambique and Madagascar. Poaching
of nesting females or post-nesting females (i.e., on land or at sea)
reduces both abundance (through loss of nesting females) and
productivity (through loss of reproductive potential). Such impacts are
high because they directly remove the most productive individuals from
DPS, reducing current and/or future reproductive potential. Egg
poaching reduces productivity. We conclude that overutilization, as a
result of poaching of turtles and eggs, poses a threat to the DPS.
Disease or Predation
While we could not find any information on disease for this DPS,
predation is a threat to the SW Indian DPS. In South Africa, nest
predators include feral dogs, side-striped jackals, honey badgers, and
ghost crabs (Hughes 1996; Nel 2006). In the 1960s, the removal of feral
dogs greatly reduced nest predation. Similarly, jackals were once a
threat (Hughes 1996). However, nest predation by jackals has not been
observed for 17 years (R. Nel, pers. comm. April 15, 2019). Nel (2006)
reports current rates of predation as relatively low. Nel et al. (2013)
reports that there is no evidence for significant beach predation on
South African beaches. Describing nest predation as minimal in South
Africa, de Wet (2012) found that 15.7 percent of nests were depredated
in the 2009/2010 and 2010/2011 nesting seasons; ants and ghost crabs
were the main cause of egg mortality. During the two seasons, ghost
crabs consumed 3.2 percent of hatchlings as they made their way to the
sea (de Wet 2012).
While all eggs and hatchlings have some exposure to predation, the
species compensated for a certain level of natural predation by
producing a large number of eggs and hatchlings. For this DPS, the
primary impact is to productivity (i.e., reduced egg and hatching
success). We conclude that, though much reduced, predation still poses
a threat to the SW Indian DPS.
Inadequacy of Existing Regulatory Mechanisms
The SW Indian DPS is protected to some degree by several regulatory
mechanisms. For each, we review the objectives of the regulation and to
what extent it adequately addresses the targeted threat.
Despite efforts to reduce impacts, fisheries bycatch continues to
be the primary threat to this DPS (Petersen et al. 2009; Nel et al.
2013; Wallace et al. 2013; Fossette et al. 2014; Angel et al. 2014; Nel
et al. 2015; Harris et al. 2018). To minimize the impacts from longline
fisheries, the FAO published guidelines for sea turtle protection,
entitled Technical Consultation on Sea Turtle-Fishery Interactions (FAO
2004; Huang and Liu 2010). The UN 1995 Code of Conduct for Responsible
Fisheries (FAO 2004) provides guidelines for the development and
implementation of national fisheries policies, including gear
modification (e.g., circle hooks, fish bait, deeper sets, and reduced
soak time), new technologies, and management of areas where fishery and
sea turtle interactions are more severe. The guidelines stress the need
for mitigation measures, data on all fisheries, fishing industry
involvement, and education for fishers, observers, managers, and
compliance officers (FAO 2004; Honig et al. 2007). These guidelines,
however, are rarely enacted in full. The ICCAT has adopted a resolution
for the reduction of sea turtle mortality (Resolution 03-11),
encouraging States to submit data on sea turtle interactions, release
sea turtles alive wherever possible, and conduct research on mitigation
measures. The responsibility to implement mitigation measures remains
within each nation, and many nations have not implemented such measures
(Honig et al. 2007). South Africa, Namibia, and Angola signed the
Memoranda of Understanding concerning Conservation Measures for Marine
Turtles of the Atlantic Coast of Africa. Though South African vessels
are required to carry a dehooker and line-cutter (Honig et al. 2007)
and has instituted an observer program (Petersen et al. 2009), few
other at-sea conservation measures have been implemented (Honig et al.
2007). For Taiwanese fishing vessels operating within the range of this
DPS, Taiwan has regulations to limit the number of vessels in the area
and to require vessels to carry de-hookers. However, bycatch and
mortality remain high (Huang and Liu 2010). Similarly, though the
extent of shark nets off South African beaches has been reduced from 44
km in the early 1990s to 23 km in 2007, bycatch and mortality continue
to occur (Brazier et al. 2012), and Nel et al. (2015) identify bather
protection nets, together with boat strikes, as the second greatest
threat to the DPS, after longline fisheries. Regarding shark nets,
Brazier et al. (2012) concludes that bycatch is low and rates are
stable, but because the leatherback population is small, a further
reduction in bycatch is desirable. Because the offshore longline
fishery contributes more than the shark nets to leatherback mortality,
Brazier et al. (2012) also recommends further introduction of bycatch
reduction techniques in the longline fishery. Because longline threats
are proportionally large and possibly increasing, Harris et al. (2018)
concludes that bycatch mitigation measures in this industry remain
first and most important management action. Thus, existing regulations
have been inadequate to meet their objectives.
Beach habitat is protected throughout a portion of the nesting
range of this DPS. In South Africa, approximately 280 km of nesting
beaches benefit from intensive and effective protection as part of the
iSimangaliso Wetland Park, a World Heritage Site since 1999 (UN
Educational, Scientific and Cultural Organization 1999; Nel et al.
2015). iSimangaliso includes 280 km of beaches, rocky shores,
mangroves, lakes,
[[Page 48378]]
estuaries, and coastal waters out to three nautical miles (5 km) and
200 m depth. Regulations prevent coastal development and commercial
fishing within this area. However, Harris et al. (2015) estimated that
66 percent of leatherback turtles nest outside of the protected
monitoring area (i.e., only 300 km of the 900 km nesting area is
monitored and protected). In addition, leatherback turtles use coastal
waters that are not protected under the marine reserve. In Mozambique,
much of the nesting habitat is protected, including: The Ponto du
Ouro--Kosi Bay Transfrontier Marine Conservation Area; the Maputo
Special Reserve; the Bazaruto Archipelago National Park; and the
Quirimbas Archipelago National Park. However, nest protection only
occurs over nine percent of the Mozambique coastline (Videira et al.
2008; Garnier et al. 2012). Thus, regulations to protect the nesting
habitat of the DPS have been successful. However, leatherback turtles
nesting outside these areas receive no protection.
In addition, South Africa hosts several marine protected areas and
has proposed to add 20 new marine protected areas to expand protection
to five percent of its EEZ (https://www.marineprotectedareas.org.za/).
Two of these were proposed in order to protect leatherback marine
habitat: The 1200 km\2\ iSimangaliso Marine Protected Area (off nesting
beaches); and the 6200 km\2\ Agulhas Front Marine Protected Area
(encompassing core foraging habitat). These initiatives are likely to
protect leatherback turtles within the proposed areas. However, the DPS
has a large range that extends well beyond protected areas. Harris et
al. (2018) identifies the Mozambique Channel as an additional key
priority area to protect.
In South Africa, a 1963 ban on egg and turtle harvest has been
effective in virtually eliminating overutilization (Hughes 1996). The
current law, Regulation 58(7) of the MLRA (1998), provides full
protection to sea turtles and their products. In Mozambique, national
legislation includes: Diploma Legislativo 2627 (7 August 1965), Forest
and Wildlife Regulation (Decree 12/2002 of 6 June 2002) and
Conservation Law (Law 5/2017 of 11 May). These laws protect turtles and
eggs and impose fines for poaching or possession. For example, the
Forest and Wildlife regulation prohibits the killing of turtles and the
possession of their eggs, with fines up to US $1,000 (Decree 12/2002 of
6 June 2002; Costa et al. 2007). In 2008, there were at least 13
conservation programs focusing on protection and education. Despite
these efforts, illegal poaching of eggs and turtles remains prevalent
in Mozambique (Fernandes et al. 2014) due to limited implementation and
enforcement of the environmental legislation (Costa et al. 2007; Louro
2006; Williams et al. 2016; Fernandes et al. 2018). In Madagascar, all
sea turtles are protected from exploitation by Presidential Decree
(2006-400). However, fishers continue to target and consume turtles
captured at sea (Ratsimbazafy 2003; Epps 2006; Humber et al. 2010). The
effectiveness of the Malagasy law is limited due to lack of
enforcement, a reluctance to manage the local, cultural fishery, and
the size of the coastline (Rakotonirina and Cooke 1994; Okemwa et al.
2005; Humber et al. 2010). Thus, while regulations to prevent the
harvest of turtles and eggs have been adequate in South Africa,
regulatory protections in Mozambique and Madagascar are inadequate.
In summary, numerous regulatory mechanisms protect leatherback
turtles, eggs, and nesting habitat throughout the range of this DPS.
Though the regulatory mechanisms provide some protection to the
species, many do not adequately reduce the threat that they were
designed to address, generally as a result of limited implementation or
enforcement. As a result, bycatch, incomplete nesting habitat
protection, and poaching in Mozambique and Madagascar remain threats to
the DPS. In summary, we consider the inadequacy of the regulatory
mechanisms to be a threat to the SW Indian DPS.
Fisheries Bycatch
Fisheries bycatch is the primary threat to the SW Indian DPS
(Wallace et al. 2013; Fossette et al. 2014; Angel et al. 2014; Nel et
al. 2015; Harris et al. 2018). Bycatch occurs in commercial and
artisanal, coastal and pelagic fisheries. Gear types include: Longline,
purse seine, pelagic trawl, shrimp trawl, gillnets, and beach seines
(Honig et al. 2007; Petersen et al. 2009; Nel et al. 2013; Nel et al.
2015).
Of all gear types, longline fisheries likely have the largest
impact on the DPS (Petersen et al. 2009; Nel et al. 2013; Angel et al.
2014; Nel et al. 2015; Harris et al. 2018). Leatherback turtles are
exposed to longline fisheries throughout their foraging range,
including the Benguela Current in the Atlantic Ocean, the Agulhas
Current in the Indian Ocean, and coastal waters off South Africa,
Mozambique, and Madagascar (Honig et al. 2007; Peterson et al. 2009;
Huang and Liu 2010; Harris et al. 2018). Flag states include: South
Africa, Mozambique, Japan, and Taiwan (Honig et al. 2007; Peterson et
al. 2009; Huang and Liu 2010).
Harris et al. (2018) found a positive, significant relationship
between the longline fisheries' extent of overlap with leatherback
migratory corridors and threat intensity (F1,8 = 184.7, P
<0.001, R2 = 0.95), which was defined as a product of the turtles
utilization distribution and the normalized fishing effort. They
concluded that incidental capture in longline fisheries was the most
important offshore threat to leatherbacks and supports the hypothesis
that longlining is suppressing growth of this DPS (Nel et al. 2013;
Harris et al. 2018). Harris et al. (2018) calculated longline bycatch
rates, around Southern Africa, to be 1,500 leatherback turtles
annually. Though this estimate likely includes turtles from other DPSs
(SE Atlantic and NE Indian), the authors concluded that even low
absolute bycatch has a disproportionately large effect in slowing
population growth rates, due to the small nesting female abundance of
the SW Indian DPS (Harris et al. 2018). Additional reason for concern
is that the threat intensity of longlining was especially high in the
last 5 years of the study (ICCAT and IOTC data from 2004 to 2013),
suggesting that the threat and its impacts on the DPS are increasing
(Harris et al. 2018). Throughout the SE Atlantic and SW Indian Oceans,
Harris et al. (2018), Wallace et al. (2013), deWet (2012), Thorson et
al. (2012), and Peterson et al. (2009) analyze longline bycatch over a
large portion of the DPS's foraging range. Wallace et al. (2013)
categorize the longline fishing effort as medium to high and conclude
that such effort leads to a high risk and high bycatch impact for the
SW Indian DPS. Thorson et al. (2012) used data from the IOTC (1954 to
2009) and South African fishery (2006 to 2009) in a model of
leatherback turtle survival and availability. Their model did not find
that leatherback survival declined during the period when longline
fishing effort increase. However, the authors state that their null
result could be explained by an imprecise index of longline effort or
using newer bycatch rates for the South African longline fishery (i.e.,
Petersen et al. 2009). For example, based on fisheries data from 30
South African and Asian pelagic longline vessels operating in the South
African EEZ between 2006 and 2010, De Wet (2012) estimates the mean
annual bycatch to be 7.8 (7.8 S.D.) leatherback turtles,
based on 39 leatherback turtle captures reported over 5 years. Other
studies estimate bycatch to be higher. Based on extrapolations from
[[Page 48379]]
independent observer bycatch reports from 1998 to 2005 (n = 2,256
sets), Peterson et al. (2009) estimates that the South African pelagic
longline fishery for tunas and swordfish captures 50 leatherback
turtles annually, many of which likely belong to the SW Indian DPS (the
remainder belong to the SE Atlantic DPS). Though most (84 percent) were
caught alive, Peterson et al. (2009) estimates the long-term survival
of affected turtles at 50 percent (based on an estimated range of 25 to
75 percent; Aguilar et al. 1995). Peterson et al. (2009) thus estimates
total mortality from the South African pelagic longline fishery to be
25 turtles annually, or around two percent of the total population
(based on a total population size of 1,200 leatherback turtles), which
they conclude is enough to hamper recovery of the SW Indian population.
Nel et al. (2013) agrees with this conclusion, citing a 30 year (1965
to 1995) increasing trend in nesting female abundance that stalled as
the longline fishery expanded from 1990 to 1995. Huang and Liu (2010)
come to a similar conclusion. They report that the longline fishery
operated at a relatively low level until 1995, when South Africa,
Japan, and Taiwan started a joint venture fishing program.
In the Indian Ocean, Huang and Liu (2010) evaluated the Taiwanese
longline fishery bycatch, and Louro (2006) described illegal longlining
in Mozambique waters. Huang and Liu (2010) evaluated observer data from
77 trips (4,409 sets) on Taiwanese large-scale longline fishing
vessels. They identified 84 leatherback turtles captured from 2004 to
2008, with 48 mortalities (57 percent; Huang and Liu 2010).
Extrapolating to the entire Taiwanese longline fishery in the Indian
Ocean, they estimated an average bycatch of 173 leatherback turtles
between 2004 and 2007. This number likely included individuals from the
SW and NE Indian DPSs. In addition to commercial longlining, artisanal
longlining also occurs in the SW Indian Ocean. Illegal longlining off
Mozambique targets sharks and leatherback turtles. The level of take
and mortality is unknown. A program called Eyes on the Horizon reports
such events, when observed (Louro 2006).
In the SE Atlantic Ocean, Honig et al. (2007) and Angel et al.
(2014) evaluate longline bycatch. Honig et al. (2007) evaluated turtle
bycatch by longline fisheries in the Benguela Large Marine Ecosystem by
using data from observer reports, surveys, and specialized trips from
the coastal nations of South Africa, Namibia and Angola. They estimated
bycatch at 672 leatherback turtles annually (based on an annual bycatch
estimate of 4,200 turtles, of which approximately 16 percent are
leatherback turtles) in the southern and central regions and as many as
5,600 leatherback turtles (based on an annual bycatch estimate of
35,000 turtles) for the entire Benguela Large Marine Ecosystem (Honig
et al. 2007). These estimates likely include many leatherback turtles
from the much larger SE Atlantic DPS, but telemetry studies indicate
that the turtles of the SW Indian DPS use this foraging area too
(Luschi et al. 2006; Robinson et al. 2016). Evaluating ICCAT data,
Angel et al. (2014) confirms exposure to high longline fishing effort
but reports that bycatch of this population is low relative to other
leatherback populations. Although Thorson et al. (2012) found that
increased fishing effort had no explanatory power regarding changes in
leatherback survival, other studies identify longline fisheries as the
primary threat to the DPS (Petersen et al. 2009; Nel et al. 2013; Angel
et al. 2014; Nel et al. 2015; Harris et al. 2018). Based on the weight
of evidence, we agree with the latter and conclude that longline
fisheries pose a major threat to the DPS throughout its foraging range.
Other fisheries also impact the SW Indian DPS, possibly resulting
in substantial mortalities. However, these fisheries are not as well
studied, and mortality estimates are not available (Honig et al. 2007;
Nel et al. 2013). Leatherback turtles are caught in artisanal and
commercial shrimp trawl, pelagic trawl, gillnet, purse seine, and beach
seine fisheries (Honig et al. 2007; Petersen et al. 2009; Nel et al.
2013). Citing Walker (2005) and Rakotonirina (1994), Nel (2013) reports
that the number of sea turtles (all species) caught in artisanal
fisheries of the Mozambique Channel could exceed commercial fishery
catches. Honig et al. (2007) echoes this concern for the Benguela
Current Large Marine Ecosystem, citing high mortality rates for these
fisheries in other regions. The Mozambican shrimp trawl fishery
operates in the Sofala Bank of the Mozambique Channel, near leatherback
nesting, migrating, and foraging areas (Luschi et al. 2006; Robinson et
al. 2016). The fishery supports 50 to 96 vessels that employ standard
otter trawl nets in a single or quad-net configuration with an average
tow-time of three hours (Brito 2012). It does not employ TEDs and
incidentally captures several (i.e., at least two to six but possibly
many more) leatherback turtles annually (Louro 2006; Videira et al.
2010; SWOT 2017). In 2001, one shrimp trawler captain reported
capturing more than six leatherback turtles since fishing season
opened; all were captured alive (Gove et al. 2001). Based on 39
interviews with observers, enforcement officers, and vessel operators,
the fleet (n = 50) captures approximately 56 (40)
leatherback turtles; the overall estimated mortality rate for bycaught
turtles is 14 percent (Brito 2012). Given the overlap between the
fishery and an important foraging area, M. Pereira (CTV, pers. comm.,
2019) concludes that the Mozambican shrimp trawl fishery may be one of
the main threats to this DPS. The South African shrimp trawl fishery
has been reduced to two vessels, with an average annual bycatch of less
than one leatherback (Honig et al. 2007; Petersen et al. 2009; Nel et
al. 2013). Domestic shrimp trawling in Eritrea is considered a major
threat to sea turtles, and bycatch is underreported. However,
leatherback turtles are relatively rare in these waters, as
demonstrated by the foreign trawl fleet, which has 100 percent observer
coverage and bycatch records indicating 39 leatherback turtles between
1996 and 2005 (Pilcher et al. 2006).
During a small random sampling exercise in 2013 by onboard
observers from the Research Division of Eritrea, one leatherback turtle
(of 48 sea turtles total) was captured and released (Mebrahtu 2015). On
June 20, 2019, the European Union passed a regulation (PE-CONS 59/1/19
Rev 1) that requires shrimp trawl fisheries to use a turtle excluder
device in European Union waters of the Indian and West Atlantic Oceans.
Gillnets in Macaneta, Mozambique, killed two leatherback turtles
during the 2010 nesting season (Videira et al. 2010) and captured one
in the 2003 nesting season (Louro 2006). In Madagascar, leatherback
turtles are a ``common'' bycatch of the set gillnet shark fishery
(Robinson and Sauer 2013); mortality is likely high given the 24-hour
soak time and propensity for consuming turtle meat. Purse seine
fisheries have a much lower impact than longline fisheries (Angel et
al. 2014); two leatherback turtles were captured (alive) between 1995
and 2010 in the Indian Ocean (Clermont et al. 2012). In the EEZ of all
Indian Ocean French Territories (mostly from the Mozambique Channel),
40 leatherback turtles were captured in unspecified fisheries from 1996
to 1999; 92 percent were released alive (Ciccione 2006).
Shark or bather nets, which are gillnets installed off beaches in
South Africa to limit human-shark interactions, incidentally capture
[[Page 48380]]
leatherback turtles. According to Nel et al. (2015), bather protection
nets and boat strikes together present the second greatest threat to
the DPS, after fisheries. Three of nine leatherback turtles equipped
with satellite tags between 1996 and 2006 were caught in shark nets (de
Wet 2012). Between 1981 and 2008, 150 leatherback turtles were captured
(mean = 5.36; SE = 0.60), of which 20 were mature females and 39 were
mature males (Brazier et al. 2012). Total mortality was 62.7 percent,
with an annual range of 1 to 12 mortalities (mean = 3.4; SE = 0.47;
Brazier et al. 2012). Most turtles were captured in December, the peak
month for nesting, which together with the prevalence of mature
individuals, suggests that bycatch is dominated by adults from nearby
nesting and breeding areas (Brazier et al. 2012). Analyzing these data
over an additional 2 years (1981 to 2010), de Wet (2012) found that 157
leatherback turtles (mean = 5.26; SD = 2.7) were captured in the nets,
with a 62.4 percent mortality rate (mean = 3.3; SD = 1.8).
To reduce bycatch mortality in longlines, South African regulations
require vessels to carry a dehooker and line cutter (Honig et al.
2007). To reduce bycatch in the shark nets, effort was reduced from 44
km of nets in the early 1990s to 23 km in 2007 (Brazier et al. 2012).
Despite these efforts, a previously increasing trend in nesting female
abundance has stalled and ``declined recently'' (Nel et al. 2013).
Individuals (immature and adult turtles) of this DPS are exposed to
high fishing effort throughout their foraging range. Estimates of
bycatch rates, when available, range considerably. For example, Harris
et al. (2018) estimated the annual longline bycatch rates around
Southern Africa to be 1,500 leatherback turtles annually; whereas, de
Wet (2012) estimated the mean annual bycatch to be 7.8 (7.8
S.D.) leatherback turtles. We have annual mortality estimates for few
individual fisheries: n = 25 for South African longline (Peterson et
al. 2009); n = 12 for Taiwanese longline (Huang and Liu 2010); n = 1 to
12 for shark nets (Brazier et al. 2012). Adding in other longline
fisheries and additional gear types may result in more than 100
mortalities annually. These estimates likely include individuals from
other DPSs (i.e., the SE Atlantic and NE Indian). However, because of
the small nesting population, even small levels of mortality have the
potential to slow population growth (Harris et al. 2018). Mortality
reduces abundance, by removing individuals from the population; it also
reduces productivity, when potential nesting females are killed.
Several studies conclude that bycatch has prevented continued
population growth and/or contributed to the recent slight decline in
nesting (Petersen et al. 2009; Huang and Liu 2010; Brazier et al. 2012;
Nel et al. 2013; Harris et al. 2018). We conclude that fisheries
bycatch is the primary threat to the SW Indian DPS.
Vessel Strikes
Vessel strikes are a threat to the SW Indian DPS. According to Nel
et al. (2015), vessel strikes and bather protection nets together
present the second greatest threat to the DPS, after fisheries.
Together these threats kill up to 10 leatherback turtles annually (Nel
et al. 2015). One of 24 leatherback turtles stranded along the South
African coastline between 1972 and 2010 was struck by a boat propeller
(Nel 2008). However, additional mortalities or injuries may go
unnoticed or unreported. Vessel strikes affect adult females returning
to nest, removing individuals and their future reproductive potential.
Thus, this threat reduces the abundance and productivity of the DPS. We
conclude that vessel strikes pose a threat to the DPS.
Pollution
Pollution includes contaminants, marine debris, and ghost fishing
gear. As with all leatherback turtles, entanglement in and ingestion of
marine debris and plastics are threats that likely kill several
individuals a year. For six stranded hatchlings and 24 stranded adults
over the past 40 years, the cause of death was generally unknown.
However, fishery-related injuries, ghost-fishing (i.e., entanglement in
discarded fishing gear), disease, or pollution may be responsible (de
Wet 2012). Plastic pollution may be a main threat in the waters off
Mozambique (M. Pereira, pers. comm., 2019). Outer accumulation of the
Indian Ocean ``garbage patch'' (Cozar et al. 2014) overlaps with
foraging areas in the Mozambique Channel and occurs in waters offshore
from nesting areas in South Africa and Mozambique. Though we were
unable to find ingestion or entanglement data for SW Indian leatherback
turtles, 51.4 percent of gut and fecal samples from loggerhead turtles
(n = 74) captured as bycatch in the Reunion Island longline fishery
contained marine debris, of which plastic comprised 96.2 percent
(Hoarau et al. 2014). Ryan et al. (2016) found that 24 of 40 loggerhead
turtle post-hatchlings had ingested plastics or other anthropogenic
debris. Based on the foraging behavior of leatherback turtles and the
proximity of the ``garbage patch,'' we conclude that the ingestion and
entanglement of marine debris are threats to this DPS.
In addition, State of the World's Sea Turtles (SWOT 2017)
identifies hydrocarbon extraction along the eastern African seaboard,
including northern Mozambique, as the greatest emerging concern for
this DPS. They report that the impact of such activities remain to be
seen (SWOT 2017). However associated oil spills are likely to modify
habitat off nesting beaches and reduce prey availability for all life
stages. Harris et al. (2018) found that the hydrocarbon industry poses
a moderate threat to the DPS because of its spatial overlap with
migratory corridors (second in extent, after longline fisheries). They
expressed concern over the expansion of the hydrocarbon extraction
along the coasts of southern Mozambique and northeastern South African
and the possibility of an oil spill in these areas (Harris et al.
2018). Pretorius (2018) identified 28 significant impacts to sea
turtles as a result of hydrocarbon exploration and production; these
included: Potential water pollution, light pollution, noise pollution,
and habitat destruction. However, Du Preez et al. (2018) reports that
metal and metalloid contaminants do not appear to be a problem for this
DPS. We conclude that pollution poses a threat to the DPS.
Climate Change
Climate change is a threat to the SW Indian DPS. The impacts of
climate change include: Increases in temperatures (air, sand, and sea
surface); sea level rise; increased coastal erosion; more frequent and
intense storm events; and changes in ocean currents.
Angel et al. (2014) identifies coastal erosion as the main beach-
based threat to this population and one that is likely to increase with
climate change. Though coastal erosion is a natural process, sea level
rise (as a result of climate change) increases the rate of erosion and
the amount of beach affected. In Bazaruto Archipelago, Mozambique,
coastal erosion and rising sea levels destroyed approximately 12
percent of nests over 10 seasons of monitoring (Videira and Louro 2005;
Louro 2006). Because leatherback turtles nest lower on the beach than
other sea turtles, their eggs are more at risk of being exposed and
destroyed by increases in sea level and coastal erosion (Boyes et al.
2010). Thus, erosion and rising sea level as a result of climate change
are a threat to the DPS.
Sand temperatures influence leatherbacks' egg viability and sex
determination. Temperatures over 32 [deg]C
[[Page 48381]]
result in death and temperatures below 29.2 [deg]C produce only males
(Rimblot et al. 1985; Rimblot-Baly et al. 1986). Temperature probes on
South African beaches reveal that nests are already close to pivotal
temperatures, with an average of 29.04 [deg]C (S.D. 0.86 [deg]C; range
27.62 to 29.69 [deg]C; Boyes et al. 2010). A modeling study suggests
that even if South African beaches experience a temperature increase of
5 [deg]C, hatching success and emergence success may not be
significantly reduced (Santidri[aacute]n Tomillo et al. 2015). Instead,
nesting females may shift their nesting season to months (e.g., July
through October) when temperature and precipitation would be similar to
current conditions of the current nesting season (i.e., October through
January). However, the authors cautioned that because nesting females
do not change their nesting habits in response to oceanographic
conditions, they may not change their nesting habits in response to
climate change either (Santidri[aacute]n Tomillo et al. 2015). In
addition, a shift in the nesting season could have impacts beyond
hatching success, such as reduced post-hatchling survival and
suboptimal foraging conditions for post-nesting females. We therefore
conclude that increased temperatures may be a threat to the DPS, and
will likely result in impacts ranging from nesting season shifts to
significant nest losses.
The threat of climate change may modify the nesting conditions for
the entire DPS. Impacts likely range from small, temporal changes in
nesting season to large losses of productivity. Because we are already
seeing small impacts due to coastal erosion and sea level rise, we
conclude that climate change is a threat to the SW Indian DPS.
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply to the SW Indian DPS (for a
description of each effort, please see the section on conservation
efforts for the overall taxonomic species): African Convention on the
Conservation of Nature and Natural Resources (Algiers Convention),
Convention on the Conservation of Migratory Species of Wild Animals,
Convention on Biological Diversity, Convention on International Trade
in Endangered Species of Wild Fauna and Flora, Convention on the
Conservation of European Wildlife and Natural Habitats, Convention for
the Co-operation in the Protection and Development of the Marine and
Coastal Environment of the West and Central African Region (Abidjan
Convention) and Memorandum of Understanding Concerning Conservation
Measures for Marine Turtles of the Atlantic Coast of Africa (Abidjan
Memorandum), Convention Concerning the Protection of the World Cultural
and Natural Heritage (World Heritage Convention), FAO Technical
Consultation on Sea Turtle-Fishery Interactions, Indian Ocean Tuna
Commission, The Indian Ocean Tuna Commission, Indian Ocean--South-East
Asian Marine Turtle Memorandum of Understanding, MARPOL, IUCN, Nairobi
Convention for the Protection, Management and Development of the Marine
and Coastal Environment of the Eastern African Region, Ramsar
Convention on Wetlands, UNCLOS, and UN Resolution 44/225 on Large-Scale
Pelagic Driftnet Fishing. Although numerous conservation efforts apply
to the turtles of this DPS, they do not adequately reduce its risk of
extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the SW Indian DPS is at high risk of extinction. The DPS exhibits
a total index of nesting female abundance of 149 females. Such a
limited nesting population size places this DPS in danger of stochastic
or catastrophic events that increase its extinction risk. This DPS
exhibits a slightly decreasing nest trend at monitored nesting beaches
in South Africa. This declining trend has the potential to further
lower abundance and thereby increase the risk of extinction. With only
one nesting aggregation, the DPS lacks spatial structure, and its
genetic diversity is low. Thus, stochastic events could have
catastrophic effects on nesting for the entire DPS, with no potential
source subpopulations to buffer losses or provide additional diversity.
However, the DPS uses multiple, distant, and diverse foraging areas,
providing some resilience against reduced prey availability. Based on
these factors, we find the DPS to be at risk of extinction, likely as a
result of past threats.
Current (ongoing) threats further contribute the risk of extinction
of this DPS. The primary threat to this DPS is bycatch in commercial
and artisanal, pelagic and coastal, fisheries. Longline fisheries
constitute the greatest threat. Though poorly studied, other fisheries
together may have overall mortality rates for affected turtles from
this DPS that rival those from longline fisheries. Fisheries bycatch
reduces abundance by removing individuals from the population. Because
several fisheries operate near nesting beaches, productivity is also
reduced when nesting females are prevented from returning to nesting
beaches. Exposure and impact of this threat are high. Poaching is also
a threat to the DPS. Egg and turtle poaching, while no longer a threat
in South Africa, likely continues in Mozambique. In Madagascar, turtles
are illegally captured at sea and consumed in local villages. Vessel
strikes also pose a threat. Vessel strikes kill several leatherback
turtles each year, including females returning to beaches to nest.
While exposure is low, impacts are high, affecting both abundance and
productivity. Coastal erosion and beach driving in Mozambique modify
nesting habitat and are believed to result in minor reductions in
productivity currently. However, these threats are likely to increase
over time as climate change and tourism increases. Climate change is
likely to result in reduced productivity due to greater rates of
coastal erosion and nest inundation. Predation of eggs and hatchlings
is also a threat. However, although predation has the potential to
reduce productivity, the DPS has likely adapted to predation by native
species, which account for most of the predation at present. Ingestion
of plastics and entanglement in marine debris are threats to all
leatherback turtles, most likely resulting in injury and reduced
health, though sometimes mortality. Though many regulatory mechanisms
are in place, they do not reduce the impact of these threats to levels
that allow the DPS to continue its previous increasing nesting trend.
Thus, the Team unanimously concluded, that the SW Indian DPS is at
high risk of extinction. The total index of nesting female abundance of
149 females makes the DPS highly vulnerable to threats. We determine,
consistent with the team's findings, that the DPS is currently ``in
danger of extinction.'' The slightly declining nest trend and lack of
spatial structure and diversity further contribute to its risk of
extinction. While this small population had an increasing or stable
nesting trend for decades, the lack of continued population growth and
recent decline may indicate that threats have outpaced productivity.
Past egg and turtle harvest initially reduced the nesting female
abundance of this DPS and likely confined its nesting habitat to a
relatively small geographic area, with little diversity or spatial
structure. Currently, fisheries bycatch is the primary present, ongoing
threat. It reduces abundance and productivity (i.e., imminent and
substantial demographic risks) by removing mature and immature
individuals from the
[[Page 48382]]
population at rates exceeding replacement. Though numerous conservation
efforts apply to this DPS, they do not adequately reduce the risk of
extinction. We conclude that the SW Indian DPS is in danger of
extinction throughout its range and therefore meets the definition of
an endangered species. The threatened species definition does not apply
because the DPS is at risk of extinction currently (i.e., at present),
rather than on a trajectory to become so within the foreseeable future.
NE Indian DPS
The Team defined the NE Indian DPS as leatherback turtles
originating from the NE Indian Ocean, south of 71[deg] N, east of
61.577[deg] E, and west of 120[deg] E. The western boundary occurs at
the border between Iran and Pakistan, where the Somali Current begins.
This current, and the cold waters of the Antarctic Circumpolar Current,
likely restrict the nesting range of this DPS. We placed the eastern
boundary at 120[deg] E to approximate the Wallace and Huxley lines,
which are established biogeographic barriers to gene flow between
Indian and Pacific Ocean populations of numerous species. While the
genetic differences between the NE Indian and West Pacific DPSs
demonstrate discreteness, genetic sampling is unavailable from areas
where the nesting range of the DPSs likely meet, preventing us from
defining the boundary more specifically.
The range of the DPS (i.e., all areas of documented occurrence)
extends throughout the Indian Ocean and possibly into the Pacific
Ocean. Records indicate that the species occurs in the waters of the
following nations: India, Sri Lanka, Bangladesh, Myanmar, Thailand,
Malaysia, Indonesia, Vietnam, China, and Philippines (Hamann et al.
2006). Given the range of the DPS, leatherbacks may also occur in the
waters of Pakistan, Australia, Brunei, Cambodia, Philippines, and
Taiwan.
Leatherback turtles of the NE Indian DPS nest on beaches scattered
throughout the NE Indian Ocean. The largest abundance of nesting occurs
on beaches of the Andaman and Nicobar Islands in India. The sandy
beaches of the Andaman and Nicobar Islands consist of soft limestone
formed of coral and shell (Lal 1976; Bandopadhyay and Carter 2017). A
moderate amount of nesting occurs in Sri Lanka, and even less occurs in
Thailand and Sumatra, Indonesia (Hamann et al. 2006; Nel 2015).
Information on this DPS is limited, but foraging appears to occur
throughout the Indian Ocean (Andrews et al. 2006; Hamann et al. 2006).
The foraging range extends throughout the Bay of Bengal, south of Sri
Lanka, and along the west coast of Sumatra, Indonesia, as indicated by
satellite telemetry data and fisheries reports (NMFS and FWS 2013).
Nesting females at Little Andaman Island likely use a variety of
foraging areas and have been tracked to: South and east of the Andaman
and Nicobar Islands; along the coast of Sumatra; beyond Cocos (Keeling)
Island towards Western Australia; and across the Indian Ocean towards
Madagascar and the African continent (Namboothri et al. 2012;
Swaminathan et al. 2017; Swaminathan et al. 2019). Stranding data also
indicate the use of diverse foraging areas: 15 individuals stranded or
were caught in fishing gear along the mainland coast of India (Shanker
2013). Leatherback turtles have also stranded along the coasts of
Mindanao, Philippines and Pakistan (Firdous 2006; Lucero et al. 2011).
Abundance
The total index of nesting female abundance of the NE Indian DPS is
109 females. We based this total index on the nesting aggregations at
South and West Bays, Little Andaman Island, India (K. Shanker pers.
comm., 2018). Our total index does not include 14 unquantified nesting
aggregations in Bangladesh, India, Indonesia, Malaysia, Myanmar, Sri
Lanka, Thailand, Philippines, and Vietnam. To calculate the index of
nesting female abundance, we divided the total number of nests at South
and West Bays, Little Andaman Island between the 2015/2016 and 2017/
2018 nesting seasons (i.e., a 3-year remigration interval; Andrews
2002) by the clutch frequency (3.8 clutches/season; Andrews 2002;
Eckert et al. 2015). This number represents an index of abundance for
this DPS, and is likely to be an underestimate, because it only
includes available data from recently and consistently monitored
nesting beaches. Additional nesting occurs at other locations but is
unquantified.
Published estimates of total nesting female abundance are not
available for this DPS. The IUCN Red List assessment did not provide an
estimate of the total number of mature individuals because monitoring
was not sufficient (Tiwari et al. 2013). Currently, the largest nesting
aggregations occur in the Andaman and Nicobar Islands of India. Nesting
in Sri Lanka may consist of about 100 to 200 nesting females per year,
and low levels of nesting occur in Thailand and Sumatra, Indonesia
(Hamann et al. 2006; Nel 2012). Low and scattered nesting occurs in
Indonesia: 1 to 14 nesting females annually at Alas Purwo in East Java;
and one to three nesting females annually on three beaches in Bali.
There are also rare reports of nesting in the Philippines (Lucero et
al. 2011; Arguelles 2013), Vietnam, and Malaysia. In Myanmar, nesting
is rare, and only one confirmed nesting event has been recorded in
recent years (i.e., December 2016; Platt et al. 2017). Historically,
there may have been nesting in Bangladesh, but no current reports exist
(Hamann et al. 2006).
Malaysia once hosted the DPS's largest nesting aggregation (Chan
and Liew 1996). It is now considered functionally extinct or extirpated
(Pilcher et al. 2013), as a result of continuous, large-scale egg
harvest and fisheries bycatch (Chan and Liew 1996; Eckert et al. 2012).
The major nesting site in Malaysia, Rantau Bang in Terengganu,
decreased drastically from 10,000 nests in the 1950s to 10 or fewer
nests in the 2010s (reviewed by Eckert et al. 2012), and to one or no
nests annually, more recently. The number of nesting females in Vietnam
has also decreased dramatically, from approximately 500 nesting females
in the 1960s to two to three nests in 2005 and 2007 (The Chu and Nguyen
2015). In the late 1970s, females nested in multiple locations of
Thailand, including: along the airport beach in Changwat Phuket; in the
Laem Phan Wa marine reserve; and in coastal Changwan Phangnga (Bain and
Humphry 1980). Settle (1995) recorded about 30 nests on the Phuket and
Phangnga coastlines from 1992 to 1993. Aureggi et al. (1999) found nine
nests between 1997 and 1998, during a survey of Phra Thong Island in
the south.
Our total index of nesting female abundance (109 females) places
the DPS at risk for environmental variation, genetic complications,
demographic stochasticity, negative ecological feedback, and
catastrophes (McElhany et al. 2000; NMFS 2017). These processes,
working alone or in concert, place small populations at a greater
extinction risk than large populations, which are better able to absorb
losses in individuals. Due to its small size, the DPS has restricted
capacity to buffer such losses. Historic abundance estimates indicate
that the DPS was once much larger. The current abundance is likely a
result of past and current threats, which we describe below. Given the
intrinsic problems of small population size, we conclude that the
limited nesting female abundance is a major factor in the extinction
risk of this DPS.
[[Page 48383]]
Productivity
The NE Indian DPS has exhibited a drastic population decline with
extirpation of its largest nesting aggregation in Malaysia. Nest counts
from Malaysia exhibited a steep decline of 17.9 percent annually (sd =
4.2 percent; 95 percent CI = -25.5 to -8.4 percent; f = 0.998; mean
annual nests = 1,166) over the 44-year period of data collection (1967
to 2010). The drastic decline of nests observed in Malaysia is likely
representative of the overall trend for the DPS given the magnitude of
historical abundance for that site and the high confidence in the trend
estimate.
Despite the dramatic population decline, driven by the extirpation
of the largest nesting aggregation (i.e., Malaysia), productivity
parameters are similar to the species averages. However, we have a low
degree of confidence in these estimates due to limited monitoring of
existing nesting aggregations. We conclude that the NE Indian DPS
exhibits a declining nesting trend, which increases its extinction
risk.
Spatial Distribution
For the NE Indian DPS, nesting is limited to a few, scattered
nesting beaches. Currently, the majority of the nesting occurs on
beaches of the Andaman and Nicobar Islands and Sri Lanka, with small
numbers of nests on the western coast of Thailand, Sumatra, and Java
(Nel et al. 2015).
Spatial structure is unknown but presumed to be low. There are no
estimates of genetic population structure within this DPS because
published genotypes only exist for Malaysia (Dutton et al. 1999, 2007).
Genetic samples were taken from nesting females at Little Andaman
Island from 2008 through 2010, but the results are not yet available
(Namboothri et al. 2010).
The wide distribution of foraging areas likely buffers the DPS
somewhat against local catastrophes or environmental changes that would
limit prey availability. Remaining nesting is limited to a few,
scattered but broadly distributed nesting sites. The largest nesting
aggregations are clustered, thus rendering the DPS susceptible to
environmental catastrophes (e.g., tsunamis), and directional changes
(e.g., sea level rise). Thus, despite widely distributed foraging
areas, the somewhat limited nesting distribution increases the
extinction risk of the NE Indian DPS.
Diversity
Genetic diversity of the NE Indian DPS is potentially relatively
high, based on analyses of samples collected from the previously large,
but now functionally extinct, nesting aggregation in Malaysia (Dutton
et al. 1999, 2007); genetic diversity has not been assessed at other
nesting sites. The diversity of nesting sites is low, given that the
majority of the nesting currently occurs on islands (Sivasundar and
Prasad 1996). We conclude that existing diversity provides little
resilience to the DPS.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
Erosion, coastal development, and artificial lighting have
destroyed or modified the available, suitable nesting habitat and thus
are threats to the NE Indian DPS.
Erosion reduces the available nesting habitat for the DPS. Some
erosion occurs as a result of natural disasters. In 2004, a major
earthquake occurred off the west coast of Sumatra, Indonesia, resulting
in the 2004 tsunami, which destroyed many of the beaches that once
hosted over 1,000 nests (Subramaniam et al. 2009). As a result of the
tsunami, the width of the coastline was reduced by one meter, severely
modifying the beaches of South Bay, Little Andaman Island, and
resulting in very low leatherback nesting in 2005 and 2006 (Namboothri
2010). The tsunami also caused drastic changes at other leatherback
nesting beaches (Alfred et al. 2005; Ramachandran et al. 2005; Murugan
2005; Andrews et al. 2006). It caused erosion at some beaches and
accretion at others, especially in the Andaman and Nicobar Islands,
which lie closest to the epi-center of the earthquake and host the
largest numbers of nesting females in the DPS (Subramaniam et al.
2009). In addition, the beaches in Indonesia are being lost due to
erosion from high tides and monsoons (Obermeier 2002).
Sand mining and tourism-related development are the main threats to
nesting habitat (Fatima et al. 2011). While we were unable to find
specific information regarding sand mining, coastal development is
increasing in Sri Lanka, India, and Bangladesh. The beaches of Sri
Lanka are under high threat from tourism development (e.g., large
hotels and restaurants), urban and industrial development, and
artificial lighting (Kapurisinghe 2006). Along the mainland of India,
granite blocks and embankments prevent turtles from approaching many
beaches (Andrews et al. 2006). Intense coastal development, stemming
from the tourism industry, occurs in Bangladesh without environmental
review (Pilcher 2006), resulting in the alteration of sand dunes and
other activities that reduce the quality of nesting habitat (Islam
2002; Islam et al. 2011). In Vietnam, increasing tourism is expected to
result in coastal development on the beaches of Son Tra Peninsula,
QuanLan, and Minh Chau (Ministry of Fisheries 2003). In addition, most
Vietnam beaches are affected by a large amount of marine debris (e.g.,
glass, plastics, polystyrenes, floats, nets, and light bulbs), which
can entrap turtles and impede nesting activity.
Artificial lighting modifies the quality of nesting beaches because
lights over land disorient nesting females and hatchlings. Instead of
crawling toward the surf and their marine habitat, they crawl further
inland, where they may become dehydrated and die or are susceptible to
predation. Nests moved to hatcheries as part of conservation efforts
may be subject to inadequate hatchery practices, which have resulted in
skewed sex ratios and low hatching success (Chan and Liew 1996;
Kapurisinghe 2006; Rajakaruna et al. 2013; Phillott et al. 2018).
Some areas are protected. Of the 306 islands in the Andaman and
Nicobar Islands of India, 94 are designated as wildlife sanctuaries,
six of which are national parks, and two of which are marine national
parks (Andrews et al. 2006). In Sri Lanka, in 2006, sea turtle
sanctuaries were established at Rekawa (4.5 km stretch) and Godawaya
(3.8 km stretch), where high frequency leatherback nesting is observed;
the area is bounded 500 meters towards the sea and 100 meters towards
the land from the high tide level in both sites (Phillott et al. 2018).
Although laws protect sea turtles throughout Sri Lanka, most nesting
areas are not protected and hence, local communities can disturb
nesting beaches by removing sand, lighting the beaches, and cutting the
beach vegetation (Phillott et al. 2018). In Malaysia, turtle
sanctuaries have been established in Terengganu, Sabah, and Sarawak.
However, nesting habitat modification and destruction continue in many
areas.
We conclude that nesting females, hatchlings, and eggs are exposed
to the reduction and modification of nesting habitat, as a result of
erosion, coastal development, and artificial lighting. These threats
impact the DPS by reducing nesting and hatching success, thus lowering
its productivity. The most abundant remaining nesting aggregations are
protected from
[[Page 48384]]
development, but they experience high rates of erosion; other nesting
beaches are subject to anthropogenic threats. Thus, we conclude that
habitat loss and modification pose a threat to the NE Indian DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Overutilization is a threat to the NE Indian DPS. The harvest of
turtles and eggs led to the historical decline of the DPS, and poaching
continues in several areas (Phillott et al. 2018).
Regular, nearly complete egg harvest caused the functional
extinction of the once large nesting aggregation in Malaysia (Chan and
Liew 1996). In the early 1960s, the Terengganu, Malaysia nesting
beaches were leased to the highest bidder, and nearly all leatherback
eggs were harvested. In the 1980s, the State Fisheries Department tried
to buy back about 10 percent of the harvested eggs to be incubated in a
hatchery (Siow and Moll 1982; Chan and Liew 1996; Stiles 2009).
However, such efforts could not prevent the extirpation. Excessive egg
harvest, both legal and illegal, also caused declines in India, Sri
Lanka, and Thailand (Ross 1982).
The harvest of turtles and eggs continues but has not been
quantified (Nel 2012). In Sri Lanka, almost all eggs are taken from the
beach and sold at markets or to hatcheries for ecotourism (Kapurusinghe
2000, 2006; Rajakaruna et al. 2013; Phillott et al. 2018). The
conservation benefit provided by hatcheries in Sri Lanka is debatable
(Phillott et al. 2018) because they do not follow the hatchery
practices established by the IUCN (Hewavisenthi 1994; IUCN 2005;
Namboothri et al. 2012; Rajakaruna et al. 2013; Phillott et al. 2018).
Egg harvest also continues in Thailand. Commercial egg harvest is
illegal in the Andaman and Nicobar Islands, and in the Andaman Islands,
a ban on hunting and harvesting of turtles came into force in 1977.
However, the original inhabitants of the Andaman and Nicobar Islands
are exempt from the Indian Wildlife Protection Act (Shanker and Andrews
2004), and Namboothri et al. (2012) observed intense egg harvest at
Delgarno, Trilby, and East Turtle Islands. In Myanmar, despite
regulations prohibiting the consumption of turtle meat and eggs (Hamann
et al. 2006), there is illegal trade of turtles caught at sea,
including leatherback turtles (Murugan 2007). In Sri Lanka, the
historically high direct take of turtles at sea has been reduced
(Kapurushinghe 2006). Records indicate that turtle meat and parts were
once regularly exported from Tamil Nadu, India, to Sri Lanka, and then
to other nations such as the United States, Singapore, and Belgium
(Kuriyan 1950; Chari 1964; Shanmughasundarun 1968 as cited in
Agastheesapillai and Thiagarajan 1979).
Exposure to egg and turtle poaching remains high throughout the
range of the DPS. Poaching of nesting females or post-nesting females
at sea reduces both abundance (through loss of nesting females) and
productivity (through loss of reproductive potential). Such impacts are
high because they directly remove the most productive individuals from
the DPS, reducing current and future reproductive potential. Egg
harvest reduces productivity only, but, as previously demonstrated
within this DPS, can have devastating population-level impacts. We
conclude that overutilization, as a result of egg and turtle harvest,
poses a major threat to the NE Indian DPS.
Disease or Predation
While we could not find any information on disease for this DPS,
the best available data indicate that predation is a threat to the NE
Indian DPS. Multiple predators prey on eggs and hatchlings at several
nesting beaches (Andrews 2000). During a 2016 survey of the Nicobar
Islands, approximately 57 percent (n = 1,223) of leatherback nests were
lost to depredation by feral dogs, water monitor lizards, or feral pigs
(Sus domesticus; Swaminathan et al. 2017). In the South Bay of Great
Nicobar Island, wild boars and dogs prey on eggs, while fiddler crabs,
dogs, and raptors prey on hatchlings (Sivakumar 2002). Sivasundar and
Prasad (1996) documented that Asian water monitor lizards took 68.6
percent of leatherback nests in the Andaman Islands. In Sri Lanka, egg
predators include feral dogs, water and land monitor lizards, jackals,
wild boars, mongooses, and ants. Egg predation by feral pigs is a major
threat in Indonesia (Maturbongs et al. 1993; Maturbongs 1995, 1996;
Sivasundar and Prasad 1996).
A large number of eggs and hatchlings are exposed to predation.
Though leatherback turtles produce a large number of eggs and
hatchlings, published rates of predation (57 to 69 percent) are high.
The predation of eggs and hatchlings mainly impacts productivity. We
conclude that predation poses a threat to the NE Indian DPS.
Inadequacy of Existing Regulatory Mechanisms
Turtles of the NE Indian DPS are protected by several regulatory
mechanisms. For each, we review the objectives of the regulation and to
what extent it adequately addresses the targeted threat. Nearly all
nations that host nesting aggregations have legislation to protect sea
turtles.
In India, the leatherback turtle is included on Schedule I, Part II
of the Wildlife (Protection) Act, 1972 (Entry No. 11) updated by Wild
Life (Protection) Amendment Act, 2002 (No. 16 of 2003). India also bans
the hunting and trade of wild animals (India National Report to CMS,
1991 and 1994). However, the indigenous people of the Andaman and
Nicobar Islands are exempt from these laws. India has regulations to
require TEDs and minimize fisheries interactions; and much of the
Andaman and Nicobar Islands are protected as wildlife sanctuaries,
including two marine national parks (Andrews et al. 2006).
In Indonesia, Order No. 301/1991 lists leatherback turtles as a
protected species. Pursuant to the Act of 10 August 1990 on the
Conservation of Living Resources and Their Ecosystems, it is prohibited
to kill, capture, possess, transport, trade in or export protected
animals whether alive or dead, or parts of such animals. The taking,
destruction, trade or possession of the eggs or nests of protected
animals are also prohibited (ECOLEX 2003). There are no habitat
protections and no regulations to minimize fisheries interactions or
require TEDs in Indonesia.
In Sabah, Malaysia, the leatherback turtle is not listed as a
totally protected or partially protected species in the Wildlife
Conservation Enactment (No. 6 of 1997). In Sarawak, Malaysia,
leatherback turtles have been fully protected since 1958. Under the
Wildlife Protection Ordinance 1998, all marine turtles in Malaysia are
protected from hunting, killing, capture, sale, import, export,
possession of any animal, recognizable part or derivative or any nest,
except in accordance with the permission in writing of the Controller
of Wildlife for scientific or educational purposes or for the
protection or conservation of a species (Tisen and Bali 2002). The
nesting beach at Rantau Abang, Terengganu is protected. However, the
nesting aggregation that once used this beach has been extirpated. In
1994, the waters surrounding 38 offshore islands of Peninsular Malaysia
and Labuan became protected as marine parks. In
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addition, one national park in Sarawak, three in Sabah, and one state
park in Terengganu protect coastal and marine ecosystems (Malaysia
National Biodiversity Policy 1998). Additional habitat protections
include: The Turtle Trust Ordinance 1957; Land Code 1958; Turtle
Protection Rules 1962; Fisheries Prohibited Areas under section 61 of
the Fisheries Act 1985; and the Wildlife Protection Ordinance 1998
(Tisen and Bali 2002). The use of TEDs will be required in Malaysia by
2020.
In Myanmar, the Burma Wildlife Protection Act 1936 (Act No. VII of
1936) requires licenses to hunt, possess, sell, or buy wild animals
with closed hunting seasons (FAOLEX 2003). The Burma Wildlife
Protection Rules of 1941 states that the import or export of any
reptile (including parts or products) into or from Myanmar is
prohibited.
In Pakistan, the leatherback turtle is protected in Baluchistan,
Azad Kashmir and Sind (Baluchistan Wildlife Protection Act 1974 No.19/
1974; The Azad Jammu and Kashmir Wildlife Act 1975 No.23/1975; The
Sindh Wildlife Protection Ordinance 1972 No.5/1972). Possession,
transport, and/or national trade are prohibited or regulated (ECOLEX
2003).
In Sri Lanka, the leatherback turtle is protected under the Fauna
and Flora Protection Ordinance (Sri Lanka National Report to CMS 1994),
which makes it an offense to kill, wound, harm or take a turtle, or to
use a noose, net, trap, explosive or any other device for those
purposes, to keep in possession a turtle (dead or alive) or any part of
a turtle, to sell or expose for sale a turtle or part of a turtle, or
to destroy or take turtle eggs. The minister of Fisheries and Aquatic
Resources may also prohibit or regulate the import and export of
turtles or their derivatives (Parliament of the Democratic Socialist
Republic of Sri Lanka 1993). The nesting beach in Yala Reserve is also
protected.
In Thailand, the Leatherback Turtle is protected under the Animals
Protection Act B.D 2535 (The Zoological Park Organization 2003).
In summary, numerous regulatory mechanisms protect leatherback
turtles, their eggs, and nesting habitat throughout the range of this
DPS. Although these regulatory mechanisms provide some protection, many
do not adequately reduce the threat that they were designed to address,
generally as a result of limited implementation or enforcement. As a
result, bycatch, nesting habitat protection, and legal and illegal
harvest remain threats to the DPS. We conclude that the inadequacy of
the regulatory mechanisms is a threat to the NE Indian DPS.
Fisheries Bycatch
Fisheries bycatch is a threat to the NE Indian DPS. Capture in
gillnet, trawl, purse seine, and longline fisheries is a significant
cause of leatherback mortality for this DPS (Wright and Mohanty 2002;
Hamann et al. 2006; Project GloBAL 2007; Bourjea et al. 2008;
Abdulqader 2010; Wallace et al. 2010).
Gillnet fisheries pose a major threat to the DPS. A survey
conducted at 16 main fishing ports in Sri Lanka estimated that 431
leatherback turtles were caught in gillnets between 1999 and 2000
(Kapurusinghe and Cooray 2002). In Malaysia, Chan et al. (1988)
reported an average of 742 and 422 sea turtles, most of which were
leatherback turtles, caught in drift gillnets and bottom longlines,
respectively. In Bangladesh, gillnets, set bag nets, trawl nets, seine
nets, hook and line and other net types of gear capture turtles
(Hossain and Hoq 2010). Gillnet and purse seine fisheries are common
off the coasts of the Andaman and Nicobar Islands, where the largest
nesting aggregations occur (Shanker and Pilcher 2003; Chandi et al.
2012).
Trawl fisheries also pose a large threat to the DPS. In India, TEDs
are required for trawl nets. However, fishers are reluctant to use them
(Murugan 2007). Trawl fishing is also common in Bangladesh, and the use
of TEDs is not required (Ahmed et al. 2006).
Longline fisheries occur in coastal and pelagic waters. Huang and
Liu (2010) evaluated observer data from 77 trips (4,409 sets) on
Taiwanese large-scale longline fishing vessels in the Indian Ocean.
They identified 84 leatherback turtles captured from 2004 to 2008, with
48 mortalities (57 percent; Huang and Liu 2010). Extrapolating to the
entire Taiwanese longline fishery in the Indian Ocean, they estimated
an average bycatch of 173 leatherback turtles between 2004 and 2007.
This number likely includes individuals from both the SW and NE Indian
DPSs (Louro 2006). In Vietnam, longline fisheries continue to capture
leatherback turtles. However, a circle hook program has been
implemented to minimize the impact (WWF 2013).
Purse seine fisheries have a much lower impact than longline
fisheries (Angel et al. 2014); two leatherback turtles were captured
(alive) between 1995 and 2010 in the Indian Ocean (Clermont et al.
2012). In the EEZ of all Indian Ocean French Territories (mostly from
the Mozambique Channel), 40 leatherback turtles were captured in
unspecified fisheries from 1996 to 1999; 92 percent were released alive
(Ciccione 2006).
In Thailand, one of the main causes of decline in the turtle
population is bycatch in trawl, drift gillnet, and purse seine
fisheries. The rapid expansion of fishing operations is largely
responsible for the increase in adult turtle mortality due to bycatch
(Settle 1995).
In Malaysia, the Fisheries Act of 1985 prohibited capture of sea
turtles by any type of fishery. However, this merely reduced the
reporting of interactions (Yeo et al. 2011 in Dutton et al. 2011). The
1991 Regulations prohibit fishing in waters adjacent to Rantau Abang
during the leatherback nesting season (Chan 1993).
We conclude that juveniles and adults are exposed to high fishing
effort throughout their foraging range and in coastal waters near
nesting beaches. Mortality rates are likely high, especially in areas
where turtle meat is consumed. Mortality reduces abundance, by removing
individuals from the population. It also reduces productivity, when
nesting females are incidentally captured and killed. We conclude that
fisheries bycatch is a major threat to the NE Indian DPS.
Pollution
Pollution includes contaminants, marine debris, and ghost fishing
gear. Ghost fishing gear can drift in the ocean and fish unattended for
decades and kill numerous individuals (Wilcox et al. 2013). The main
sources of ghost fishing gear are gillnet, purse seine, and trawl
fisheries (Stelfox et al. 2016). In one collection event, volunteers
collected over 600 nets, ropes, and buoys from India, Maldives, Oman,
Pakistan, Sri Lanka, and Thailand (Stelfox et al. 2016). Though
educational programs created in 2014 focus on reusing and recycling
fishing gear, the threat continues throughout the range of the DPS.
Ghost nets in the Maldives primarily drift from fisheries in the Bay of
Bengal (e.g., Sri Lanka and India; Stelfox et al. 2016). Around the
Andaman and Nicobar Islands and Sri Lanka, plastics and other garbage
are washed from polluted beaches and inland waters to the sea, where
they can kill or harm sea turtles through ingestion or entanglement
(Kapurusinghe 2006; Das et al. 2016). Pollution has been identified as
a main threat to sea turtles in Iran (Mobaraki 2007) and Pakistan
(Firdous 2001). However, no specific information about the type of
pollution was provided. In Gujarat, India, increased port and shipping
traffic have resulted in oil spills and the release of other
[[Page 48386]]
pollutants, such as fertilizers and cement (Sunderraj et al. 2006).
Heavy metals and E. coli were found at relatively high levels in the
waters of Malaysia (including Terengganu) and in the pancreases and
livers of leatherback turtles (Caurant et al. 1999; Ngah et al. 2012).
It is not known how these pollutants affect leatherback physiology
(Jakimska et al. 2011).
As with all leatherback turtles, entanglement in and ingestion of
marine debris and plastics are threats that likely kill several
individuals a year. However, data specific to this DPS were not
available. We conclude that pollution is a threat to the NE Indian DPS,
albeit with effects that are unquantifiable on the basis of the best
available information.
Climate Change
Climate change is a threat to the NE Indian DPS. A significant rise
in sea level would further reduce nesting habitat, which is already
affected by erosion. The DPS is also likely to be affected by increases
in sand temperatures (Hawkes et al. 2009; Poloczanska et al. 2009).
Sand temperatures prevailing during the middle third of the incubation
period determine the sex of hatchling sea turtles (Mrosovsky and Yntema
1980). Incubation temperatures near the upper end of the tolerable
range produce only female hatchlings, while incubation temperatures
near the lower end of the tolerable range produce only males. As
temperatures increase, incubation temperatures may exceed the thermal
tolerance for embryonic development, thus increasing embryo and
hatchling mortality.
In addition, the frequency and intensity of severe storm events and
cyclones in the Bay of Bengal are predicted to increase with climate
change (Balaguru et al. 2014).
Climate change is likely to modify nesting conditions for the
entire DPS. Impacts likely range from small changes in nesting metrics
to large losses of productivity. As the DPS is already experiencing
nesting habitat loss due to coastal erosion, we conclude that climate
change is a threat to the NE Indian DPS.
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply to the NE Indian DPS (for a
description of each effort, please see the section on conservation
efforts for the overall species): Association of Southeast Asian
Nations Ministers on Agriculture and Forestry, Andaman and Nicobar
Island Environmental Team, The Centre for Herpetology/Madras Crocodile
Bank Trust, Convention on the Conservation of Migratory Species of Wild
Animals, Convention on Biological Diversity, Convention on
International Trade in Endangered Species of Wild Fauna and Flora,
Convention Concerning the Protection of the World Cultural and Natural
Heritage (World Heritage Convention), FAO Technical Consultation on Sea
Turtle-Fishery Interactions, The Indian Ocean Tuna Commission, Indian
Ocean--South-East Asian Marine Turtle Memorandum of Understanding,
MARPOL, IUCN, Memorandum of Agreement between the Government of the
Republic of the Philippines and the Government of Malaysia on the
Establishment of the Turtle Island Heritage Protected Area, Memorandum
of Understanding on Association of South East Asian Nations Sea Turtle
Conservation and Protection, The Memorandum of Understanding of a Tri-
National Partnership between the Government of the Republic of
Indonesia, the Independent State of Papua New Guinea and the Government
of Solomon Islands, National Sea Turtle Conservation Project in India,
Ramsar Convention on Wetlands, UNCLOS, and UN Resolution 44/225 on
Large-Scale Pelagic Driftnet Fishing. Although numerous conservation
efforts apply to the turtles of this DPS, they do not adequately reduce
its risk of extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the NE Indian DPS is at high risk of extinction. The once large
nesting aggregation in Malaysia is now functionally extirpated. The
total index of nesting female abundance is 109 females at all monitored
beaches. This estimate is likely low because several nesting sites were
not included in the calculation due to lack of consistent, standardized
monitoring over multiple and entire nesting seasons. Still, the low
nesting female abundance places this DPS at risk of stochastic or
catastrophic events that increase its extinction risk. The DPS once
exhibited much greater nesting female abundance, which has dramatically
declined in recent decades. It currently exhibits a slightly declining
nest trend at monitored nesting beaches in India. The DPS exhibits
average productivity metrics, such as body size, clutch size and
frequency. Though it exhibits some spatial distribution and diversity,
with multiple foraging sites and relatively high genetic diversity at
the sampled locations, nesting only occurs on islands. Based on these
factors, we find the DPS to be at risk of extinction as a result of
past threats.
Current threats further contribute to the risk of extinction of
this DPS. Major threats to the DPS include fisheries bycatch and the
harvest of turtles and eggs. There are not many nests to exploit, but
evidence suggests that if such nests are found by humans, the eggs are
at risk of being harvested. Egg harvest led to the extirpation of the
largest nesting aggregation (i.e., Malaysia), and current
overexploitation occurs in Thailand, Vietnam, and Sri Lanka. The
poaching of turtles is also a threat in Myanmar. Fisheries bycatch is a
major threat, with turtles being captured in trawl and gillnet
fisheries in Malaysia, India, Thailand, Sri Lanka, Bangladesh, and
Indonesia. Erosion on the Andaman and Nicobar Islands, as a result of
tsunami damage, has significantly reduced available nesting habitat.
Additional habitat modifications include coastal development and
artificial lighting, as a result of increases in tourism. Pollution and
climate change are threats that likely affect the DPS by reducing
abundance and productivity, though the best available data do not allow
for quantification of those effects. Though many regulatory mechanisms
are in place, they do not reduce the impact of threats to levels that
ensure the continued existence of the DPS.
We conclude, consistent with the team's findings, that the NE
Indian DPS is currently in danger of extinction. Its low nesting female
abundance makes the DPS highly vulnerable to threats. Dramatic declines
in over the past several decades contribute to our concern over the
continued persistence of the DPS. Past egg and turtle harvest initially
reduced the nesting female abundance of this DPS and likely confined
its nesting habitat to a few island beaches, with little diversity and
reduced spatial distribution. The present, ongoing threats include:
overutilization (i.e., turtle and egg harvest); fisheries bycatch; loss
of habitat; and predation. Overutilization and fisheries bycatch
reduces abundance and productivity (i.e., imminent and substantial
demographic risks) by removing mature and immature individuals from the
population at rates exceeding replacement. The loss of nesting habitat
and predation (of eggs) reduces productivity and the DPS's ability to
recover to its previous abundance. Though numerous conservation efforts
apply to this DPS, they do not adequately reduce the risk of
extinction. We conclude that the NE Indian DPS is in danger of
extinction throughout its
[[Page 48387]]
range and therefore meets the definition of an endangered species. The
threatened species definition does not apply because the DPS is at risk
of extinction currently (i.e., at present), rather than on a trajectory
to become so within the foreseeable future.
West Pacific DPS
The Team defined the West Pacific DPS as leatherback turtles
originating from the West Pacific Ocean, south of 71[deg] N, north of
47[deg] S, east of 120[deg] E, and west of 117.124[deg] W. The northern
and southern boundaries reflect the highest latitude occurrences of
leatherback turtles in each hemisphere (Goff and Lien 1988; Carriol and
Vader 2002; McMahon and Hayes 2006; Shillinger et al. 2008; Benson et
al. 2011; Eckert et al. 2012). We placed the western boundary at
120[deg] E to approximate the Wallace and Huxley lines, which are
established biogeographic barriers to gene flow between Indian and
Pacific Ocean populations of numerous species. While the genetic
differences between the Northeast Indian and West Pacific DPSs
demonstrate discreteness, genetic sampling is unavailable from areas
where the nesting ranges of the DPSs likely meet, preventing us from
defining the boundary more specifically. We placed the eastern boundary
at the border between the United States and Mexico to reflect the DPS's
wide foraging range throughout the Pacific Ocean. We chose this border
because the West Pacific DPS crosses the ocean to forage in the eastern
Pacific Ocean, including in waters of the United States, whereas the
East Pacific DPS forages primarily off the coasts of Central and South
America. The two DPSs overlap in foraging habitats off waters of Chile
and Peru (Donoso and Dutton 2010).
The range of the DPS (i.e., all areas of occurrence) extends
throughout the Pacific Ocean with specific coastal and pelagic areas in
the Indo-Pacific basin providing important foraging and migratory
habitats. Documented nesting occurs on beaches of the following
nations: Indonesia, Papua New Guinea, Solomon Islands, and Vanuatu.
Leatherback turtles of the West Pacific DPS migrate through the EEZs of
at least 32 nations including in the U.S. EEZs of California and
Hawaii, spending between 45 and 78 percent of the year on the high seas
(Harrison et al. 2018). Of the 32 nations, the West Pacific DPS
migrates through at least 18 nations or territories of the western and
southwestern Pacific Ocean: Indonesia, Papua New Guinea, Solomon
Islands, Philippines, Malaysia, Vietnam, Japan, Palau, Micronesia,
Marshall Islands, Northern Mariana Islands and Guam, Fiji, Vanuatu,
Australia, New Caledonia, New Zealand, Line Islands, and Kiribati
(Harrison et al. 2018). Foraging occurs in seven ecoregions: South
China/Sulu and Sulawesi Seas, Indonesian Seas, East Australian Current
Extension, Tasman Front, Kuroshio Extension of the Central North
Pacific, equatorial Eastern Pacific, and California Current Extension
(Benson et al. 2011). Individuals demonstrate fidelity to these
foraging areas, likely as a result of their post-hatchling dispersal
patterns and nesting season (Benson et al. 2011; Gaspar et al. 2012;
Gaspar and Lalire 2017; Harrison et al. 2018).
Leatherback turtles of the West Pacific DPS nest in tropical and
subtropical latitudes primarily in Indonesia, Papua New Guinea, and
Solomon Islands, and a lesser extent in Vanuatu (Dutton et al. 2007;
Benson et al. 2007a; Benson et al. 2007b; Benson et al. 2011). The
majority of nesting occurs along the north coast of the Bird's Head
Peninsula, Papua Barat, Indonesia at Jamursba-Medi and Wermon Beaches
(Dutton et al. 2007). A recent discovery of a previously undocumented
nesting area on Buru Island, Maluku Province, Indonesia (WWF 2018)
suggests that additional undocumented nesting habitats may exist on
other remote or infrequently surveyed islands of the western Pacific
Ocean. This DPS nests year round, and exhibits a bimodal nesting
strategy whereby a proportion of females nest during November through
February (i.e., ``winter'' nesting females) and other females nest May
through September (i.e., ``summer'' nesting females; Benson et al.
2007a; Benson et al. 2007b; Dutton et al. 2007; Tapilatu and Tiwari
2007; Benson et al. 2011).
Nesting beach habitats throughout the West Pacific are generally
dynamic, high profile beaches associated with deep water approaches and
strong waves. Beaches can be quite narrow as in parts of the Solomon
Islands or Papua New Guinea, or broad as in the case of Jamursba-Medi,
Indonesia during the summer months. Nesting females appear to prefer
coarse-grained sand free of rocks, coral, or other abrasive substrates
(reviewed by Eckert et al. 2012).
While West Pacific leatherback turtles do not have distinct
``migratory corridors,'' several areas are considered ``areas of
passage'' used by turtles traveling between nesting and foraging
locations, and there is clear separation of migratory and foraging
destinations based on nesting season (Benson et al. 2007a, b; Benson et
al. 2011; Harrison et al. 2018). Post-nesting, winter nesting females
from Papua New Guinea, Indonesia, and Solomon Islands migrate through
the Halmahera, Bismarck, Solomon, and Coral Seas, towards Southern
Hemisphere temperate and tropical foraging areas in the Tasman Sea,
East Australian Current, and western South Pacific Ocean (Benson et al.
2011; Harrison et al. 2018; Jino et al. 2018). Genetic analyses of
leatherback turtles caught in fisheries off Peru and Chile indicates
that approximately 15 percent of sampled individuals originate from the
West Pacific DPS, likely winter nesting females that have migrated
across the Southern Hemisphere to the productive waters off South
America (Donoso and Dutton 2010; NMFS unpublished data 2018). It is
unclear what proportion of the West Pacific DPS might utilize this area
and how important it might be to this DPS.
Leatherback turtles migrate through and forage in the waters of the
Philippines (Benson et al. 2007a, 2011; MRF 2010, 2014). In 2005,
Salinas et al. (2009) found a female in San Fernando (close to El Nido)
that had been previously tagged at Jamursba-Medi in July 2003. The
Marine Research Foundation (MRF) utilized aerial transects to assess
leatherback foraging area use in Palawan waters and off the coast of
Borneo (MRF 2010, 2014). They found leatherback turtles (n = 28 in 2010
and 2013/2014) foraging in nearshore waters around the NE and SE coasts
of Palawan, potentially linked to large jellyfish aggregations from
February to May, and overlapping with high density fishing activity in
Taytay Bay, off NE Palawan (MRF 2010, 2014). Additionally, numerous
leatherback turtle marine sightings, strandings, and fishery bycatch
(typically entangled in gillnet gear) exist for locations throughout
the Philippines including Marine Wildlife Watch of the local NGO,
Marine Wildlife Watch of the Philippines, from 2010 to 2018 (Bagarinao
2011; Cruz 2006; MRF 2010; MWWP unpublished data 2018).
Abundance
The total index of nesting female abundance of the West Pacific DPS
is 1,277 females. We based this total index on two nesting aggregations
in Jamursba-Medi and Wermon, Indonesia (Tapilatu et al. 2013; Tiwari et
al. in prep). Our total index does not include 18 unquantified nesting
aggregations in Indonesia, Papua New Guinea, Solomon Islands, and
Vanuatu. To calculate the index of nesting female abundance (723
females) for Jamursba-Medi (i.e., a 18 km stretch of beach that has
been monitored since 1981), we divided the total number of nests
between the 2015/2016 and 2017/2018 nesting seasons (i.e., a 3-year
remigration interval) by the clutch frequency (5.5 clutches per
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season; Tapilatu et al. 2013). We performed a similar analysis for data
from Wermon (index = 554 females), a 6 km beach that has been monitored
since 2002.
Based on the Tapilatu et al. (2013) study, the IUCN Red List
assessment estimated the total number of mature individuals (including
females and males) utilizing Jamursba-Medi and Wermon beaches to be
1,438 leatherback turtles (Tiwari et al. 2013). The IUCN estimate
includes males and thus is higher than ours. Curtis et al. (2015)
provided a minimum annual nesting female estimate of 318 females (or
954 total nesting female abundance over a 3-year remigration interval).
Dutton et al. (2007) estimated that 1,113 females may have nested
annually, or conservatively 2,700 total nesting females, in the entire
western Pacific population. At that time, they estimated 75 percent of
the population originated from Bird's Head Peninsula (or approximately
2,025 females; Dutton et al. 2007). Our total index is within the range
of published estimates of abundance for this DPS, taking into account
differences in survey methods over time, and is based on the best
available data for the DPS at this time.
Within the nesting range of this DPS, nest monitoring activities
have occurred relatively recently, with standardized methods in Papua
Barat first implemented in 2002 (Hitipeuw et al. 2007; Tapilatu et al.
2013). Outside the Bird's Head Peninsula, monitoring has been sporadic,
opportunistic, and spatially limited because the region is vast,
remote, and logistically challenging to access. Often nesting beaches
are located far from towns or cities, where there are no roads to, or
electricity in, adjacent villages. Cultural and socio-economic dynamics
confound monitoring programs, which are dependent upon fiscal
sponsorship, incentives, community buy-in, and the degree of
familiarity of local communities with concepts of sustainability or
conservation (Kinch 2006; Gjersten and Pakiding 2012). While Jamursba-
Medi and Wermon beaches have been monitored fairly consistently over
time, less is known about the status and trends of nesting beaches in
Papua New Guinea, Solomon Islands, and Vanuatu. Records are further
confounded by changes in place names and jurisdictional boundaries over
recent decades (e.g. the Indonesian province formerly known as Irian
Jaya is currently two provinces of Papua and Papua Barat). Village
names or location descriptions have also changed over time, and
geographic coordinates were not recorded historically. Therefore, all
estimates of abundance in this DPS carry substantial uncertainty.
In Indonesia, aerial surveys provided the first indication of
leatherback nesting in Papua (i.e., Irian Jaya; Salm 1982). At that
time, Salm (1982) did not provide location details out of concern that
public disclosure prior to protection would be detrimental. Follow-up
studies during the 1980s and 1990s indicated that a large nesting
population was located along the coastal beaches of northern Papua or
Papua Barat, Bird's Head Peninsula (Bhaskar 1985). Systematic
monitoring of leatherback turtles began during the early 1990s,
primarily in the form of annual nest counts (Hitipeuw et al. 2007). On
the Bird's Head Peninsula of Papua Barat, nesting occurs mainly at
Jamursba-Medi and Wermon, where a total of 1,371 nesting females were
tagged between 2002 and 2011 (Tapilatu et al. 2013). The primary
nesting season at Jamursba-Medi occurs during the summer (May to
September), while nesting occurs year round at Wermon with a small peak
in July and primary nesting activity during the winter, between
November and February (Hitipeuw et al. 2007). Historically,
approximately 60 percent of nesting activity occurs at Jamursba-Medi
with 40 percent of activity at Wermon (Tapilatu et al. 2013). While a
few females have been documented nesting at both beaches during a
nesting season (Tapilatu et al. 2013), the vast majority of females do
not appear to utilize both Jamursba-Medi and Wermon Beaches during a
single nesting season (Tapilatu and Tiwari 2007; Tapilatu et al. 2013;
Lontoh 2014). Based on nest counts and clutch frequency per season
(mean = 5.5 +/- 1.6 nests per female), approximately 464 to 612 females
nested at Jamursba-Medi and Wermon in 2011 (Tapilatu et al. 2013).
Additional low-level nesting activity in Indonesia occurs in the
Manokawari region of the Bird's Head Peninsula to the east of the
Jamursba-Medi and Wermon Beaches (Suganuma et al. 2012). Between 2008
and 2011, 84 to 135 nests were recorded, or a mean of about 117 nests
annually (Suganuma et al. 2012). However, survey effort was limited and
not consistent across years and may underestimate total nesting
activity. Further it is unknown whether interchange occurs between
turtles nesting in the Manokawari region and those of the Bird's Head
Peninsula index beaches. In 2016, nesting activity was identified in
Central Maluku at Buru Island, west of Bird's Head Peninsula. In 2017,
a monitoring program to quantify nesting activity was initiated on
three north coast beaches of Buru Island (totaling 10 km) which
documented 203 nests, and preliminary data indicates that there might
be two nesting peaks: May through July and November through February
(WWF 2018). Nesting activity in other areas of Indonesia are known or
suspected, but unquantified (Dutton et al. 2007; Tapilatu 2017).
In Papua New Guinea, the majority of known nesting activity occurs
during the winter months (November to February) along the Huon Coast on
the northeastern coast of the Morobe Province, where 576 females were
tagged between 1999 and 2013 (Pilcher 2006, 2008, 2009, 2010, 2011,
2012, 2013; Pilcher and Chaloupka 2013). Aerial surveys along the Huon
Coast in January and December between 2004 and 2006 documented 276
nests, with an estimate of 500 nests per season (Benson et al. 2007b;
Dutton et al. 2007). During the Huon Coast Leatherback Turtle Project,
which took place between 2005 and 2012, an average of 258 nests were
laid per season (range: 193 to 527) at seven beaches which comprised
approximately 35 km of nesting habitat along the Huon Coast (Pilcher
2013; WPRFMC 2015). One challenge in estimating nesting activity in
Papua New Guinea is that leatherback site fidelity appears to be
variable, with some satellite tagged animals seen visiting a number of
areas during one nesting season (Benson et al. 2007b). For example, a
number of Huon Coast nesting females visited other nearby beaches and
east-facing beaches of the Huon Peninsula, including Bougainville and
Woodlark Islands during a single nesting season (Benson et al. 2007b).
Therefore, for assessment purposes, we consider the Huon Coast to be
one nesting beach complex.
Additional nesting activity occurs in other areas of Papua New
Guinea, such as along the north coast of the Madang Province and on
several islands including Manus, Long, New Britain, Bougainville, New
Ireland, and Normanby (Prichard 1982; Spring 1982; Benson et al. 2007b;
Dutton et al. 2007). In these areas nesting activity has not been
quantified via standardized or consistent methods, but information has
been obtained via community surveys, aerial surveys, or rapid
assessments. Nesting occurs primarily in the winter months, although
low-level year-round nesting may also occur (Spring 1982; Dutton et al.
2007). Approximately 50 nests may be laid annually along the north
coast of the Madang Province (Benson et al. 2007b; TIRN 2017). The
Islands of New Britain and Bougainville may host approximately 140 to
160
[[Page 48389]]
nests per year, respectively (Benson et al. 2007b; Dutton et al. 2007;
Kinch et al. 2009). On Bougainville Island, aerial surveys conducted
during the 2005 and 2007 nesting seasons documented a mean of 68 nests
(range: 41 to 107 nests) or an extrapolated estimate of 160 to 415
nests per year (Dutton et al. 2007; Benson et al. 2007b). In 2009, a
one week full-island ground survey (conducted by boat and foot)
recorded 46 leatherback nests (Kinch et al. 2009).
In the Solomon Islands nesting activity is distributed throughout
the country with the majority of nesting activity at Sasakolo and
Litogarhira beaches on Isabel Island, and on Rendova and Tetepare
Islands in the Western Province (Pita 2005; Dutton et al. 2007; Benson
et al. 2018a). The nesting season occurs primarily during winter
(November through February), although some year-round nesting has been
documented (Pilcher 2010b; Williams et al. 2014; Jino et al. 2018; TNC-
Solomon Islands 2018 unpublished). Leatherback turtle monitoring was
begun by the Solomon Island Department of Fisheries in 1989 (Pita
2005). Between 1999 and 2006, an estimated 640 to 700 nests were laid
annually in the Solomon Islands, representing approximately eight
percent of the total western Pacific leatherback nesting at that time
(Dutton et al. 2007). At Sasokolo Beach, Isabel Island, during a 54 day
monitoring period between November 28, 2000 and January 21, 2001, 132
nests were documented with an additional 35 nests present when
monitoring began (Ramohia et al. 2001). Between December 27, 2006 and
January 2, 2007, aerial surveys provided seasonal estimates of 207
nests laid on Isabel Island, and an additional 312 nests on other
islands (Benson et al. 2018a). A January 2011 site visit resulted in
315 nests identified at Sasakolo and Litogahira (Tiwari 2011
unpublished). Recently, nesting activity has also been documented at
the southeastern side of Isabel, where approximately 52 females may
nest annually (TNC-Solomons 2018 unpublished). Since 2002, the Tetepare
Descendants' Association (TDA) has monitored nesting activity
opportunistically in the Solomon Islands, where approximately 30 to 50
leatherback nests are laid seasonally on two beaches (Goby et al.
2010). Between July 1, 2012 and April 30, 2013, TDA undertook 257 beach
surveys and found 44 leatherback nests (TDA 2013). While monitoring
efforts may be ongoing, data management and analysis remains a key
challenge for these isolated communities (TDA 2013; Pilcher 2010b). At
Rendova Island during the 2003/2004 winter nesting season, 235
leatherback turtle nests were recorded, and during the 2009/2010
season, 79 nests were laid (Pilcher 2010b; Goby et al. 2010). Likely
the most comprehensive surveys occurred from September 1, 2012 to April
30, 2013 (91 patrols, 3 days per week), which documented a total of 74
nests (TDA 2013). During the 2017/2018 winter nesting season, 29 nests
were documented (Solomon Islands Community Conservation Partnership
2018 unpublished data). The community on Vangunu Island documented a
total of 23 nests and 11 females between June 2011 and July 2014 (Jino
et al. 2018). Nesting occurred during two distinct seasons from May to
July and from November to January, and of the females tagged, one
nested successfully six times and another nested five times (Jino et
al. 2018). The other nine turtles were only observed nesting once or
twice, and it is likely that either some nesting events were not
recorded or the females nested on surrounding unmonitored beaches (Jino
et al. 2018). On Malaita Island at Waisurione beach, nesting activity
occurs during the summer (June to August), but only a few females were
determined to use the area, with five and seven nests documented in
2014 and 2015, respectively (Williams et al. 2014).
Nesting occurs in low numbers at other islands in the western
Pacific Ocean. In Vanuatu, 30 to 40 nests are laid annually on Epi and
Ambrym Islands (Dutton et al. 2007; Petro et al. 2007; WSB 2011),
although fewer nests (n = 15) were documented during the 2014/2015
nesting season (WSB 2016). Leatherback turtles have been reported in
Fiji (Rupeni et al. 2002; NMFS and USFWS 2013; Jino et al. 2018), but
these accounts involved foraging or in-water capture of animals, and it
is unclear if historic reports included nesting activity (Guinea 1993;
Benson et al. 2013). Historical nesting records also exist for the
eastern coast of Queensland, in New South Wales, and in the Northern
Territories from December to February (Dobbs 2002; Limpus 2009).
However, current information was not available at the time of the
study, and no nests have been observed since 1995 despite regular
monitoring (Flint et al. 2012). Since the 1980s, there have also been
reports of leatherback turtles nesting in the Philippines (Cruz 2006;
MRF 2010). Of recent reports, two documented cases have been confirmed
by sea turtle experts (i.e., staff of the Marine Wildlife Watch of the
Philippines). On July 15, 2013, at Barangay Yawah, Legazpi City, Albay,
NAVFORSOL (the Philippines Naval facility) personnel observed a
leatherback nesting, but the eggs failed to hatch. On August 6, 2013 at
Camp Picardo beach, Barangay, Eastern Samar, a nesting event was
aborted due to disturbance on the beach, but according to the social
media report (i.e., a Facebook post), the female was tagged and led
back to sea (MWWP unpublished 2018). Given the low-site fidelity of the
turtles in this DPS (Benson et al. 2007b), it is not surprising that
leatherbacks might distribute nests among various areas throughout the
region.
The total index of nesting female abundance of the West Pacific DPS
(i.e., 1,277 females) places it at risk for environmental variation,
genetic complications, demographic stochasticity, negative ecological
feedback, and catastrophes (McElhany et al. 2000; NMFS 2017). These
processes, working alone or in concert, place small populations at a
greater extinction risk than large populations, which are better able
to absorb impacts to habitat or losses in individuals. Due to its small
size, the DPS has restricted capacity to buffer such losses. Given the
intrinsic problems of small population size, we conclude that the
nesting female abundance is a major factor in the extinction risk of
this DPS.
Productivity
The West Pacific DPS exhibits a declining nesting trend. We
conducted trend analyses for the two index beaches in Indonesia, which
were the only two beaches with 9 or more recent years of standardized
data, with the most recent data collection in 2014 or more recently
(the standards for conducting a trend analysis in this report). The
median trend in annual nest counts estimated for Jamursba-Medi (data
collected from 2001 to 2017) was -5.7 percent annually (sd = 5.4
percent; 95 percent CI = -16.2 to 5.3 percent; f = 0.867; mean annual
nests = 2,063). While data are available for the period starting in
1999, the best available information indicates that beach monitoring
and nest protection practices improved in 2001; therefore, we used the
time series starting in 2001. For Wermon (data collected from 2006 to
2017, excluding 2002-2005 and 2013-2015 due to low or insufficient
effort), the median trend was -2.3 percent annually (sd = 8.4 percent;
95 percent CI = -19.8 to 14.9 percent; f = 0.643; mean annual nests =
1,010). As Jamursba-Medi and Wermon currently represent approximately
75 percent of nesting for this DPS, we
[[Page 48390]]
consider these declining trends to be representative of the entire DPS.
Our trend data for Indonesia yield similar results to other
published findings. The IUCN Red List assessment found a decreasing
trend of -7 percent annually (Tiwari et al. 2013). Tapilatu et al.
(2013) identified a -5.5 percent annual rate of decline at Jamursba-
Medi between 1984 and 2011 and a -11.6 percent annual rate of decline
at Wermon between 2002 and 2011. Between 1986 and 2010, Benson et al.
(2013) highlighted drastic declines in the annual number of nests at
Jamursba-Medi and Wermon. Additionally, a 27-year aerial survey study
indicates a decline in the number of leatherback turtles foraging off
central California (Benson et al. 2018b). From 1995 to 2003, an
estimated 12 to 379 individuals (mean = 178) foraged in this area
(Benson et al. 2007), while from 2004 to 2017, an estimated 23 to 112
individuals foraged in this area, representing a decline of 5.6 percent
annually (Benson et al. 2018b).
At Jamursba-Medi, nesting data have been collected for some years
since 1981. However, no data were collected during many years in the
mid-1980s and late 1990s (Tapilatu et al. 2013). There is considerable
uncertainty in the early estimates, with over 4,000 nests estimated in
1981, 14,522 nests in 1984, and a dramatic drop to 3,261 nests in 1985
(Tapilatu et al. 2013). It is unclear if there was sampling
inconsistency between years or if there was an actual decline in
nesting activity. However, if analyses are based on the 1984 data,
during which the greatest number of nests was recorded, there was a
78.3 percent decline over the past 27 years (1984 to 2011), or 5.5
percent annual rate of decline (Tapilatu et al. 2013). Alternatively,
if analysis is based on 2005 to 2011 when the Tapilatu et al. (2013)
study ensued, nesting activity declined 29 percent from 2,626 nests (in
2005) to 1,596 nests (in 2011; Tapilatu et al. 2013). Since the
Tapilatu et al. (2013) study, University of Papua scientists have
continued to engage with local communities to monitor nesting activity.
The overall nesting trend has continued to decline by 5.6 percent per
year between 2003 and 2017. However, there appears to be an increase in
nesting since 2013 (Tiwari et al. in prep).
The first comprehensive surveys at Wermon beach in 2002 found
almost as many nests laid on Wermon as on Jamursba-Medi (Hitipeuw et
al. 2007). At that time, it was hypothesized that the decline at
Jamursba-Medi may have been offset by an increase at Wermon (Hitipeuw
et al. 2007). However, Tapilatu et al. (2013) found a significant
decline in nesting at Wermon from 2,994 nests in 2002 to 1,096 nests in
2011 (62.8 percent total or 11.6 percent annual rate of decline).
Unfortunately, no monitoring activities occurred at Wermon between 2013
and 2015 due to community discord, which prevented beach access.
Between 2006 and 2017, nesting has continued to decline at
approximately 2.3 percent (Tiwari et al. in prep). However, there may
have been a slight increase in recent nesting, similar to Jamursba-Medi
(Tiwari et al. in prep).
Local residents stated that leatherback turtles were the dominant
sea turtle species nesting in Maokawari prior to the 1980s, but that
the population has declined significantly since the 1990s due to
village development and exploitation of turtles and eggs (Tapilatu et
al. 2017).
Data collection in Papua New Guinea spanned 8 years and ended prior
to 2014. Because these data did not meet our criteria for ``recent,''
we did not perform a trend analysis, but included a bar graph in the
Status Review Report. In Papua New Guinea, nesting activity along the
Huon Coast was relatively stable between 2005 and 2013, with 193 to 527
nests per year (mean annual nests = 258) and with most nesting activity
occurring at two primary areas, Busama and Kamiali (Pilcher 2013;
Benson et al. 2015; WPRFMC 2015). Given the exchange of females and
evidence of multiple beach use among females in Papua New Guinea
(Benson et al. 2007b), we consider the Huon Coast to be one nesting
area and not individual nesting beaches. Though there have been several
independent studies of abundance over time, we determined that these
data are inadequate to incorporate into a trend analysis because these
data do not meet our criteria (i.e., nest count data consistently
collected in a standardized approach for at least 9 years). For
historical perspective, leatherback turtle nesting along the Huon Coast
was first identified south of the city of Lae near the Buang River, at
an area likely between Labu Tale and Busama villages (i.e., Maus Buang
or Buang-Buassi; Bedding and Lockhart 1989; Quinn and Kojis 1985; Hirth
et al. 1993). Estimates of leatherback turtle nesting at Maus Buang
during the 1980s ranged from five to 10 turtles per night from November
to January (Quinn and Kojis 1985) or 300 nests laid annually (Bedding
and Lockhart 1989). Quinn and Kojis (1985) estimated that 300 to 500
females may nest annually in Papua New Guinea, although it is unclear
if estimates were for the Maus Buang area specifically or the Huon
Coast at large. Hirth et al. (1993) undertook the most standardized
survey at that time and recorded 76 nests and 34 females nesting at
``Piguwa'' (i.e., Maus Buang) on 725 meters of beach during a 15-day
period in December 1989. During the Huon Coast leatherback turtle
nesting beach program, an average of 35 and 114 nests were laid
annually during the 4-month nesting season in this similar area at Labu
Tale and Busama beaches, respectively (Pilcher 2013; WPRFMC 2015).
Kamiali Beach lies approximately 30 km south of the city of Lae. In
1996, the Kamiali Wildlife Management Area was declared a protected
area for leatherback turtles, and the harvest of nests was prohibited
along 2 km of beach. In 1999, village rangers began opportunistic
tagging of nesting females at Kamiali. A community-based nesting beach
monitoring program was established in 2003, which soon grew into the
Huon Coast Leatherback Turtle Conservation Program (Benson et al.
2007b; Pilcher and Chaloupka 2013; Kinch 2006). By 2005, monitoring
activities expanded from Kamiali Beach (approximately 7 km) to seven
beaches encompassing approximately 35 km of nesting beaches, which
included an agreement by participating villages to no longer harvest
eggs (Kinch 2006; Pilcher 2013). Of these seven beaches, Kamiali was
the nesting beach with the longest running, most consistent monitoring
within the Huon Coast nesting beach complex. At Kamiali, 194 females
were tagged between 1999 and 2012, and an average of 77 nests laid per
winter nesting season between 2005/2006 and 2012/2013 (Pilcher 2010,
2011, 2012, 2013; Pilcher and Chaloupka 2013). While we are unable to
interpret an overall trend from these studies, anecdotal reports from
villagers and historic information indicates that leatherback nesting
activity was significantly greater in past decades (Benson et al.
2007b, 2015; Hirth et al. 1993; Kinch 2006; Bellagio Sea Turtle
Conservation Initiative 2008).
In the Solomon Islands, it is not possible to estimate nesting
trends due to non-standardized methods and opportunistic monitoring
efforts over time. Available datasets cannot be compared due to
differences in methodology and do not meet our criteria (i.e., nest
count data consistently collected in a standardized approach for at
least 9 years). Historically, nesting was reported at more than 15
beaches in the Solomon Islands, which may have totaled several hundred
nests per season (McKeown 1977; Vaughan 1981). Currently, nesting
activity occurs
[[Page 48391]]
primarily in eight locations (Pita 2005; Dutton et al. 2007; Benson et
al. 2018a; Jino et al. 2018). However, due to the remoteness of these
areas and lack of systematic surveys, and likely additional
undocumented nesting beaches, additional low numbers of nesting
leatherback turtles are likely to exist in Solomon Islands. For
example, nesting activity was recently identified on Vanugnu Island,
where 23 nests were recorded and 11 females nested between 2011 and
2014 (Jino et al. 2018). Additionally, it is unknown to what extent
females use multiple beaches throughout the Solomon Islands, or those
in Papua New Guinea, and what proportion of females nest in the summer
versus winter (Benson et al. 2007b; Jino et al. 2018; TNC-Solomons 2018
unpublished). While we are unable to interpret an overall trend, local
villagers indicate that leatherback nesting was greater in past decades
(Bellagio Sea Turtle Conservation Initiative 2008; Benson et al. 2007b;
Benson et al. 2015).
In Vanuatu, anecdotal information suggests that nesting has
declined over time (Petro et al. 2007). During the 2010/2011 winter
nesting season, 41 nests were laid at Votlo Beach, Epi Island, and,
during the 2014/2015 nesting season, three females laid 15 nests (WSB
2011, 2016). It is not possible to estimate nest trends due to non-
standardized methods and opportunistic monitoring efforts over time,
which render existing data incomparable and do not meet our criteria
(i.e., nest count data consistently collected in a standardized
approach for at least 9 years).
In addition to an overall declining nest trend, the West Pacific
DPS exhibits low reproductive output (i.e., low hatching success), due
in part to a combination of past and current threats (e.g., beach
erosion, predation, and beach temperatures).
The DPS exhibits low productivity (i.e., low hatching success), and
the overall nest trend is declining, likely due to anthropogenic and
environmental impacts at nesting beaches and in foraging habitats
(Tiwari et al. 2013). We conclude that the declining nest trend and low
reproductive output place the DPS at elevated extinction risk,
especially given the low nesting female abundance.
Spatial Distribution
The West Pacific DPS nests throughout four countries with a broad,
diverse foraging range. It exhibits metapopulation dynamics and fine-
scale population structure.
Aerial surveys conducted between 2004 and 2007 identified
Indonesia, Papua New Guinea and Solomon Islands as the core nesting
areas for the DPS (Benson et al. 2007a; Benson et al. 2007b; Benson et
al. 2011; Benson et al. 2018b). During the nesting season, nesting
females generally stayed within 300 km or less of these nesting
beaches, although a few females were documented visiting multiple
beaches during a nesting season (Benson et al. 2007b). Distributing
nesting activity among various habitats may help to buffer some of the
population from impacts at a single nesting area, but the majority of
females utilize one nesting area during a nesting season (Benson et al.
2011).
Migration and foraging strategies vary based on nesting season,
likely due to prevailing offshore currents and seasonal monsoon-related
effects experienced by the turtles as hatchlings (Gaspar et al. 2012).
The lack of crossover among seasonal nesting populations suggests that
leatherback turtles develop fidelity for specific foraging regions,
likely based on juvenile dispersal patterns (Benson et al. 2011; Gaspar
et al. 2012; Gaspar and Lalire 2017). Oceanic currents help to
structure the spatial and temporal distribution of juveniles and lead
them to foraging and developmental habitats (e.g., the North Pacific
Transition Zone) and to undertake seasonal migrations seeking favorable
oceanic habitats/temperatures and abundant foraging resources, such as
the central California ecoregion (Gaspar and Lalire 2017). Inter-annual
or long-term variability in dispersal patterns can influence population
impacts or resilience to regional or Pacific Ocean perturbations (e.g.,
exposure to fisheries, ENSO events, etc.). Stable isotopes, linked to
particular foraging regions, confirm nesting season fidelity to
specific foraging regions (Seminoff et al. 2012). Size differences are
also apparent, with slightly larger adults appearing to exploit distant
temperate foraging habitats regardless of nesting season (Benson et al.
2011; Lontoh 2014).
Summer nesting females forage in Northern Hemisphere habitats in
Asia and the Central North Pacific Ocean, while winter nesting females
forage in tropical waters of the Southern Hemisphere in the South
Pacific Ocean (Benson et al. 2011; Harrison et al. 2018). This variance
in foraging strategy results in a foraging range that covers much of
the Pacific Ocean: Tasman Sea; East Australian Current; eastern and
western South Pacific Ocean; Indonesian, Sulu and Sulawesi, and South
China Seas; North Pacific Transition Zone; equatorial currents; and
central California ecoregion (Benson et al. 2011; Lontoh 2014; Harrison
et al. 2018; Jino et al. 2018). Different strategies result in
demographic differences within the DPS which may affect productivity
and reproductive output. For example, leatherback turtles that exploit
distant temperate foraging habitats (e.g., central California) may
require multiple years of seasonal foraging before returning to nesting
beaches, due to greater energetic demands. In contrast, leatherback
turtles exploiting geographically closer, year-round prey resources in
more tropical habitats (e.g., Sulu Sulawesi and South China Seas) may
remigrate annually (Lontoh 2014).
The DPS also exhibits genetic population structure. While mtDNA
analyses of 106 samples from Indonesia, Papua New Guinea, and Solomon
Islands did not detect genetic differentiation among nesting
aggregations (Dutton et al. 2007), microsatellite DNA analyses indicate
fine-scale genetic structure (Dutton 2019; NMFS SWFSC unpublished
data).
The wide distribution and variance in foraging strategies likely
buffers the DPS to some degree against local catastrophes or
environmental changes that would limit prey availability. The
distribution of nesting beaches throughout four countries, although
primarily concentrated in three, helps to buffer the entire DPS from
major environmental catastrophes, because disturbances are not likely
to similarly affect all countries during the same seasons.
Additionally, the fine-scale genetic structure among nesting
aggregations is indicative of metapopulation dynamics, which may also
provide the DPS with some resilience.
Diversity
The West Pacific DPS exhibits genetic diversity, with six
haplotypes identified in 106 samples from Solomon Islands, Papua Barat
Indonesia, and Papua New Guinea (Dutton 2006; Dutton et al. 2007;
Dutton and Squires 2008). This may provide the DPS with the raw
material necessary for adapting to long-term environmental changes,
such as cyclic or directional changes in ocean environments due to
natural and human causes (McElhany et al. 2000; NMFS 2017). The
population also exhibits temporal nesting diversity, with various
proportions of the population nesting during different times of the
year (summer versus winter) which helps to increase resilience to
environmental impacts. The foraging strategies are also diverse, with
turtles using seven
[[Page 48392]]
ecoregions of the Pacific Ocean. Diverse foraging strategies likely
provide some resilience against local reductions in prey availability
or catastrophic events, such as oil spills or typhoons, by limiting
exposure from a single event to only a portion of the DPS. We conclude
that diversity within the DPS provides it with some level of resilience
to threats.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
The destruction or modification of habitat is a threat to this DPS.
Primary impacts to nesting beaches include erosion and ocean
inundation, which may be caused by natural processes.
Nesting beaches of the West Pacific DPS are dynamic, high profile
beaches that are subject to erosion, such as during King Tides
(naturally occurring, predictable highest tides), which are common
seasonal occurrences. In Indonesia, the Bird's Head Peninsula beaches
are also subject to seasonal patterns of erosion and accretion. Changes
in the currents brought on by monsoons beginning in September cause
major erosion at Jamursba-Medi that often removes the entire beach,
making the habitat unsuitable for nesting until accretion begins again
in March (Hitipieuw et al. 2007). This natural erosion has been
documented to impact many nests at Jamursba-Medi (Hitipeuw et al.
2007). Arguably, western Pacific leatherbacks have been dealing with
such changes in beach habitats over time, and a turtle's long
reproductive lifespan in general is designed to sustain nest loss
during a few bad years or seasons. For example, during the 2003/2004
nesting season, 80 percent of marked nests at Jamursba-Medi (Warmamedi
beach) washed away before they hatched (Hitipeuw et al. 2007). However,
given the low abundance of the population, the loss (or continued loss
over time) of nests is a concern.
At Wermon, the inundation of nests from high tides is a threat
during the winter months. During the 2008/2009 winter nesting season,
26 percent of nests laid at Wermon were inundated by tidal activity
(Wurlianty and Hitipeuw 2009). During the 2004/2005 nesting season, 23
percent of nests were lost to inundation (Wurlianty and Hitipeuw 2005).
During the 2003/2004 nesting season, 10.7 percent of all nests at
Wermon were below the high water mark and were subsequently washed away
by high tides (Hitipeuw et al. 2007). Tapilatu and Tiwari (2007)
stressed that any management plan developed for Papua will need to
address the impact of inundation and beach erosion.
Beach erosion is also a threat to nests in Papua New Guinea, where
strong storms and tidal surges result in substantial erosion and
changes to beaches throughout the Huon Coast. For example, much of the
Labu Tale nesting beach was lost to erosion during the 2012/2013
nesting season (Pilcher 2013). The differences in beach width along the
Huon Coast place some beaches at more risk of inundation and erosion,
such as Kamiali Beach, which is half the width and significantly
narrower than Busama Beach (Pilcher 2008). At Kamiali, the average
distance of nests to the sea was 3.2 m, compared to 6.2 m at Busama;
the distances to the vegetation line were comparable across sites (1.3
m and 1.7 m, respectively; Pilcher 2013).
In Vanuatu, there has been low hatching success in some years due
to storms, floods, and high water (Petro et al. 2007; WSB 2016).
In recent years, management and conservation practices have
included relocating erosion-prone nests to bolster hatchling
production. However, these projects are funding-dependent throughout
the range of the West Pacific DPS. At Jamursba-Medi, ``doomed'' nests
(i.e., those that are likely to be lost to erosion or inundation) are
sometimes relocated to a more stable section of beach; 15 nests were
relocated during the 2017 summer nesting season (Tiwari et al. in
prep.). At Wermon, nests are relocated to avoid erosion and tidal
inundation, and increasingly due to Ipomea root invasion (Tiwari et al.
in prep), but beach management activities are project-dependent. At
Wermon during the 2017/18 winter nesting season, nests could not be
relocated because of the lack of permission from the beach owners, and
all but three nests washed away (Tiwari et al. in prep). In Papua New
Guinea, 22 of 47 nests (47 percent) at Kamiali beach were relocated to
protect them from storm surge and erosion during the 2011/2012 nesting
season, and 41 percent of nests were relocated during the 2009/2010
season (Pilcher 2012). In the Solomon Islands, efforts to relocate
``doomed'' nests is an ongoing and necessary management strategy to
help bolster hatchling production, given that a large proportion of
nests are inundated or have very low hatching success (Goby et al.
2010; TDA 2013; Jino et al. 2018).
A large, significant portion of nests (i.e., 10.7 percent to nearly
all) are exposed to the reduction and modification of nesting habitat,
as a result of erosion and inundation. This threat impacts the DPS by
reducing nesting and hatching success, which has been documented
throughout the nesting range of the DPS (NMFS and USFWS 2013; Bellagio
Sea Turtle Conservation Initiative 2008). While West Pacific
leatherback turtles have undoubtedly evolved to sustain changes in
beach habitats given their proclivity to select highly dynamic and
typically narrow beach habitats, and therefore at the population level
can sustain some level (albeit unquantified level) of nest loss.
However, the increasing frequency of storms and high water events
perhaps as a result of climate change can result in increased and
perhaps unnatural loss of nests. Such impacts may lower the
productivity of the DPS. Based on the information presented above, we
conclude that habitat loss and modification is a threat to the DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
The primary threat to the West Pacific DPS is the harvest (both
legal and illegal) of leatherback turtles and their eggs. Leatherback
turtles are protected by regulatory mechanisms in all four nations
where the DPS nests, but laws are largely ignored and not consistently
enforced. This is due to the extreme remoteness of beaches, customary
and traditional community-based ownership of natural resources (which
includes sea turtles), and overall lack of institutional capacity and
funding for enforcement. Furthermore, the cultural and socio-economic
dynamics in these nations confound community buy-in and conservation
efforts (Kinch 2006; Gjersten and Pakiding 2012; von Essen et al.
2014). Additionally, there are nuances related to indigenous harvest
(and the definition thereof), some of which is permitted in these
nations.
Turtle poaching affects both nesting females on beaches and turtles
in their foraging habitats (Bellagio Sea Turtle Conservation Initiative
2008; Kinch 2009; Suarez and Starbird 1996; Tiwari et al. 2013; WWF
2018). Turtle poaching has been documented in all four countries where
this DPS nests. Egg poaching is a well-documented threat (past and
current) and is widespread throughout the range of the DPS (Bellagio
Sea Turtle Conservation Initiative 2008; NMFS and USFWS 2013; Tiwari et
al. 2013; Tapilatu et al. 2017).
In Indonesia, the poaching of turtles and eggs continues to occur,
though egg harvest and exploitation of females has been minimized at
Jamursba-Medi and Wermon beaches due to the presence of monitoring
programs and educational outreach. Large-scale egg poaching
[[Page 48393]]
occurred at Jamursba-Medi between 1980 and 1993, whereby approximately
4 to 5 boats per week (from May to August) collected 10,000 to 15,000
eggs per boat (Tapilatu et al. 2013). Commercial egg harvest has been
effectively eliminated since beach monitoring was established at that
beach in 1993 (Hitipeuw et al. 2007). However, recent survey efforts
indicate that most, if not all, sea turtle eggs (including leatherback
turtles) are poached at other Bird's Head Peninsula beaches and sold in
local markets (Tapilatu et al. 2017). At Buru Island, Indonesia,
between 2016 and 2017, eight females were poached (WWF 2018), and over
the past 20+ years, three to five nesting females have likely been
taken annually (J. Wang, NMFS, pers. comm., 2018). In 2017, 114 of 203
leatherback nests were harvested at Buru Island (WWF 2018). In 2018,
due to education provided by the newly established WWF program on Buru
Island, local community-based efforts in four villages now prohibit
female and egg harvest. While protective laws exist in Indonesia,
enforcement is largely lacking in areas where monitoring programs do
not exist.
In Indonesia, foraging leatherback turtles are also harvested in
the waters of the Kei Islands, Maluku Province, where a recognized
indigenous subsistence harvest of immature and adult turtles (average
size 145 to 170 cm; range 52 to 203 cm) occurs and has likely been a
key feature of the local traditional culture for centuries (Compost
1980; Hamman et al. 2006; Hitipeuw and Lawalata 2006, 2008). Within the
Kei Islands, customary law (``hak adat'') authorizes a ritual
leatherback turtle hunt in the nine villages of the traditional kingdom
of the Nufit people. Starbird and Suarez (1994) brought attention to
this hunt when they reported that approximately 200 turtles were
harpooned in three months (October to December) of 1994, with as many
as 13 taken in one day. Over the past three decades, sporadic
monitoring efforts have estimated that up to 100 individuals are
harvested annually (Suarez and Starbird 1996; Hitipeuw and Lawalata
2008; WWF 2018). At one point, it was assumed that harvest pressure had
declined and was no longer an issue (NMFS and USFWS 2013). However,
recent surveys indicate that harvest continues, with conservative
estimates of 431 turtles killed over an 8-year period (an average of
53.9 turtles annually), typically between August to February (Hitipeuw
and Lawalata 2008), and at least 103 turtles harvested in 2017 (WWF
2018). Most concerning perhaps is that some of the turtle meat
harvested may be commercially sold as dried meat (i.e., leatherback
``jerky'' locally known as dendeng), which is illegal to sell and
inconsistent with indigenous traditional practices. Of four genetic
samples acquired in 1995 from turtles harvested in the Kei Islands,
three were assigned to Birds Head Indonesian region and the fourth
sample was not definitive (66 percent probability, with 34 percent
probability to Solomon Islands), although it could also be from the
Indian Ocean or from an undetermined location (NMFS SWFSC unpublished
data 2018).
In Papua New Guinea, turtle and egg poaching is a major threat
despite the fact that leatherback turtles have been protected since the
1976 Fauna (Protection and Control) Act. The illegal take of both eggs
and turtles likely continues throughout the country due to lack of
community-based awareness, reliance on traditional community-based
practices, institutional capacity, and law enforcement (Bellagio Sea
Turtle Conservation Initiative, 2008). The killing of nesting females
has also been well documented throughout Papua New Guinea (Bellagio Sea
Turtle Conservation Initiative 2008; Kinch 2009; Pilcher 2013). For
example, at Bougainville Island, surveys of community members
identified that 21 nesting females were poached during the last decade
(Kinch 2009). However, the harvest of eggs is likely the most prolific
threat in Papua New Guinea. If unprotected, egg harvest (compounded by
intense dog predation described below) resulted in the loss of 70 to
100 percent of nests (Quinn and Kojis 1985; Hirth 1993; Bellagio Sea
Turtle Conservation Initiative 2008; Pilcher 2013). For example, during
a one week survey in January 2009 at Bougainville Island, almost 100
percent of the 46 documented nests were poached (Kinch 2009). It is
likely that near total egg collection occurred throughout the Huon
Coast between World War II and the establishment of the Huon Coast
Leatherback Turtle Monitoring and Conservation Program in 2003
(Bellagio Sea Turtle Conservation Initiative 2008; Pilcher and
Chaloupka 2013; Pilcher 2013). The Huon Coast Project, which operated
between 2003 and 2013, helped to reduce egg and turtle harvest due to
program involvement and community incentive funds received in exchange
for non-harvest agreements (Pilcher 2013). As a result of the program,
hatchling production (i.e., percent of eggs yielding hatchlings)
increased from zero to approximately 60 percent (Pilcher 2009, 2011,
2013; WPRFMC 2015). The Project ended in 2013, and unfortunately egg
harvest resumed since there was no incentive for communities to
maintain their no-harvest agreements (John Ben, Huon Coast Leatherback
Turtle Project, pers. comm., 2020).
In Vanuatu and the Solomon Islands, the poaching of females and
collection of eggs is also well documented (Bellagio Sea Turtle
Conservation Initiative 2008; NMFS and USFWS 2013). In Vanuatu, MacKay
et al. (2014) reported the harvest of five nesting females between 1999
and 2008. However there is a general understanding that nesting females
were typically harvested (Petro et al. 2007). Of the 315 nests
documented on Isabel Island, Solomon Islands during a January 2011 site
visit at Sasokolo and Litogahira beaches, the majority of nests had
been poached (Tiwari 2011 unpublished data). Historically, nearly all
nesting females and eggs were poached on Redova for consumption (Tiwari
2011 unpublished data). In response, financial incentive programs have
been established to protect nests and females whereby villagers are
paid a financial reward for each nest that hatches successfully (TDA
2013). On Vangunu Island, 10 to 20 nesting females were poached
annually, in addition to near-total egg collection (Jino et al. 2018).
In response to declining population trends, the community declared a
moratorium on the harvest of leatherback turtles in 1999 (Jino et al.
2018), and a community incentive program providing financial awards has
helped to reduce harvest pressure (TDA 2013). Despite these efforts and
protective legislation, the poaching of females and eggs likely
persists throughout the Solomon Islands (TDA 2013: Tiwari 2011
unpublished; MacKay et al. 2014).
Within the West Pacific DPS, many nesting females, foraging
turtles, and eggs are exposed to both illegal poaching and legal
harvest. The taking of turtles reduces abundance. The taking of nesting
females reduces both abundance and productivity. Such impacts are high
because they directly remove the most productive individuals from the
DPS, reducing current and/or future reproductive potential. Egg harvest
reduces productivity; the persistent, and near-total (at some
locations) collection of eggs guarantees that future population
recruitment (i.e., nesting female abundance) will be reduced or
eliminated. Given the declining nesting trend and current nesting
female abundance of this DPS, the continued and unregulated poaching
[[Page 48394]]
or harvest of leatherback turtles and eggs is unsustainable. Further,
the harvest of approximately 100 foraging leatherback turtles annually
at the Kei Islands, Indonesia is likely an unsustainable practice given
the current low abundance of the population. We conclude that
overutilization is a major, and the primary, threat to the West Pacific
DPS, accelerating its risk of extinction.
Disease or Predation
While we could not find any information on disease for this DPS,
predation of eggs is a major and well-documented threat to the West
Pacific DPS, likely second to poaching (i.e., nests not taken by humans
are typically predated; Bellagio Sea Turtle Conservation Initiative
2008).
In Indonesia, predation of eggs by feral pigs, feral dogs, and
monitor lizards has been documented, with feral pig predation being the
most detrimental (Hitipeuw and Maturbongs 2002; Tapilatu and Tiwari
2007; Bellagio Sea Turtle Conservation Initiative 2008). Nest predation
by domestic and/or feral dogs has been recorded in both Jamursba-Medi
and Wermon. Predation of nesting females by crocodiles has also been
documented at Wermon beach (Bellagio Sea Turtle Conservation Initiative
2008; UNIPA, pers. comm., 2018). At Jamursba-Medi, between June and
July of 2005, 29.3 percent of nests were destroyed by pigs (Tapilatu
and Tiwari 2007). Intensive management effort at Jamursba-Medi reduced
feral pig predation of nests to five percent during the 2016 and 2017
nesting seasons (Tiwari et al. in prep). Feral pigs and dogs depredated
17.5 percent of all nests at Wermon during the 2003 and 2004 winter
nesting season (Hitipeuw et al. 2007). At Wermon, 21 percent of nests
were lost to predation during the 2004/2005 nesting season (Wurlianty
and Hitipeuw 2005). At Buru Island in 2017, 16 nests were lost to
predation by dogs, wild boar, lizards, or saltwater crocodiles (WWF
2018).
In Papua New Guinea, predators of eggs include feral dogs, monitor
lizards, and ghost crabs (Kinch 2009). Depredation of nests by village
dogs was determined to be an intense threat to nests, with dogs
consuming all nests laid during the 2003/2004 and 2004/2005 nesting
seasons at Kamiali beach (Pilcher 2006; I. Kelly, NMFS, pers. comm.,
2018). Predation of nesting females by crocodiles has also been
documented in a number of locations in Papua New Guinea (Bellagio Sea
Turtle Conservation Initiative 2008; Kinch 2009). To protect nests,
Huon Coast communities developed and placed bamboo grids over nests to
prevent dogs from preying on the eggs (Pilcher 2006; 2009). This, along
with efforts to reduce egg harvest by humans, resulted in increased
hatching production from zero to approximately 60 percent between 2006
and 2013, with over 2,300 nests saved producing approximately 100,000
hatchlings (Pilcher 2009; 2011; 2013; WRFMC 2015). However, this
project ended in 2013, and it is unknown if egg protection continues,
or if nest predation has resumed.
In this DPS, a large proportion of eggs are exposed to predation,
especially by dogs and pigs. Predation primarily results in the loss of
eggs, and the impact of this threat is a reduction of productivity.
Though leatherback turtles generally produce a large number of eggs and
hatchlings, predation is widespread throughout the range of the DPS,
and in some areas, predation rates are as high as 100 percent. We
conclude that predation poses a threat to the West Pacific DPS.
Inadequacy of Existing Regulatory Mechanisms
The West Pacific DPS is protected by several regulatory mechanisms.
For each, we review the objectives of the regulation and to what extent
it adequately addresses the targeted threat.
Leatherback turtles are protected by legislation in all four of the
nations where the West Pacific DPS nests (Indonesia, Papua New Guinea,
Solomon Islands, and Vanuatu). It is generally illegal to harvest
leatherback turtles and their eggs. However, laws are not typically
enforced or followed given customary marine tenure systems that dictate
near-shore rights. Lack of enforcement or implementation of protective
laws may be due to: Overall lack of in-country institutional capacity
and funding for enforcement; the extreme remoteness of beaches;
customary marine tenure or traditional community-based ownership of
natural resources in these nations (which includes sea turtles; Kinch
2006; McDonald 2006) and regulatory government-led legislation, which
may be incompatible with traditional practices (von Essen et al. 2014).
There are also nuances related to indigenous harvest (and the
definition thereof), which is not prohibited in these nations. As a
result, most leatherback nesting beaches with the exception of
Jamursba-Medi and Wermon (i.e., beaches with established long-term
monitoring programs) are not currently protected (or only minimally
protected) from harvest or poaching of eggs, nesting females, or other
anthropogenic threats.
In Indonesia, all sea turtles are protected by law, but there are
allowances for indigenous peoples (although indigenous provisions are
not clearly defined). The 1990 Government Regulation Act number 5
concerning the Conservation of the Natural Resources and the Ecosystem,
makes the trade of protected wildlife illegal, and those found liable
can be punished to a maximum of 5-year prison term and fined 100
million Indonesia Rupiah (approximately 6,500 USD). The protection of
all sea turtle species (Government Regulation No. 7 on Preserving Flora
and Fauna Species) came into effect in 1999 (Zainudin et al. 2007). The
use of protected wildlife is allowed for the purposes of research,
science, and rescue of the wildlife itself. While the trade and
exploitation of turtles is illegal in Indonesia, there still exists a
documented harvest of green turtles in Bali, which contributes to
public confusion regarding sea turtle protections (Westerlaken 2016).
In Papua New Guinea, the leatherback turtle is the only species
protected under the 1976 Fauna (Protection and Control) Act, which
makes killing of leatherback turtles or taking of leatherback turtle
eggs illegal, with fines of 500 to 1000 kina (approximately 100 to 300
USD). Any person who buys or sells or offers for sale, or has in
possession leatherback turtle eggs or meat can also be fined. The Act
makes provisions for persons with customary rights to take turtles, but
states that sea turtles cannot be taken, killed, or sold from May
through July (Kinch 2006). This is typically the nesting season for
hard-shelled sea turtle species, but leatherback turtles nest primarily
during the winter months (November to February). As with most
Melanesian countries, lands are locally-owned and managed, and the
national government has little influence outside major cities (Kinch
2006).
The Solomon Islands Fisheries Act (1993) regulations protect
nesting turtles and eggs during the breeding season (June to August and
November to January); prohibit the sale, purchase, or export of sea
turtle species or their parts; and contain specific protections for
leatherback turtles. In the Solomon Islands, more than 85 percent of
the land is held under customary (locally-managed) marine tenure, and
the vast majority of the population still lives in rural areas making a
living from the natural resources on those lands. For centuries,
communities have practiced traditional models of resource stewardship,
making implementation and enforcement of national regulations nearly
impossible. Instead, natural
[[Page 48395]]
resource governance must originate from chiefs and village leaders,
which requires extensive educational outreach to encourage traditional
approaches that may be supported by legal or `modern' enforcement
measures (McDonald 2006).
Fisheries Regulations under the Vanuatu Fisheries Act (2009)
prohibit the take, harm, capture, disturbance, possession, sale,
purchase of or interference with any turtle nest (or any turtle in the
process of nesting) and the import, or export of green, hawksbill, and
leatherback turtles or their products (shell, eggs, or hatchlings). The
Act also prohibits the possession of turtles in captivity. A person may
apply in writing to the Director of Fisheries for an exemption from all
or any of these provisions for the purposes of carrying out customary
practices, education, and/or research. Similar to Papua New Guinea and
the Solomon Islands, natural resource governance in Vanuatu is best
directed, realized, and implemented at the community level and not via
national legislation. Fortunately, traditional practices are
experiencing a renaissance in Vanuatu and may complement current
regulatory marine resource management efforts (Hickey et al. 2006).
Throughout the foraging range of the DPS, there are numerous
regulatory mechanisms that protect turtles within the DPS. These
include: RFMOs such as the Western and Central Pacific Fisheries
Commission (WCPFC) and the IATTC and fisheries management regulations
in 32 nations where this DPS may occur (Harrison et al. 2018). The
WCPFC adopted a Sea Turtle Conservation and Management Measure (CMM
2008-03) to mitigate the impacts on turtles from commercial shallow-set
fisheries operating in the Western and Central Pacific Ocean. The
measure included the adoption of FAO (2009) guidelines to reduce sea
turtle mortality through safe handling practices and to reduce bycatch
by implementing one of three methods by January 2010. The three methods
to choose from are: (1) Use only large circle hooks with offsets of
<=10[deg]; (2) use whole finfish bait; or (3) use any other mitigation
plan or activity that has been approved by the Commission. This sea
turtle conservation measure is specific to self-identified shallow-
setting, swordfish-targeting fleets. It does not apply to the
international Pacific longline deep-set tuna-targeting fisheries, which
comprise the majority of the longline fisheries and are also known to
interact with leatherback turtles (Lewison et al. 2004; Beverly and
Chapman 2007; Roe et al. 2014; Wallace et al. 2013). Technical analysis
of the sea turtle conservation measure found a very small percentage of
shallow-set fisheries to be in compliance, with less than one percent
of Western and Central Pacific Ocean longline effort implementing
mitigation measures, even though approximately 20 percent of longline
effort consists of shallow sets (Clarke 2017). Further, many RFMO
members are not meeting the five percent observer coverage requirement
resulting in limited bycatch reporting (Clarke 2017).
In summary, regulatory mechanisms exist to protect leatherback
turtles and their eggs throughout the range of this DPS. However, most
are inadequate to reduce the threat that they were designed to address
due to a lack of implementation or enforcement or inclusion of
provisions for indigenous harvest. Regulations are also misaligned with
established traditional practice and management systems. As a result,
poaching and bycatch remain major threats to the DPS. In summary, we
consider the inadequacy of the regulatory mechanisms to be a threat to
the DPS.
Fisheries Bycatch
Fishery bycatch in coastal and pelagic fisheries is a major threat
to the West Pacific DPS, which is exposed to domestic and international
fisheries throughout its extensive foraging range. At-sea bycatch of
leatherback turtles has been documented for a variety of gillnet and
longline fisheries in the Pacific Ocean, but little is known about the
total magnitude or full geographic extent of mortality. Satellite
telemetry studies have identified movements and revealed fidelity to
foraging regions of the DPS, specifically in habitats of the North
Pacific Ocean, southwestern Pacific Ocean, and Indo-Pacific tropical
seas (Bailey et al. 2012; Benson et al. 2011, Seminoff et al. 2012; Roe
et al. 2014). The summer nesting component of the population exhibits
strong site fidelity to the central California foraging area (Benson et
al. 2011) which puts them at risk during migrations of interacting with
U.S. and international pelagic longline fleets operating throughout the
Central and North Pacific Oceans. For example, several of the turtles
tagged in Papua Barat, Indonesia were known or suspected to have been
killed in fisheries operating off Japan, Philippines, and Malaysia
(Benson et al. 2011).
Historically, significant leatherback bycatch was documented in the
North Pacific high seas driftnet fishery, which expanded rapidly during
the late 1970s but was banned in 1992 by a UN resolution (summarized in
Benson et al. 2015). Wetherall et al. (1993) estimated that over 750
leatherback turtles were killed in Japanese, Korean, and Taiwanese
driftnet fisheries during the 1990 to 1991 season, with potentially
5,000 to 10,000 leatherback turtles bycaught between the late 1970s and
1992. Based on current knowledge of movement patterns (Benson et al.
2011), the majority of these bycaught turtles would have originated
from western Pacific nesting beaches after their boreal summer nesting
period. Thus, high seas driftnet fishery bycatch was likely a
significant contributor to the population declines observed at nesting
beaches during the 1980s and 1990s (Benson et al. 2015).
Many nations are involved in longline fishing in the Pacific Ocean,
where two types of vessels are used: (1) Large distant-water freezer
vessels that undertake long (months) voyages and operate over large
areas of the region; and (2) smaller offshore vessels with ice or chill
capacity that typically undertake trips of about one month. Target
species are yellowfin, bigeye, albacore tuna, and swordfish. The total
annual number of longline vessels in the western and central Pacific
region has fluctuated between 3,000 and 6,000 for the last 30 years,
including the 100 to 140 vessels in the Hawaii longline fisheries (NMFS
2018).
Pelagic Fisheries
International longline fisheries are characterized by inconsistent
reporting and traditional gear configurations, including J-style hooks
with squid bait, which result in higher interaction and mortality rates
than for modified gear (Beverly and Chapman 2007; Lewison et al. 2004;
Swimmer et al. 2017). For example, the Taiwan and China tuna longline
fisheries are estimated to have bycatch rates several times higher than
Hawaii longline fisheries (Bartram and Kaneko 2008; Chan and Pan 2012).
Analyzing multi-national turtle bycatch data from 1990 to 2004, Molony
(2005) found that the purse seine fishery and the deep, shallow, and
albacore longline fisheries (operating between 15[deg] N and 31[deg] S)
take an average of about 100 leatherback turtles annually. Lewison et
al. (2004) collected fish catch data from 40 nations and turtle bycatch
data from 13 international observer programs to estimate global
longline bycatch of loggerhead and leatherback turtles in 2000. In the
Pacific Ocean, they estimated 1,000 to 3,200 leatherback turtle
(juvenile and adult) mortalities from pelagic longlining in 2000
(Lewison et al. 2004). Using effort data from Lewison et al. (2004) and
bycatch data from Molony (2005), Beverly and
[[Page 48396]]
Chapman (2007) estimated sea turtle longline bycatch to be
approximately 20 percent of that estimated by Lewison et al. (2004),
approximately 200 to 640 leatherback turtles annually. These estimates
include turtles from the East and West Pacific DPS. While the results
of each of these studies may be feasible, the Lewison et al. 2004
estimates were based on available data at that time (i.e., less than 30
percent of longline fishing effort) that was skewed toward fishing
fleets with relatively better management and data reporting systems,
and hence extrapolations may have overestimated interaction rates
(Clarke et. al. 2014). However, Beverly and Chapman (2007) applied
different catch per unit effort (CPUE) estimates in calculations
differentiated between deep-set and shallow-set fisheries which have
different interaction rates and, hence, their estimates may be more
realistic.
Despite scientific evidence showing that use of circle hooks and
finfish bait significantly reduces leatherback turtle bycatch rates in
longline fisheries (Gilman et al. 2007; Swimmer et al. 2017), nations
are not required to use this hook/bait combination. The WCPFC Sea
Turtle Conservation and Management Measure (CMM 2008-03) only applies
to fleets using shallow-set gear targeting swordfish. Additionally,
observer program coverage levels in WCPFC longline fisheries have not
reached the required five percent coverage rate, resulting in limited
bycatch reporting and likely underreporting (Clarke 2017). Further,
existing sea turtle mitigation measures are currently only being
applied to approximately one percent of shallow-set longline fisheries
in the Convention Area, even though approximately 20 percent of the
longline effort consists of shallow-sets (Clarke 2017).
A workshop convened to assess the effectiveness of WCPFC's Sea
Turtle Conservation and Management Measure found limited reductions in
interactions and mortalities (Clarke 2017). Fishery observer data
collected between 1989 and 2015 of 34 purse seine and longline fleets
across the Pacific documented a total of 2,323 sea turtle interactions,
of which 331 were leatherback turtles (Clarke 2017). Two bycatch
hotspot areas were identified: One in central North Pacific (which
likely reflects the 100 percent observer coverage in the Hawaii
shallow-set longline fishery) and a second hotspot in eastern Australia
(Clarke 2017). However, analysis of the data also found that overall
conservation benefits would have been greater had mitigation measures
also been applied to deep-set gear and not only to shallow-set
swordfish fisheries (Clarke 2017).
While bycatch in pelagic shallow-set swordfish-targeting longline
fisheries has received the most attention to date, comparable studies
for deep-set tuna-targeting fisheries are not available due to the more
complex nature of these fisheries. There may be fewer interactions
because deep-set fisheries (operating at depths more than 60 m)
generally have lower bycatch rates, but they also have higher mortality
rates than shallow-set gear (Lewison et al. 2004; Kaplan 2005; Gilman
et al. 2007). Pelagic deep-set tuna-targeting fisheries cannot be
ignored because they also have the potential to interact with
leatherback turtles and constitute four times greater effort than
shallow-set fisheries yet do not have RFMO gear mitigation requirements
(Clarke 2017).
Wallace et al. (2013), and a global review based on that study (FAO
2014), categorized longline and gillnet fisheries interactions with
West Pacific leatherback turtles as high risk but low impact for
longline and gillnet gear, likely due to insufficient data from this
data-poor region. Bycatch in small-scale coastal fisheries has been a
significant contributor to population declines in many regions (Kaplan
2005; Peckham et al. 2007; Alfaro-Shigueto et al. 2011), yet there is a
significant lack of information from coastal and small-scale fisheries,
especially from the Indian Ocean and Southeast Asian region (Lewison et
al. 2014).
Southeast Asian Fisheries
Waters of Southeast Asia are heavily fished by a variety of
gillnets, trawls, fish traps, and a range of different hook and line
gears, involving hundreds of thousands of fishers (FAO 2011). The West
Pacific DPS nests, migrates, and forages throughout this densely
populated and heavily exploited coastal region (Bellagio Sea Turtle
Conservation Initiative 2008; Benson et al. 2011; Lewison et al. 2014;
Roe et al. 2014; Harrison et al. 2018).
There are few quantitative estimates of fisheries interactions near
nesting beaches of this DPS, and existing reports provide only brief
snapshots of impacts or are outdated. In Indonesia, between 1980 and
1993, shark gillnets off the nesting beaches of Jamursba-Medi killed
two to three nesting females weekly (Tapilatu et al. 2013). As a member
of the WCPFC and the IOTC, Indonesia must comply with reporting
requirements and conservation measures as required by these RFMOs. In
2006, of the 85 sea turtle interactions observed in 539 sets on 10 tuna
longline vessels, 3 were adult leatherback turtles (Zainudin et al.
2007). Leatherback turtles are known to migrate through and forage
within Philippine waters (Benson et al. 2011), and in 2014, aerial
surveys observed leatherback turtles foraging in high density fishing
areas (130 to 381 boats; MRF 2010, 2014). Leatherback turtles have also
stranded dead or injured on Philippine beaches as a result of fishery
interactions, typically with gillnet gear (Bagarinao 2011; Cruz 2006;
MRF 2010; MWWP 2018 unpublished). In Malaysia, bycatch studies using an
interview-based approach revealed that four leatherback turtles were
caught in gillnets the prior year (Pilcher et al. 2008).
Fisheries operating out of Australia and New Zealand may result in
high bycatch and mortality rates for the winter nesting component of
the DPS that migrates into the Southern Hemisphere (MacKay et al. 2014;
Harrison et al. 2018). In Australia, some bycatch records exist for
pelagic longline fisheries (Robins et al. 2002; Stobutzki et al. 2006),
prawn trawls off Queensland and Northern Territory, gillnet fisheries
off Queensland and Tasmania, and pot gear off Tasmania (Limpus 2009).
Gillnet sea turtle bycatch is reported as widespread and includes
anecdotal reports of leatherback turtles taken in Tasmanian tuna
gillnet fisheries (Limpus 2009).
Between 2004 and 2014, the Australian shallow-set fishery had an
estimated 29 to 178 leatherback interactions, based on two to 10
observations (average = 4.6 interactions) and four to 10 percent
observer coverage (MacKay et al. 2014). These data are similar to
bycatch information extrapolated from interviews with Australian
fishers (Robins et al. 2002) which identified 162 leatherback turtles
interactions in 2001 (MacKay et al. 2014). Australia has a sea turtle
mitigation plan for its Eastern Tuna and Billfish Fishery which sets
``trigger level'' interaction rates of <=0.0048 turtles per 1,000 hooks
for each turtle species or 0.0172 turtles per 1,000 hooks overall (DAFF
2009 in Clarke et al. 2014). In 2013, Australia reported that the
trigger levels had been exceeded for the third year in a row and as a
consequence the Australian Fisheries Management Authority required that
shallow-set vessels in these fisheries use large circle hooks
consistent with the WCPFC sea turtle measure (CMM 2008-03; Clarke et
al. 2014).
In New Zealand, records document 288 instances of stranding or
commercial and recreational bycatch of leatherback turtles from 1892 to
2015 (Godoy et al. 2016). New Zealand's surface longline fishery
captured 90
[[Page 48397]]
leatherback turtles between 2008 and 2015 (Godoy et al. 2016). This is
likely an underestimate because data were based on low observer
coverage (5.8 percent overall), with limited observer overage during
the peak time of leatherback abundance in New Zealand waters (January
to March). Strandings can also provide opportunities for researchers to
identify fisheries interactions. MacKay et al. (2014) identified 19
mortalities in New Zealand and 29 mortalities in Australia. Although
the cause of most strandings was often unknown, leatherback turtles
have been found entangled in crab pot gear and monofilament fishing
nets and ropes. Longline fishing is concentrated off southern
Queensland and New South Wales, Australia and is the suspected cause of
41 percent of strandings (n = 12). In Victoria, Tasmania and South
Australia, 61 percent of strandings (n = 17) involved suspected
entanglement in inshore fishing gear and crab pots (MacKay et al.
2014).
U.S. Pacific Pelagic Fisheries
Detailed bycatch data are available for U.S.-managed pelagic
fisheries operating in the central and eastern Pacific Ocean due to
regulatory mandates and high levels of observer coverage. Longline
fisheries, based out of Hawaii and American Samoa, may interact with
foraging turtles of the West Pacific DPS. However, only two
interactions involved individuals of the East Pacific DPS in 1995 and
2011 (P. Dutton, NMFS, pers. comm., 2018). Prior to 2001, the Hawaii
longline fishery was estimated to capture about 110 leatherback turtles
annually, resulting in approximately 9 annual mortalities (McCracken
2000). Since 2005, the fishery has reduced its estimated mortality to
seven leatherback turtles annually, and data confidence increased
significantly due to increased observer coverage (NMFS 2018). The
fishery was closed in 2001 under court order and re-opened in 2004 as
two separate fisheries: A shallow-set swordfish-targeting fishery and a
deep-set tuna-targeting fishery. Management requirements include: Gear
modification (e.g., circle hooks and fin-fish bait) and handling
measures designed to reduce sea turtle bycatch rates and post-hooking
mortality in both fisheries; an annual hard-cap limit on the number of
allowable interactions in the shallow-set fishery; 100 percent observer
coverage in the shallow-set fishery; and 20 percent observer coverage
in the deep-set fishery (50 CFR 665 (Subparts A-C); NMFS 2012, 2014,
2015). The shallow-set fishery has been closed three additional times
since reopening in 2004: In 2006, after reaching the hard cap for
loggerhead turtle interactions (n = 17); in 2011, after reaching the
hard cap for leatherback turtle interactions (n = 16); and in 2018
under a stipulated settlement after the Ninth Circuit Court of Appeals
held that NMFS' no jeopardy determination for loggerheads in the 2012
biological opinion (9th Circuit 2017) was arbitrary and capricious. See
Turtle Island Restoration Network v. U.S. Dep't. of Commerce, 878 F.3d
725 (9th Cir. 2017). Since 2004, leatherback turtle interactions in the
shallow-set component of the fishery have been reduced by 84 percent
from 0.03 to 0.01 BPUE as a result fisheries regulations (Swimmer et
al. 2017). Between 2004 and 2017, there have been 99 total leatherback
turtle interactions in the shallow-set fishery (or approximately 8
turtles annually), based on 100 percent observer coverage (WPRFMC
2018). Between 2002 and 2016, an estimated 168 interactions may have
occurred in the Hawaii deep-set fishery (or approximately 12 annually),
based on an extrapolation of data collected at a level of 20 percent
observer coverage (WPRFMC 2018). Observer coverage of the American
Samoa longline fishery has varied over time from 5 to 40 percent and
has had an estimated 59 interactions between 2010 and 2017 (WPRFMC
2018).
The U.S. tuna purse seine fishery operating in the Western and
Central Pacific Ocean anticipates up to 11 leatherback turtle
interactions annually (NMFS 2006). However, the fishery had fewer
interactions, with approximately 16 leatherback turtle interactions
between 2008 and 2015 based on observer coverage ranging from 20 to 100
percent (NMFS unpublished data).
From 1990 to 2009, there were 24 observed leatherback turtle
interactions in the California drift gillnet fishery based on 15.6
percent per year observer coverage (Martin et al. 2015). Genetic
analyses indicated that almost all originated from the West Pacific DPS
(Dutton et al. 1999; NMFS SWFSC unpublished). In 2001, NMFS implemented
regulations (i.e., a large time/area closure in Central California)
that reduced interactions by approximately 80 to 90 percent, with only
two leatherback turtle interactions (both alive) observed based on 20
to 30 percent observer coverage since regulations were implemented
(NMFS West Coast Region unpublished). Drift gillnet fishing is
prohibited annually from August 15 to November 15 within the California
leatherback turtle conservation area. Currently, NMFS anticipates up to
10 interactions (or 7 mortalities) over a 5-year period (NMFS 2013).
In addition, nine fixed gear fisheries operate off the U.S. West
Coast, including the Federally-managed sablefish pot fishery and the
state-managed California Dungeness crab fishery. Since 2008, only one
leatherback interaction has been documented in the sablefish fishery
(NMFS 2013). The state-managed Dungeness crab fishery may be a newly
emerging threat: Two documented leatherback entanglements in pot gear
(mainline or surface buoy) occurred in 2015 and 2016. Fishing effort
was high, and the fishery had shifted into the Central California
region, which overlaps somewhat with leatherback foraging habitat (S.
Benson, NMFS, pers. comm., 2018). In 2019, the State of California
settled with a non-profit organization in response to a complaint that
the commercial Dungeness crab fishery was taking leatherback sea
turtles (and other large whales) without authorization under section 10
of the ESA. The California Dungeness crab fishery closed in mid-April
2019 as part of the settlement agreement and again on May 15, 2020
(just the Central Management Area), due to significant risk of marine
life entanglement. The northern part of California remains open until
mid-July unless CDFW decides to take further management action (i.e.,
if risks to large whales and/or leatherbacks is elevated in that area).
East Pacific Pelagic Fisheries
The West Pacific DPS has a vast trans-Pacific range. Some
individuals forage in the East Pacific Ocean, where leatherback turtles
are caught in fisheries of Peru and Chile (Donoso and Dutton 2010;
Alfaro-Shigueto et al. 2007, 2011, 2018). Of 59 leatherback turtles
caught in East Pacific fisheries, an estimated 15 percent of
individuals sampled originated from the West Pacific DPS (Dutton et al.
2000; Donoso and Dutton 2010). Information compiled by IATTC on sea
turtle interactions with pelagic longline fisheries operating in the
East Pacific is limited, given that requirements for longline observer
coverage of five percent was only implemented in January 2013 (Clarke
et al. 2014). Additional information on East Pacific fisheries are
presented in the bycatch section for the East Pacific DPS.
Summary of Fisheries Bycatch
We conclude that individuals of this DPS are exposed to high
fishing effort throughout their foraging range, in coastal waters near
nesting beaches, and when migrating to and from nesting
[[Page 48398]]
beaches, though very little fisheries data are available for coastal
areas. Bycatch rates in international pelagic and coastal fisheries are
high, and these fisheries have limited management regulations despite
hotspots of high interactions in Southeast Asia (Lewison et al. 2004,
2014; Alfaro-Shigueto et al. 2011; Wallace et al. 2013; Clarke 2017).
Annual interaction and mortality estimates are only available for U.S.-
managed pelagic fisheries, which operate under extensive fisheries
regulations that are designed to minimize the capture and mortality of
endangered and threatened sea turtles (NMFS 2013; Swimmer et al. 2017;
NMFS 2018). Mortality reduces abundance, by removing individuals from
the population; it also reduces productivity, when nesting females are
killed. We conclude that fisheries bycatch is a major threat to the
West Pacific DPS.
Vessel Strikes
Vessel strikes are a threat to the West Pacific DPS. Between 1981
and 2016, there were 11 documented vessel strikes in central California
(NMFS West Coast Region, unpublished data 2018). Many vessel strikes
are not reported, and turtles are not recovered.
The range of the DPS overlaps with many high-density vessel traffic
areas. Though the potential for exposure is high, we are only aware of
11 vessel strikes in recent decades. Vessel strikes resulting in
mortality would lower the abundance of the DPS. However, available data
does not support characterizing this as a high or moderate impact. We
conclude that vessel strikes pose a threat to the DPS, albeit of less
concern than other impacts such as overutilization and fisheries
interactions.
Pollution
Pollution includes contaminants, marine debris, and ghost fishing
gear. Leatherback turtles can ingest small debris, causing internal
damage and blockage. Larger debris can entangle animals, leading to
reduced mobility, starvation, and death. Given the amount of floating
debris in the Pacific Ocean (Lebreton et al. 2018), marine debris has
the potential to be a significant threat to the DPS. Presently
available data do not allow for quantifying the precise extent of the
threat.
Leatherback turtles feed exclusively on jellyfish and other
gelatinous organisms and as a result may be prone to ingesting plastics
resembling their food source (Schuyler et al. 2013). Lebreton et al.
(2018) estimated plastic debris accumulation to be at least 79,000
(45,000 to 129,000) tonnes in the Great Pacific Garbage Patch, a 1.6
million km\2\ of subtropical waters between California and Hawaii. This
figure is four to 16 times greater than previously reported.
Entanglement in ghost fishing gear is also a concern (Gilman et al.
2016), and derelict nets made up approximately 46 percent by piece, and
86 percent by weight, of debris floating in this area (Lebreton et al.
2018). The highest risk areas within the range of the West Pacific DPS
where animals may encounter significant amounts of debris includes the
north Pacific gyre, the South China Sea, and off of the east coast of
Australia (Schuler et al. 2015). However, Wedemeyer-Strombel et al.
(2015) found no plastics in the gastrointestinal tracts of two
leatherback carcasses from American Samoan and Hawaiian longline
fisheries from 1993 to 2011. Clukey et al. (2017) found no plastics in
the gastrointestinal tracts of three leatherback carcasses from Pacific
longline fisheries captured between 2012 and 2016. However, it is very
difficult to obtain dead leatherback turtles to study these effects,
and given the great amount of plastics within environment, such results
may underestimate ingestion impacts.
Few studies of pollutants and their effect on leatherback turtles
were available within the range of this DPS. Harris et al. (2011) found
the heavy metal exposure in leatherback turtles foraging off the coast
of California to be nine times higher than the St. Croix nesting
population, although levels were not expected to be lethal. We do not
know if there were sub-lethal effects. Stewart et al. (2011) found that
PCBs are more likely to be transferred from females to their eggs than
from the environment to eggs.
Given the large amount of marine debris within the range of the
DPS, we expect exposure to be high for all life stages despite low
sample sizes of leatherback turtles with ingested marine debris.
Potential impacts include death and injury. However, quantitative
estimates of such impacts are not available. We conclude that pollution
may be a threat to the DPS.
Natural Disasters
The best available scientific and commercial data indicate that
natural disasters are a threat to the DPS but do not allow the impact
to be quantified. Natural disasters within the range of this DPS
include: Tsunamis, typhoons, earthquakes, and flash floods. Such
environmental events are periodic, with localized impacts that do not
persist over time. These events may reduce nest incubation and hatching
success in one season or at few locations. While leatherback turtles
have undoubtedly evolved to sustain such natural impacts, the
increasing frequency of environmental events as a result of a changing
climate, which can affect the frequency and intensity of high tides and
large storms, may hamper productivity and conservation activities (Goby
et al. 2010; S. Benson, NMFS, pers. comm., 2018). Such events may pose
additional threats by depositing marine debris on nesting beaches and
in occupied waters. The 2011 Japan tsunami and the 2006 Indonesian
earthquake and resulting tsunami likely deposited large amounts of
debris (i.e., millions of tons) into the foraging and migrating
habitats of the DPS (Hafner et al. 2014; NOAA 2015). We conclude that
natural disasters pose a potential threat to the West Pacific DPS.
Climate Change
Climate change is a threat to the West Pacific DPS. A warming
climate and rising sea levels can impact leatherback turtles through
changes in beach morphology, increased sand temperatures leading to a
greater incidence of lethal incubation temperatures, changes in
hatchling sex ratios, and the loss of nests or nesting habitat due to
beach erosion (Benson et al. 2015).
Elevated egg incubation temperatures can lead to mortality. During
the 2009/2010 nesting season at the Huon Coast (Papua New Guinea),
Pilcher (2010) found higher incubation temperatures (32 to 33 [deg]C)
in exposed nests compared to shaded nests (29 to 30 [deg]C). Sea
turtles exhibit temperature-dependent sex determination. The incubation
temperature determines sex ratios and the duration of incubation (i.e.,
thermosensitive period). Along the Huon Coast, incubation duration
decreased during the nesting season as beach temperatures warmed.
During the 2006/2007 nesting season, nests laid in November hatched in
61.8 4.2 days, and nests laid in February hatched in 55.8
3.4 days (n = 171 nests; Steckenreuter et al. 2010).
Assuming that hatchlings were male at temperatures less than 29.2
[deg]C and female at temperatures greater than 30.5 [deg]C,
Steckenreuter et al. (2010) estimated that only 7.7 percent of the
hatchlings were female, indicating a highly male-skewed sex ratio.
However, given the Pilcher (2010) results, sex ratios are likely
variable over time and space.
Climatic change may also alter rainfall levels, which may cool
beaches and offset increases in sand temperature. At Wermon, the sand
is black, yet beach temperatures are lower, perhaps because
[[Page 48399]]
peak nesting coincides with the monsoon season (Tapilatu and Tiwari
2007). Sand temperatures fluctuate between 28.6 and 34.9 [deg]C at
Jamursba-Medi and between 27.0 and 32.7 [deg]C at Wermon (Tapilatu and
Tiwari 2007). Hatching success of nests undisturbed by feral pig
predation was significantly lower in Jamursba-Medi (25.5 percent) than
Wermon (47.1 percent). Although there was significant variation between
beaches, Tapilatu and Tiwari (2007) concluded that high sand
temperatures may exceed the thermal tolerance of leatherback embryos,
resulting in high embryo mortality and low hatching success at
Jamursba-Medi. Further, Tapilatu and Tiwari (2007) concluded that high
average sand temperatures may suggest a female-biased population at
Jamursba-Medi. However, the mean incubation period of 61.5
4.7 days (Tapilatu and Tiwari 2007) was similar to the length of
incubation recorded in Papua New Guinea during the cooler November
period, which Steckenreuter et al. (2010) suggested produced a male-
biased sex ratio.
Tapilatu et al. (2013b) found that the daily average sand
temperatures during the boreal summer (from 2005 to 2012) ranged from
26.5 to 34.9 [deg]C, suggesting the production of female-biased sex
ratios and potentially lower hatching success. Further, histological
examination of dead hatchlings from both summer and winter nesting
seasons from 2009 to 2019 produced a female-biased sex ratio, which is
consistent with the relatively warm thermal profiles of the nesting
beaches (Tapilatu et al. 2013b). Additional impacts of climate change
include increased sea level rise and storm frequency, resulting in
greater nest inundation and beach erosion. As sea level rises, King
Tides are likely to have a greater effect on nests. Climate change may
also affect prey availability. Saba et al. (2007, 2012) identified a
correlation between the reproductive frequency of the East Pacific DPS
and ENSO events. Because the West DPS also forages in the East Pacific
Ocean, it too may be exposed to variability in productivity.
The threat of climate change is likely to modify the nesting and
foraging conditions for turtles of the DPS. Impacts are likely to
affect productivity. Negative impacts and low hatching success due to
high beach temperatures and coastal erosion have already been
documented and are likely to become worse, and thus we conclude that
climate change is a threat to the West Pacific DPS.
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply to turtles of the West Pacific DPS
(for a description of each effort, please see the section on
conservation efforts for the overall species): Convention on the
Conservation of Migratory Species of Wild Animals, Convention on
Biological Diversity, Convention on International Trade in Endangered
Species of Wild Fauna and Flora, Convention for the Protection of the
Marine Environment and Coastal Area of the South-East Pacific (Lima
Convention), Convention for the Conservation and Management of Highly
Migratory Fish Stocks in the Western and Central Pacific Ocean (WCPF
Convention), Convention for the Protection of the Natural Resources and
Environment of the South Pacific Region, Convention Concerning the
Protection of the World Cultural and Natural Heritage (World Heritage
Convention), Eastern Pacific Leatherback Network, Eastern Tropical
Pacific Marine Corridor Initiative, FAO Technical Consultation on Sea
Turtle-Fishery Interactions, IAC, MARPOL, IUCN, The Memorandum of
Understanding of a Tri-National Partnership between the Government of
the Republic of Indonesia, the Independent State of Papua New Guinea
and the Government of Solomon Islands, Ramsar Convention on Wetlands,
RFMOs, Secretariat of the Pacific Regional Environment Programme,
UNCLOS, and UN Resolution 44/225 on Large-Scale Pelagic Driftnet
Fishing. Although numerous conservation efforts apply to the turtles of
this DPS, they do not adequately reduce its risk of this DPS, they do
not adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the West Pacific DPS is at high risk of extinction. The DPS
exhibits a total index of nesting female abundance of 1,277 females at
two currently monitored beaches over the most recent remigration
interval. These beaches may represent 75 percent of total DPS nesting
activity. This abundance makes the DPS vulnerable to stochastic or
catastrophic events that increase its extinction risk. This DPS
exhibits low hatching success and decreasing nest and population trends
due to past and current threats, which are likely to further lower
abundance and increase the risk of extinction. The DPS exhibits genetic
diversity and metapopulation structure, with nesting aggregations
distributed throughout four nations. Nesting occurs during two seasons
(winter and summer), with year-round nesting at some locations and uses
multiple foraging areas, throughout the Pacific Ocean. Thus, the DPS
has some resilience to stochastic events and environmental
perturbations at nesting beaches and foraging areas. However, its
abundance and declining trends place the DPS at risk of extinction as a
result of past threats.
Current threats also contribute to the risk of extinction of this
DPS. The overutilization of turtles and eggs, as a result of legal and
illegal harvest, is the primary threat to this DPS, reducing abundance
and productivity. Abundance and productivity are further reduced by
fisheries bycatch. Juvenile and adult turtles are taken by numerous,
international, coastal, and pelagic fisheries throughout the extensive,
pan-Pacific foraging range of the DPS. Predation (especially by dogs
and pigs) reduces productivity at high rates. Erosion and inundation
result in habitat loss and modification that reduces productivity and
contributes to low hatching success. Additional threats include:
Pollution, vessel strikes, and natural disasters. Climate change is an
increasing threat that results in reduced productivity. Though many
regulatory mechanisms exist, they do not adequately reduce threats.
We conclude, consistent with the team's findings, that the West
Pacific DPS is at risk of extinction. Its nesting female abundance
makes the DPS highly vulnerable to threats. The declining nesting trend
further contributes to its risk of extinction. While the DPS has
spatial structure and diversity, the resilience provided by those
factors is likely to be eroded by the reduced and declining abundance.
Past egg and turtle harvest reduced the abundance and productivity of
this DPS and remains a primary threat. Fisheries bycatch is also a
primary threat that reduces abundance by removing mature and immature
individuals from the population. Predation is also a major threat to
productivity. Though numerous conservation efforts apply to this DPS,
they do not adequately reduce the risk of extinction. We conclude that
the West Pacific DPS is in danger of extinction throughout its range
and therefore meets the definition of an endangered species. The
threatened species definition does not apply because the DPS is
currently in danger of extinction (i.e., at present), rather than on a
trajectory to become so within the foreseeable future.
[[Page 48400]]
East Pacific DPS
The Team defined the East Pacific DPS as leatherback turtles
originating from the East Pacific Ocean, north of 47[deg] S, south of
32.531[deg] N, east of 117.124[deg] W, and west of the Americas. In the
south, the cold waters of the Antarctic Circumpolar Current likely
restrict the nesting range of this DPS. We placed the northern and
western boundaries at the border between the United States and Mexico
because this DPS forages primarily in the East Pacific Ocean, off the
coasts of Central and South America.
The range of the DPS (i.e., all documented areas of occurrence) is
centered in the eastern Pacific Ocean but may include distant waters
for foraging, as demonstrated by a turtle satellite-tracked to waters
off the Tonga Trench and a turtle captured by the Hawaii longline
fishery, genetically assigned to the population we refer to in this
finding as the East Pacific DPS (P. Dutton, NMFS, pers. comm., 2018).
Records indicate that the DPS occurs in the waters of the following
nations: Chile; Colombia; Costa Rica; Ecuador; El Salvador; France
(Clipperton Island); Guatemala, Honduras; Mexico; Nicaragua; Panama;
Peru; and the United States (Hawaiian Islands) (Wallace et al. 2013).
Leatherback turtles of the East Pacific DPS nest primarily on
beaches in Mexico, Costa Rica, and Nicaragua. In Mexico, where the
largest nesting aggregations occur, nesting beaches are found in 11
states, over 7,828 kilometers as far north as Baja California Sur
(Sarti 2002). The following beaches in Mexico host approximately 40 to
50 percent of total nesting for the nation: Mexiquillo
(Michoac[aacute]n), Tierra Colorada (Guerrero), and Cahuit[aacute]n,
Chacahua, and Barra de la Cruz (Oaxaca; Gaona Pineda and
Barrag[aacute]n Rocha 2016). In Costa Rica, approximately 75 percent of
nesting occurs within the Parque Nacional Marino Las Baulas (Guanacaste
Province) at three nesting beaches: Playa Ventanas; Playa Grande; and
Playa Langosta (based on recent abundance estimates from 2011-2015;
Santidri[aacute]n Tomillo et al. 2017). In Nicaragua, small numbers of
leatherback turtles nest on Playa Salamina-Costa Grande and Veracruz de
Acayo (Chacocente Wildlife Refuge) (FFI 2018). Rare nesting events have
been documented in Guatemala (n = 6), El Salvador (n = 4), and Panama
(n = 4), with none in Honduras (Sarti et al. 1999).
Generally, the nesting season starts in October and ends in March
(Santidri[aacute]n Tomillo et al. 2007; Eckert et al. 2012). Nesting is
generally bound between 10[deg] N and 20[deg] N, falling within the
northeast corner of the Intertropical Convergence Zone. The nesting
beaches share similarly warm temperatures, moderate annual rainfall,
and seasonal dynamics (Saba et al. 2012). In general, nesting beach
habitat for leatherback turtles is associated with deep water and
strong waves and oceanic currents, but shallow water with mud banks are
also used by leatherback turtles. Beaches with coarse-grained sand and
free of rocks, coral, or other abrasive substrates also appear to be
selected by leatherback turtles (reviewed by Eckert et al. 2012).
Foraging areas of the East Pacific DPS include coastal and pelagic
waters of the southeastern Pacific Ocean. Leatherback turtles are
widely dispersed on the high seas throughout the eastern Pacific Ocean
(Shillinger et al. 2008). They also forage in coastal areas off the
coast of Peru and Chile (Alfaro-Shigueto et al. 2007; Eckert 1997;
Donoso and Dutton 2010). Using satellite telemetry, Morreale et al.
(1996) tracked the movements of eight post-nesting females and
identified a persistent southbound migration corridor from Las Baulas
National Park toward the Galapagos Islands. Eckert (1997) found a
similar pattern, tracking seven post-nesting females from Mexiquillo in
a similar direction; while three continued to the same foraging habitat
as the Costa Rican nesting females, four shifted their movements away
from the South American coast, when a strong El Ni[ntilde]o caused a
warm water anomaly. Additional tracking of 46 post-nesting females from
Las Baulas National Park over a 3-year period (2004/2005 to 2006/2007)
confirmed the persistent migratory corridor (Shillinger et al. 2008).
The turtles navigated the equatorial current system, south to around
5[deg] S latitude and negotiated the strong alternating eastward-
westward flows of the equatorial current, swimming predominantly in a
southward direction and moving rapidly through the productive
equatorial region. They then dispersed throughout the South Pacific
Gyre ecosystem, which is characterized by low phytoplanktonic biomass.
The South Pacific Gyre contains ample mesoplankton forage base, as
demonstrated by tuna longline fisheries effort in the eastern tropical
Pacific Ocean (Shillinger et al. 2008). Of the 46 turtles, only one
leatherback moved into coastal foraging areas, which had been
documented earlier by Eckert (1997). During the course of the tracking
duration, this female occupied nearshore foraging habitats along the
coast of Central America, which represents highly productive areas when
compared with oceanic areas. Researchers have hypothesized that high
bycatch along the coastal areas of Central and South America could have
extirpated a coastal migratory phenotype in this population (Saba et
al. 2007). Recently, Harrison et al. (2018) determined that post-
nesting females from Las Baulas National Park spent 78.2 percent of
their time on the high seas, 17.8 percent of their time in Costa Rica's
EEZ, and 3.7 percent of their time around the Galapagos Islands.
In summary, preferred foraging areas for the East Pacific DPS are
characterized by low sea surface temperatures and high mesoscale
variability. Post-nesting females migrate relatively quickly through
areas that contain the strong equatorial currents as well as high
chlorophyll-a concentrations, likely because of the strong currents.
While swimming speed was significantly higher in areas of high
chlorophyll levels, the association between these two variables was
weak (Shillinger et al. 2008). Once past this area, they appear to
forage in the southern part of their range in the South Pacific
Subtropical Convergence, where there is a sharp gradient in primary
production. In this area, Ekman upwelling may accelerate the transport
of nutrients and consequently increase prey availability. Seasonally,
leatherback turtles from the East Pacific DPS foraged at higher
southerly latitudes during the austral summer (November to February),
which may reflect seasonal patterns in prey abundance during higher
latitudes (Bailey et al. 2012).
Abundance
The total index of nesting female abundance for the East Pacific
DPS is 755 females. We based this total index on 13 nesting
aggregations in: Mexico (Mexican Commission for Natural Protected
Areas; L. Sarti, CONANP, pers. comm. 2018); Costa Rica
(Santidri[aacute]n Tomillo et al. 2017; Leatherback Trust 2018); and
Nicaragua (FFI 2018). Our total index does not include several
unquantified nesting aggregations in Mexico, Costa Rica, and Nicaragua.
To calculate the index of nesting female abundance for nesting beaches
in Mexico (i.e., 572 females), we added the total number of nesting
females between the 2013/2014 and 2016/2017 nesting seasons (i.e., a 4-
year remigration interval; L. Sarti, CONANP, pers. comm., 2018) at each
beach. We performed a similar calculation for Costa Rica (n = 165
females). To
[[Page 48401]]
calculate the index of nesting female abundance in Nicaragua (i.e., 20
females), we divided the total number of nests between the 2014/2015
and 2017/2018 nesting seasons (i.e., a 4-year remigration interval;
Santradi[aacute]n Tomillo et al. 2007) by the clutch frequency (7.2
clutches/season; Santradi[aacute]n Tomillo et al. 2007).
This number represents an index of nesting females for this DPS
because it only includes available data from recently and consistently
monitored nesting beaches. While rare or sporadic nesting may occur on
other beaches, consistent and standardized monitoring only occurs at
these beaches, which are for the most part protected.
Our total index of nesting female abundance is similar to published
abundance estimates for this DPS. The IUCN Red List assessment
estimated the total number of mature individuals (males and females) at
633 turtles, based first on dividing the average annual number of nests
(n = 926) by the estimated clutch frequency (n = 7.2, Reina et al.
2002) to obtain an average annual number of nesting females. This value
was then multiplied by the average remigration interval (n = 3.7 years,
Reina et al. 2002; Santidri[aacute]n Tomillo et al. 2007) to obtain a
total number of adult females that included nesting as well as non-
nesting turtles. In order to account for adult males, the authors
assumed that the sex ratio of hatchlings produced on nesting beaches in
the East Pacific (approximately 75 percent female, or 3:1 female:male
ratio) reflected the natural adult sex ratio (Wallace et al. 2013). A
more recent analysis of primary sex ratios that included multiple years
of data and considered hatching success (i.e., lower in hot nests)
estimated primary sex ratios at Playa Grande, Costa Rica as
approximately 85 percent female (Santidri[aacute]n Tomillo et al.
2014). In Mexico, the female to male ratio is closer to 1.1:1 (A.
Barragan, Kutzari, pers. comm., 2019).
In Mexico, the beaches included in our total index represent
approximately 70 to 75 percent of total nesting in that nation (Gaona
Pineda and Barragan Rocha 2016). However, our total index does not
include nesting females from Agua Blanca (40 km in Baja California);
Playa Ventura (6 km), Playa San Valent[iacute]n (21 km), Piedra de
Tlacoyunque (44 km in Guerrero), and La Tuza (16 km in Oaxaca) (Sarti
et al. 2007). These beaches are not regularly monitored for nesting,
which is thought to be rare or of low abundance (L. Sarti, CONANP,
pers. comm., 2018).
In Costa Rica, 75 percent of nesting occurred at Las Baulas
National Park (summarized in Santidri[aacute]n Tomillo et al. 2017),
although the recent nesting at other beaches may lower this percentage.
These beaches include: Naranjo, Cabuyal, Nombre de Jes[uacute]s,
Ostional, and Caletas. The longest data set was provided for Naranjo,
which has been intermittently covered from 1971 to 2015. Limited
nesting has been documented at Playa Coyote and at Playa Caletas, which
is a high energy eight kilometer beach located on the Nicoya Peninsula
(Squires 1999). Given the lack of nesting events for Caletas in recent
years, it may no longer host leatherback nesting, despite the fact that
the Playa Caletas/Ario National Wildlife Refuge was created in 2004 to
protect leatherback turtles (Gaos et al. 2008).
In Nicaragua, leatherback turtles nest at three beaches. Salamina
Costa Grande and Veracruz de Acayo (in the Rio Escalante Chacocente
Wildlife Refuge) host the most nesting and have been subject to the
most consistent monitoring. Small numbers of females also nest at Juan
Venado National Reserve, which is not consistently monitored (V. Gadea,
FFI, personal communication, 2018).
Nesting is rare in other nations (Sarti et al. 1999). Nesting is
very uncommon in Ecuador with one record of a female attempting to nest
(according to local reports) in Atacames, a province of Esmeraldas
(Salas 1981). Sarti et al. (1999) reported six nests at Playa Puntilla,
El Salvador, but overall nesting is low and/or unknown throughout the
nation. In Guatemala, nesting is rare, with reports by Sarti et al.
(1999) recording only eight nests during an entire season, and more
recently, zero to six nests per year along the Pacific coast of
Guatemala (Muccio and Flores 2015). Past nesting sites included Hawai
beach, La Candelaria, Taxico, Santa Rosa, and the zone adjacent to the
border with El Salvador, as reported by Chac[oacute]n-Chaverri (2004).
Although nesting has been documented at Barqueta National Refuge,
little is known about nesting in Panama (Chac[oacute]n-Chaverri 2004).
Our total index of nesting female abundance (755 females) places
the DPS at risk for environmental variation, genetic complications,
demographic stochasticity, negative ecological feedback, and
catastrophes (McElhany et al. 2000; NMFS 2017). These processes,
working alone or in concert, place small populations at a greater
extinction risk than large populations, which are better able to absorb
losses in individuals. Due to its small size, the DPS has relatively
little capacity to buffer such losses. Historical abundance estimates
were much greater (e.g., 75,000 leatherback nesting females estimated
in Pacific Mexico from a 1980 aerial survey ((Pritchard 1982). However,
this estimate was derived from a brief aerial survey and may have been
an overestimate (Pritchard 1996)), indicating that this population at
one time had the capacity for a much larger nesting population.
Therefore, the current nesting female abundance is likely an indicator
of past and current threats, and given the intrinsic problems of small
population size, elevates the extinction risk of this DPS.
Productivity
The East Pacific DPS exhibits a decreasing nest trend since
monitoring began, with a 97.4 percent decline since the 1980s or 1990s,
depending on the nesting beach (Wallace et al. 2013). Despite intense
conservation efforts, the decline in nesting had not been reversed as
of 2011 (Benson et al. 2015). We found a declining nest trend at some
of the remaining, small nesting aggregations. Abundance at Las Baulas,
Costa Rica (previously the single largest nesting aggregation) at its
peak was seven times the current abundance at Playa Barra de la Cruz/
Playa Grande, Mexico (currently the largest nesting aggregation). From
1988/1989 to 2015/2016, the number of nesting females at Las Baulas
declined -15.5 percent annually (sd = 3.8 percent; 95 percent CI = -
23.1 to -7.8 percent; f = 0.998; mean annual nests = 315).
In recent decades (after a historical decline), nest counts have
increased at some beaches in Mexico. The Playa Tierra Colorada nest
trend has increased by 0.6 percent annually (sd = 8.9 percent; 95
percent CI = -17.1 to 18.9 percent; f = 0.536; mean annual nests = 153)
between the 1996/1997 and 2016/2017 nesting seasons. Over the same time
period, nesting at Playa Barra de la Cruz/Playa Grande increased by 9.5
percent annually (sd = 8.0 percent; 95 percent CI = -6.5 to 25.8
percent; f = 0.918; mean annual nests = 122). In contrast, nest counts
at Cahuit[aacute]n decreased from 1997/1998 through 2016/2017, with a
median trend of -4.3 percent annually (sd = 9.7 percent; 95 percent CI
= -22.1 to 17.6 percent; f = 0.716; mean annual nests = 123).
We lack adequate data on nesting in Nicaragua to estimate trends.
Our trend analysis yields similar results to other published
findings. The IUCN Red List assessment concluded that this
subpopulation is decreasing and has declined by -97.4 percent over the
past three generations (Wallace et al. 2013). The number of nests at
Mexico nesting beaches has declined precipitously in recent decades
(Benson et al. 2013). Historically, Mexico hosted
[[Page 48402]]
the largest leatherback turtle nesting aggregation in the world, with
75,000 nesting females estimated during an aerial survey in 1980
((Pritchard 1982). However, this estimate was derived from a brief
aerial survey and may have been an overestimate (Pritchard 1996)).
Prior to that aerial survey, Marquez et al. (1981) reported that the
nesting beach of San Juan Chacahua (Oaxaca) was the most important
nesting site in Mexico, with approximately 2,000 females nesting each
season. Researchers also identified Tierra Colorada and Mexiquillo as
important nesting sites, with approximately 3,000 to 5,000 nests per
season. Monitoring of the nesting assemblage at Mexiquillo has been
continuous since 1982. During the mid-1980s, more than 5,000 nests per
season were documented along 4 km of this nesting beach. By 1993, less
than 100 nests were counted along the entire 18 km beach (Sarti 2002).
According to Sarti et al. (1996), nesting declined at this location at
an annual rate of over 22 percent from 1984 to 1995. Researchers from
the National University of Mexico recorded 3,000 to 5,000 nests
annually from 1982 to 1989 at primary nesting beaches, with sharp
declines observed in 1993 to 1994 at the nesting sites at Mexiquillo,
Tierra Colorada, Chacahua and Barra de la Cruz. These early reports
were generally snapshots (e.g., local unpublished data) of leatherback
nesting activity in Mexico, until 1995, when a more coordinated
conservation effort took shape in the form of complete nesting surveys
for the entire Pacific coast of Mexico (Eckert 1997). In 1995,
``Proyecto Laud'' (Leatherback Project) was formed to estimate the
population size using comprehensive surveys. In 1995 and 1996, Proyecto
Laud estimated approximately 1,100 females nesting throughout Mexico;
the next two seasons, they estimated between 236 and 250 nesting
females, and declines continued. Currently, based on data from 2014
through 2018 (preliminary) between 100 and 250 females nest at all the
protected beaches in Mexico.
In Costa Rica, the number of nesting females per season declined
from 1,367 females in 1988 to 117 females in 1998 (Spotila 2000). While
there were increases in the number of nesting females during the 1999/
2000 season (224 females) and 2000/2001 season (397 females), the
population has shown a steady decline, with less than 30 nesting
females in recent years (i.e., through 2016; The Leatherback Trust
2018).
In Nicaragua, 108 leatherback turtles nested on Playa Chacocente
from October to December, 1980; in January 1981, 100 turtles nested in
a single night on Playa El Mogote (Arauz 2002). An aerial survey of
Playa El Mogote during the 1998/1999 nesting season revealed a nesting
density of 0.72 turtles per kilometer (Sarti et al. 1999 in Arauz
2002). During the 2000/2001 nesting season, community members near
Playa El Mogote reported that 210 leatherback nests had been deposited.
That number decreased to 29 nests during the 2001/2002 nesting season
(Arauz 2002). At Playa Veracruz 48 nesting females were identified
between 2002 and 2010 (Urteaga et al. 2012). Between 2002 and 2014,
Salazar et al. (2019) recorded 340 nests, indicating a downward trend.
Considering the best available data, nesting has declined in Nicaragua.
Nesting females of the East Pacific DPS are generally smaller and
produce fewer eggs per clutch than turtles from other leatherback
populations (Sarti et al. 2007; Piedra et al. 2007; Santidri[aacute]n
Tomillo et al. 2007). For example in Mexico, nesting females have a
mean size of 144 cm CCL and 62 eggs per clutch; the average total
fecundity per females was estimated to be 341 eggs per season, with a
maximum of 744 eggs deposited in a season (Sarti et al. 2007). The low
productivity parameters, drastic reductions in overall nesting female
abundance, and current declines in nesting place the DPS at risk of
extinction, especially given the limited nesting female abundance.
Spatial Distribution
The DPS is characterized by somewhat continuous and low density
nesting across long stretches of beaches along the coast of Mexico and
Central America. Santidri[aacute]n Tomillo et al. (2017) found a
contraction of the Costa Rica's overall nesting distribution since the
1990s.
The best available genetic data indicate a high degree of
connectivity among nesting aggregations. Dutton et al. (1999) did not
find any genetic differentiation between nesting populations in Mexico
(Playa Mexiquillo) and Costa Rica (Playa Grande) based on analysis of
mtDNA control region sequences. Additional analyses of mtDNA sequences
and nuclear DNA (microsatellites) from three index nesting beaches in
Mexico also failed to find genetic differentiation (Barragan and Dutton
2000; Dutton et al. unpublished).
Based on monitoring of tagged nesting females, researchers
documented female interchange between nesting beaches within Mexico and
within Costa Rica. However, only one interchange has been documented
between Mexico and Costa Rica (Sarti et al. 2007). Interchange between
nesting beaches may occur during or between nesting seasons and may
depend on the distance between nesting sites, which can be fairly
large, especially in Mexico. For example, the distance between Tierra
Colorada and Cahuit[aacute]n is 25 kilometers, and up to 18.7 percent
of nesting females visit both beaches within a season (average of nine
percent). Mexiquillo is located approximately 475 kilometers from the
closest other nesting beach (Tierra Colorada), and researchers found no
interchange of females within seasons. However, a few females were
found to nest in either Mexiquillo and/or Tierra Colorado between
seasons (Sarti et al. 2007).
In Costa Rica, nesting females move among the three nesting beaches
of Las Baulas National Park, within and between seasons, particularly
between Playa Grande and Playa Langosta, although researchers study
both Playa Grande and Playa Ventanas in combination. According to data
gathered over 10 years of research (mid 1990s through the mid-2000s),
an average of 71 percent of females nested only on Playa Grande, 10
percent nested only on Playa Langosta, and 18 percent nested on both
beaches in a given season. In other seasons, females have been shown to
shift and nest primarily on a different beach. Within two seasons, 82
percent of nesting females at Playa Langosta also nested at Playa
Grande and 100 percent of nesting females at Playa Langosta within
three seasons occasionally also nested at Playa Grande
(Santidri[aacute]n Tomillo et al. 2007). At the less abundant nesting
beaches in Costa Rica, the exchange rate between females ranged between
7 and 28 percent. For example, at Ostional, 12 out of the 43 identified
females were observed at least once at other sites (28 percent), while
at Naranjo, 4 out of 21 identified females were also observed at other
beaches (19 percent). At Cabuyal, 2 out of 15 turtles were observed at
other beaches (13 percent), while 1 out of 15 females at Caletas were
observed elsewhere (7 percent) (Santidri[aacute]n Tomillo et al. 2017).
The foraging range of the DPS extends into coastal and pelagic
waters of the southeastern Pacific Ocean. Individuals forage in the
Pacific Gyre ecosystem and along the coasts of Peru and Chile, with
variation resulting from the location of upwelling and ENSO effects.
Researchers have hypothesized that high bycatch along the coastal
foraging phenotype in this population (Saba et al. 2007). Recently,
Harrison et al. (2018) determined that post-nesting females from Las
Baulas National Park spent 78.2 percent of their time on the
[[Page 48403]]
high seas, 17.8 percent of their time in Costa Rica's EEZ, and 3.7
percent of their time around the Galapagos Islands.
Multiple nesting and foraging distributions likely help to buffer
the DPS against local catastrophes or environmental changes that would
otherwise modify nesting habitat or limit prey availability. Nesting
aggregations are largely connected. However, there is less exchange
among distant nesting beaches. Foraging turtles are vulnerable to
perturbations in ocean conditions due to climate change, ENSO, and the
Pacific Decadal Oscillation.
Diversity
The East Pacific DPS exhibits genetic diversity, as demonstrated by
moderate to high mtDNA haplotypic diversity (h = 0.66-0.71; Dutton et
al. 1999). Such diversity likely provides the DPS with some capacity
for adapting to long-term environmental changes, such as cyclic or
directional changes in ocean environments due to natural and human
causes (McElhany et al. 2000; NMFS 2017). Nesting habitat is mainly
restricted to mainland beaches along the same coast. The DPS does not
exhibit temporal or seasonal nesting diversity, with most nesting
occurring between October and March. This limits resilience. For
example, short-term spatial and temporal changes in the environment are
likely to affect all nesting females in a particular year. The foraging
strategies are somewhat diverse, with turtles foraging in coastal and
oceanic waters. However, most turtles forage in the East Pacific Ocean,
where they are similarly exposed to the effects of climate change,
ENSO, or the Pacific Decadal Oscillation. Thus, the DPS has limited
resilience.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
The destruction or modification of habitat is a threat at many
nesting beaches used by turtles of the East Pacific DPS. Foraging
habitat has also been characterized as marginal, particularly in the
eastern tropical Pacific Ocean (pelagic environment) due to relatively
low productivity. Coastal habitat, which is normally associated with
high productivity, may have been marginalized due to high levels of
interactions with coastal artisanal fisheries.
Development threatens the DPS by modifying the preferred beach
habitat for nesting. Sustained and substantial development along the
northern and southern ends of the nesting beach at Playa Grande in Las
Baulas National Park, and in adjacent areas, has resulted in the loss
of nesting beach habitat in addition to the removal of much of the
natural beach vegetation. As a result, erosion has increased and led to
other environmental damages to sand that are associated with human
development, including significant changes to elevation, water content,
particle size, pH, salinity, organic content and calcium carbonate
content (Clune and Paladino 2008). Within the past two decades,
beachfront development in the town of Tamarindo (across Tamarindo Bay
from Playa Grande) has resulted in the degradation of nesting beach
habitat, including: Pollution from artificial light, solid and chemical
wastes, beach erosion, unsustainable water consumption, and
deforestation. Hotels in this area have replaced a significant
leatherback nesting area at Playa Tamarindo, which hosted significant
nesting in the 1970s and 1980s (Wallace and Piedra 2012). Playa
Langosta, which is just across from Tamarindo, is inundated with lights
and noise from the town (Wallace and Piedra 2012). Currently,
development has been curtailed due mainly to water issues (i.e.,
drought). Any additional development would damage the current
hydrology. The Leatherback Trust, a local nonprofit working at Las
Baulas National Park, has acquired some properties to prevent
development, but property costs have increased over time. At Las Baulas
National Park, 10 percent of nests were being inundated by tidal flows.
To mitigate this threat, nests at risk of tidal inundation were
relocated to another site on the same beach or into a hatchery.
Hatchling production slightly increased due to the establishment of the
hatchery, where approximately two percent of hatchlings were produced
from 1998 to 2004 (Santidri[aacute]n Tomillo et al. 2007). We conclude
that coastal development in Costa Rica is a threat to this DPS.
In Mexico, the extent of development near nesting beaches is
generally low, given the remoteness of the beaches in Baja California
and on the mainland. Reviewing the location of these nesting beaches,
we found very few roads or development nearby. The main nesting beaches
remain somewhat isolated, with very few roads or development adjacent
to the nesting beaches. Thus, there is limited threat due to artificial
lighting and generally little to no beach driving except perhaps that
associated with monitoring efforts (L. Sarti, CONANP, pers. comm.,
2018). In 2002, the Commission for Natural Protected Areas designated
two of the index beaches (Mexiquillo and Tierra Colorada) as natural
protected areas (turtle sanctuaries), which helped protect nesting
habitat. Subsequently, in 2003, three of the index beaches (Mexiquillo,
Tierra Colorada, and Cahuit[aacute]n) were listed as Ramsar Sites,
which are wetland sites designated to be of international importance
under the Ramsar Convention.
At Veracruz de Acayo beach in Nicaragua, Salazar et al. (2019) note
that while conservation efforts has reduced the threat of poaching, the
establishment of tourism-focused coastal development that do not comply
with the existence of management plans could threaten the nesting
habitat.
While nesting beaches within this DPS are generally remote and/or
protected due to monitoring and existence of national parks and
wildlife refuges, nesting females, hatchlings, and eggs at Las Baulas
National Park (Costa Rica) nesting beaches are exposed to the
modification of nesting habitat, as a result of development. This
threat impacts the DPS by reducing nesting and hatching success, thus
lowering the productivity of the DPS. We conclude that habitat loss and
modification is a threat to the East Pacific DPS.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
The harvest of nesting females and eggs was the primary cause of
the historical decline in abundance of the East Pacific DPS. Since
then, laws have been passed to protect eggs and turtles. However,
poaching still occurs.
In Mexico, Sarti et al. (2007) attributed the decline of nesting
females to the killing of adult females and intensive egg harvest.
Adult females were historically killed at nesting beaches and in open
waters (Sarti et al. 1994; Sarti et al. 1998). Since 1990, the harvest
of turtles and eggs has been prohibited by national legislation.
However, poaching pressure remains high wherever beach patrols do not
occur (Santidri[aacute]n Tomillo et al. 2017). For example, Mexiquillo
produced hatchlings every season in the 1980s. However, even with
efforts to protect the nests in place, 60 to 70 percent of the total
number of clutches were poached. Nichols (2003) notes that leatherback
turtles were once harvested off Baja California, but their meat is now
considered inferior for human consumption. At present, leatherback
turtles are not generally captured for their meat or skin, but the
poaching of nesting females has been known to occur on beaches such as
Piedra de Tlacoyunque, Guerrero (Sarti et al. 2000).
[[Page 48404]]
Although poaching of turtles and eggs has been consistently reduced
over the years, it still occurs at high levels. Effective conservation
and protection depends on human presence at the nesting beaches
(Santidri[aacute]n Tomillo et al. 2017). Without such protection,
poaching is likely to escalate. This may have occurred at one of the
primary nesting beaches (Mexiquillo), where monitoring and conservation
has not taken place in recent years due to safety concerns (L. Sarti,
CONANP, pers. comm., 2018). Since the mid-1990s, Proyecto La[uacute]d
has been relocating clutches (usually within 1-2 hours of deposition)
to protected fenced areas and releasing hatchlings in different areas
of the beach. These efforts are intended to protect the eggs from
poachers/predators and the hatchlings from predators (Sarti et al.
2007).
In Costa Rica, the population decline was predominantly caused by
egg harvest. Ninety percent of eggs were collected on one of the major
nesting beaches, Playa Grande, a decade or more prior to the reduction
of nesting females (Santidri[aacute]n Tomillo et al. 2007). In the
1950s, there were few nesting females at Playa Grande (Wallace and
Piedra 2012). In the late 1960s and early 1970s, the number of nesting
turtles increased to more than 100 nesting females nightly (Wallace and
Piedra 2012). In the early 1970s, newly constructed roads provided
access to people from distant villages and cities, and egg harvest
increased to more than 90 percent by the late 1970s (Wallace and Piedra
2012). Such high levels of egg harvest persisted for nearly two decades
(Wallace and Saba 2009). Despite protection of nesting beaches at Las
Baulas National Park, illegal poaching of eggs still occurs, though
rarely. The black market for eggs remains strong; local bars throughout
Guanacaste and elsewhere continue to offer shots of raw sea turtle egg
yolks accompanying beer or liquor (Wallace and Piedra 2012).
In 1991, the Parque Nacional Marino Las Baulas was created and
subsequently ratified by law in 1995. The Park consists of three
leatherback nesting beaches: Playa Grande, Playa Ventanas, and Playa
Langosta. The establishment of the park ensured increased protection at
all three nesting beaches, greatly reducing egg poaching in the area.
Poaching of eggs was reduced from 90 percent prior to 1990/1991, to 50
percent in 1990/1991, 25 percent in 1991 through 1993, and near 0
percent in 1993/1994 (Santridi[aacute]n Tomillo et al. 2007). To
mitigate poaching, nests are often relocated. However, relocation may
reduce hatching success (reviewed in Hern[aacute]ndez et al. 2007;
Eckert et al. 2012). In Playa Grande, Costa Rica, fewer females were
produced in translocated nests; cooler nests due to a lower number of
metabolizing embryos may have reduced hatchling success (Sieg et al.
2011).
In Nicaragua, prior to protection in the early 2000s, poachers took
nearly 100 percent of the nests at the three nesting beaches. Nesting
beach protection has occurred at Veracruz since 2002, Juan Venado since
2004, and Salamina since 2008. An average of ten community team members
(mostly ex-poachers) monitor beaches seasonally. From 2002 to 2010, up
to 420 nests were recorded and an estimated 94 were protected (Urteaga
et al. 2012). While Veracruz de Acayo and Salamina are protected at 100
percent, Isla Juan Venado is not permanently monitored. Therefore,
poaching is likely to occur. Poaching occurs at high levels at other
beaches, such as Playa El Mogote. During the 2001/2002 nesting season,
23 of 29 nests were poached (79 percent), and the remaining six nests
were protected in a hatchery (Arauz 2002). Due to the high level of
poaching in this area, when possible, researchers from Flora & Fauna
International relocated 98 nests between 2002 and 2004. However, these
nests had a low emergence rate (22 percent; Urteaga and Chac[oacute]n
2008).
Extensive and prolonged effects of comprehensive egg harvest have
depleted the leatherback population in Costa Rica and Mexico, with egg
harvest levels of nearly 90 percent for about two decades (Sarti et al.
2007; Santidri[aacute]n Tomillo et al. 2008; Wallace and Saba 2009).
Currently, nesting females and eggs of the East Pacific DPS are exposed
to poaching. Though efforts have reduced the levels of poaching of both
eggs and nesting turtles, egg poaching remains high and affects a large
proportion of the DPS. Poaching of nesting females reduces both
abundance (through loss of nesting females) and productivity (through
loss of reproductive potential). Such impacts are high because they
directly remove the most productive individuals from DPS, reducing
current and/or future reproductive potential. Egg harvest reduces
productivity only, but over a long period of time, this also reduces
recruitment and thus abundance. Given the high exposure and impacts, we
conclude that overutilization, as a result of poaching, poses a major
threat to the DPS.
Disease or Predation
Little is known about diseases and parasites in leatherback
turtles, although fibropapillomatosis has been described as a major
epizootic disease in hard shelled turtles. A fibropapilloma tumor (in
regression) was found on one nesting female at Mexiquillo, Mexico in
1997 (Huerta et al. 2002). Various bacteria have also been documented
in leatherback eggs. Soslau et al. (2011) sampled eggs laid on a Costa
Rican beach to determine if bacteria were contributing to the low
hatching rate (50 percent). The bacteria identified (i.e., species of
the Bacillus, Pseudomonas, and Aeromonas genera) are known pathogens to
humans and may account for developmental arrest of the turtle embryo
(Soslau et al. 2011).
Numerous predators prey on East Pacific leatherback turtles
throughout their life stages. Eggs and hatchlings are eaten by crabs,
ants, birds, reptiles, mammals, and fish (Eckert et al. 2012). In Costa
Rica, during the 1993/1994 nesting season, several nests were lost to
predation and infestation by maggots (Schwandt et al. 1996). In the
Nicoya Peninsula, on the Pacific coast of Costa Rica, Squires (1999)
documented evidence of potential nest predation by dogs, coyote, and
raccoon. Predation of hatchlings by dogs and raccoons has increased in
Playa Grande due to an increase in development in the area (P.
Santridi[aacute]n Tomillo, The Leatherback Trust, pers. comm., 2019).
For adult turtles, principal predators at sea include killer
whales, crocodiles (Pritchard 1981), and sharks, while nesting females
are taken by crocodiles (Bedding and Lockhart 1989), tigers, and
jaguars (Pritchard 1971). Sarti et al. (1994) observed a lone male
killer whale feeding on a single gravid female near Michoac[aacute]n,
Mexico, apparently consuming only certain parts of the turtle and
discarding others (e.g., female reproductive organs). In summary, eggs,
hatchlings, and some adults are exposed to predation. For this DPS, the
primary impact is to productivity (i.e., reduced egg and hatching
success). Predation on nesting females, while rare, reduces abundance
and productivity. Nest predation is mitigated through screening of
nests, relocation of nests to hatcheries and releasing hatchlings in
safer areas of the beach, and protecting nesting females from large
predators such as dogs and jaguars (Sarti et al. 2007); some of these
efforts are funded through the MTCA. We conclude that predation is a
threat to the East Pacific DPS.
Inadequacy of Existing Regulatory Mechanisms
Several international regulatory mechanisms apply to turtles in
this DPS. The IAC, in particular, prohibits the harvest of turtles and
eggs. CITES
[[Page 48405]]
limits all international trade of the species. There are also
international efforts to reduce fisheries bycatch.
In 2015, at the 7th Conference of the Parties, the IAC resolved to
prioritize conservation actions in their work programs that would help
``reverse the critical situation of the leatherback sea turtle in the
Eastern Pacific.'' Specifically, parties were urged to: (1) Submit
leatherback bycatch information annually to the IAC Secretariat; (2)
improve leatherback turtle fishery monitoring efforts through the use
of on-board observers; (3) report annually on the measures they have
taken to reduce leatherback bycatch in their fisheries; (4) enhance
leatherback nest monitoring and protection to increase hatchling
survival and protect nesting beach habitat; (5) foster safe handling
and release of incidentally bycaught leatherback turtles in fisheries;
and (6) agree to a five-year strategic plan containing key activities
related to the resolution (CIT-COP7-2015-R2). The strategic plan was
patterned after the Regional Action Plan for Reversing the Decline of
the Eastern Pacific Leatherback (https://savepacificleatherbackturtles.org) and included measures to reduce
fisheries bycatch of adult and subadult leatherback turtles, the
identification of high risk areas with fisheries and leatherback
turtles, the identification and protection of important areas for
leatherback turtle survival in different life stages, the elimination
of any consumption and illegal use of leatherback turtles, and nesting
site protection.
As mandated by the 1994 North American Agreement for Environmental
Cooperation, the Commission for Environmental Cooperation (CEC)
encourages Canada, the United States, and Mexico to adopt a continental
approach to the conservation of flora and fauna. In 2003, this mandate
was strengthened as the three North American nations launched the
Strategic Plan for North American Cooperation in the Conservation of
Biodiversity. The North American Conservation Action Plan (NACAP)
initiative began as an effort promoted by the three nations, through
the CEC, to facilitate the conservation of marine and terrestrial
species of common concern. In 2005, the CEC supported the development
of a NACAP for Pacific leatherback turtles by Canada, the United
States, and Mexico. Identified actions in the plan addressed three main
objectives: (1) Protection and management of nesting beaches and
females; (2) reducing mortalities from bycatch throughout the Pacific
Basin; and (3) waste management, control of pollution, and disposal of
debris at sea.
In 2015, the Eastern Pacific Leatherback Network (also known as La
Red de la Tortuga La[uacute]d del Oc[eacute]ano Pacifico (Red
La[uacute]d OPO) (www.savepacificleatherbacks.org)) was formed to
address the critical need for regional coordination of East Pacific
leatherback conservation actions to track conservation priorities and
progress at the population level. This network has brought together
conservationists, researchers, practitioners and government
representatives from 22 institutions across nine East Pacific nations
with varying priorities, capacities and historical experiences in
leatherback research and conservation to contribute to shared
activities, projects, and goals. Through these efforts, Red La[uacute]d
OPO now has mutually-agreed upon mechanisms for sharing information and
data, as well as standardized protocols for nesting beach monitoring
and bycatch assessments/fishing practices.
The Convention for the Protection of Natural Resources and
Environment of the South Pacific, also known as the Noumea Convention,
has been in force since 1990 and includes 26 Parties (as of 2013). The
purpose of the Convention is to protect the marine environment and
coastal zones of the South-East Pacific, and beyond that area, the high
seas up to a distance within which pollution of the high seas may
affect that area.
In 2015, the IATTC passed a resolution that requires large longline
vessels fishing in the eastern tropical Pacific Ocean to carry
observers. Cooperating parties that have documented interactions with
sea turtles in their longline fleet are required to maintain at least
five percent observer coverage and provide an annual report to the
IATTC. Unfortunately, the forms used by observers to report incidents
are not standardized, so in some cases, the reports did not include
species identification, condition of the released turtles, and location
of the interactions, and the five percent minimum coverage is often not
met. Nations without reported bycatch of sea turtles simply provided a
statement to that effect. In the few reports we reviewed, leatherback
turtles comprised some of the bycatch in the eastern tropical Pacific
Ocean, but there were few details on the events (C. Fahy, NMFS, pers.
comm., 2018). In 2007, the IATTC passed a resolution requiring nations
to conduct research on sea turtle bycatch reduction measures in their
longline fleets (e.g., use of circle hooks and fish bait). Despite
results in both the Atlantic and Pacific longline fleets showing that
use of circle hooks/fish bait significantly reduced leatherback bycatch
rates (Swimmer et al. 2017), nations are not required to use this hook/
bait combination. In 2017, at an IATTC sea turtle bycatch reduction
workshop, the United States presented findings on longline bycatch
reduction and proposed a stronger resolution that would require use of
this methodology. However, some nations resisted, and the resolution
did not move forward for consideration at the annual IATTC meeting.
Throughout the world, illegal, unreported, and unregulated (IUU)
fishing leads to underestimates of bycatch. In Mexico, there is a lack
of effective fisheries governance, resulting in highly uncertain
fishery statistics. For example, from 1950 to 2010, total fisheries
catch, including estimated IUU catch and discarded bycatch, was nearly
twice as high as the official statistics (Cisneros-Montemayor et al.
2013). Thus, the bycatch threat of commercial fisheries in Mexico may
be higher than currently estimated.
In addition, several international treaties and/or regulatory
mechanisms protect East Pacific leatherback turtles. While no single
law or treaty can be 100 percent effective at minimizing anthropogenic
impacts to sea turtles in these areas, there are several international
conservation agreements and laws in the region that, when taken
together, provide a framework within which sea turtle conservation
advances can be made (Frazier 2012). In addition to protection provided
by local marine reserves throughout the region, sea turtles may benefit
from the following broader regional effort: (1) The Eastern Tropical
Pacific (ETP) Marine Corridor (CMAR) Initiative supported by the
governments of Costa Rica, Panama, Colombia, and Ecuador, which is a
voluntary agreement to work towards sustainable use and conservation of
marine resources in these nations' waters; (2) the ETP Seascape Program
managed by Conservation International that supports cooperative marine
management in the ETP, including implementation of the CMAR; (3) the
IATTC and its bycatch reduction efforts through resolutions on sea
turtles, observer coverage, etc.; (4) the IAC, which is designed to
lessen impacts on sea turtles from fisheries and other human impacts;
and (5) the Permanent Commission of the South Pacific (Lima
Convention), which has developed an Action Plan for Sea Turtles in the
Southeast Pacific.
[[Page 48406]]
Most nations within the range of the East Pacific DPS have laws
prohibiting the harvest of turtles and eggs. This applies to nesting
turtles and those captured at sea. National laws in Mexico (1990
Presidential Decree), Costa Rica (2002 Presidential Decree N[deg]8325:
The Law of Protection, Conservation, and Recuperation of Marine
Turtles), and Nicaragua (Law No. 651 and Ministrial Resolution No. 043-
2005) protect nesting females and eggs and nesting beaches. However,
poaching remains a major threat. Although laws prohibit the harvest of
turtles in Peru, fishermen consume leatherback turtles bycaught in
small-scale fisheries (Alfaro-Shigueto et al. 2011), indicating
inadequate enforcement of existing laws. In other nations where
leatherback turtles of this DPS are bycaught, the turtles are released
and not retained (e.g., Chile; Donoso and Dutton 2010).
Several protected areas have been established throughout the range
of the DPS. Most of the nesting beaches in Mexico and Costa Rica are
protected from egg and turtle poaching, with effective monitoring to
ensure low levels of poaching. Poaching likely continues at unprotected
and remote beaches, and at those that contain an extensive coastline
that is difficult to monitor and protect. Protected nesting beaches in
Mexico include: Mexiquillo (until 2013); Playa de Tierra Colorada,
Playa Cahuit[aacute]n, Playa San Juan, Bahia de Chacahua, and Playa
Barra de la Cruz. Protected nesting beaches in Costa Rica include: Las
Baulas National Park (Playa Grande, Playa Langosta, and Playa
Ventanas), Naranjo (National Park), Cabuyal (under no official
management category), Nombre De Jes[uacute]s (under no official
management category), Ostional (wildlife refuge), and Caletas (wildlife
refuge). Protected nesting beaches in Nicaragua include: Salamina-Costa
Grande, Veracruz de Acayo (Chacocente Wildlife Refuge).
Marine protected areas also exist. The waters of the Las Baulas
National Park, which represents a hotspot for inter-nesting females and
breeding males, are protected out to 22.2 km as a no-take zone for all
fishing activity. However, satellite telemetry data for nesting females
at these beaches over three seasons revealed that the turtles move well
outside these boundaries during their inter-nesting period, which makes
them vulnerable to fisheries outside the park (Shillinger et al. 2010).
Data from 44 females that were tagged off Las Baulas National Park
revealed a high use habitat within 6 nm from the nesting beaches, but
overall revealed a generally large range, covering over 33,000 km\2\,
from the Nicoya Peninsula, east into the Gulf of Nicoya in Costa Rica,
and north to coastal habitats within 30 kilometers offshore from
southern Nicaragua. The marine areas adjacent to this protected
boundary are not managed under any type of status (Shillinger et al.
2010). Fisheries within Costa Rica and Nicaragua's EEZ include trawl,
gillnet and longline that continue to operate.
In summary, numerous regulatory mechanisms exist to protect
leatherback turtles, eggs, and nesting habitat throughout the range of
this DPS. Although the regulatory mechanisms provide some protection to
the species, many do not adequately reduce the threat that they were
designed to address, generally as a result of limited implementation or
enforcement. As a result, bycatch, incomplete nesting habitat
protection, and poaching remain threats to the DPS. We conclude that
the inadequacy of existing regulatory mechanisms is a threat to the
East Pacific DPS.
Fisheries Bycatch
Bycatch in commercial and recreational fisheries, both on the high
seas and off the coasts, is the primary threat to the East Pacific DPS.
This threat affects the DPS by reducing the abundance of all life
stages of the DPS (with the likely exception of hatchlings).
Integrating catch data from over 40 nations and bycatch data from
13 international observer programs, Lewison et al. (2004) estimated the
numbers of leatherback turtles taken globally by pelagic longliners to
be more than 50,000 leatherback turtles in just one year (2000). With
over half of the total fishing effort (targeting tuna and swordfish)
occurring in the Pacific Ocean, an estimated 20,000 to 40,000
leatherback turtles interacted with longline fishing during the year
studied. Fishing effort was highest in the central South Pacific Ocean
(south of Hawaii), which overlaps with the foraging range of this DPS.
Because observers are in place on only a fraction of longline vessels
in the eastern tropical Pacific Ocean, and a requirement came into
effect only recently through an IATTC resolution, these estimates are
likely a minimum. More recently, Molony (2005) and Beverly and Chapman
(2007) estimated sea turtle longline bycatch to be approximately 20
percent of that estimated by Lewison et al. (2004), or approximately
200 to 640 leatherback turtles annually. Where tuna species are
targeted, bycatch of turtles in the deep-set longline gear often
results in mortality due to drowning. Additional studies indicate the
high impact of industrial longline fleets on leatherback turtles (e.g.,
Spotila et al. 1996, 2000).
In their global study of sea turtle bycatch, where available,
Wallace et al. (2013) found that longline bycatch had a low impact, but
that net bycatch had a high impact on the East Pacific RMU. The impact
of local artisanal fleets (using gillnets and longlines) that fish
closer to shore is less documented.
In Mexico, leatherback turtles wash to shore entangled in longlines
and driftnet, indicating interaction and mortality (Sarti et al. 2007).
Ortiz-Alvarez et al. (2019) conducted a bycatch survey across 48
different ports (933 fishers) in Mexico, Nicaragua and Costa Rica
between October 2016 and July 2017 in an effort to improve the
understanding of leatherback bycatch in artisanal fisheries,
particularly where data are lacking. The surveys represented on average
over 30 percent of the fishing fleet per port for both Nicaragua and
Costa Rica and 6 percent per port for Mexico. In Mexico, where gillnets
were the most frequently reported gear, fishers (n = 709) reported an
estimated bycatch of 300 leatherback turtles in the previous year, with
65 percent in ``good condition;'' 76 percent of fishers released
turtles alive (three percent consumed or sold the turtles). Estimated
average bycatch rates per vessel were 1.0 for Costa Rica and Nicaragua
and 2.3 for Mexico. In Costa Rica, leatherback turtles were primarily
caught in longlines and released alive; 75 percent of the Costa Rican
fishermen reported that bycaught leatherback turtles were in ``good
condition.'' In Nicaragua, where gillnets were the most frequently
reported gear, 18 percent of fishers reported that leatherback turtles
were in ``good condition;'' 76 percent of fishers released turtles
alive (six percent consumed or sold the turtles; Ortiz-Alvarez et al.
(2019).
Recent surveys of 765 Ecuadorian, Peruvian, and Chilean fishermen
(at 43 ports, representing 28 to 63 percent of ports) reported the
following leatherback interaction rates (as a percentage of total
interactions with sea turtles): 2.81 percent of 40,480 interactions
(32.5 percent mortality) in Ecuador, 14.87 of 5,828 interactions (50.8
percent mortality) in Peru, and 27.83 percent of 170 interactions (3.2
percent mortality) in Chile (Alfaro-Shigueto et al. 2018). Mortality
rates reported for all sea turtles were 3.2 percent in Chile, 32.5
percent in Ecuador, and 50.8 percent in Peru (Alfaro-Shigueto et al
2018).
The swordfish gillnet fisheries in Peru and Chile may have
contributed to the decline of the DPS. The decline in the nesting
population at Mexiquillo
[[Page 48407]]
occurred at the same time that effort doubled in the Chilean driftnet
fishery (Eckert 1997). Using data collected from Frazier and Montero
(1990) regarding leatherback takes in a swordfish gillnet fishery from
one port in Chile (San Antonio), and extrapolating to other ports in
Chile and Peru, with an increased level of effort observed through the
mid-1990s, Eckert (2007) estimated that a minimum of 2,000 leatherback
turtles were killed annually by the combined swordfish fishing
operations (only gillnet) off Peru and Chile. After some fleets
switched from large mesh gillnet to longline to target swordfish, this
estimate has declined by at least an order in magnitude. Research
conducted in the Chilean large-mesh gillnet fishery to reduce bycatch
of marine mammals and sea turtles indicates that less than five
leatherback turtles have interacted with the fishery (on observed
vessels) since 2014, and all were released alive (C. Fahy, NMFS, pers.
comm., 2018).
In Peru, the capture of leatherback turtles has been prohibited
since 1976, although retention of bycaught leatherback turtles
continues (FAO 2004). From 1985 to 1999, based on field books, diaries,
specimen data sheets, fishery statistics files and unpublished reports,
30 leatherback turtles were captured in fisheries (in Alfaro-Shigueto
et al. 2007). From July 2000 to November 2003, observers at 8 ports,
from Mancora in northern Peru to Morro Sama in the south, reported 133
leatherback turtles caught by artisanal fishing gear, with 76 percent
caught in gillnets and 24 percent caught in longlines targeting fish,
sharks, and rays (Alfaro-Shigueto et al. 2007). Of the total caught,
41.4 percent (n = 55) were released alive and 58.6 percent (n = 78)
were retained for human consumption. Of the leatherback turtles
retained and measured (n = 6), the size ranged from 98 to 123 cm curved
carapace length (CCL), indicating that both subadults and adults are
encountered by artisanal fisheries off Peru. Researchers recently
assessed and quantified sea turtle mortality levels in one fishing
village in central-southern Peru (San Andr[eacute]s) through sampling
dump sites (97.3 percent) and strandings (2.7 percent) over a 5-year
period (2009 to 2014). Of 953 carapaces recorded, leatherbacks
comprised only 1.4 percent of sea turtles (n = 13). However, this study
still confirmed that they were consumed or sold for human consumption.
With a mean CCL of 113.0 cm (range: 80 to 135, n = 10), 70 percent of
the leatherbacks were juveniles and 30 percent were sub-adults. There
were no adults. Researchers noted that the meat was used to support
separate demands: Fishermen families' consumption, local trade, and
``special'' orders from Lima (Quispe et al. 2019). Using data from
shore-based and on-board observers, Alfaro-Shigueto et al. (2011)
estimated the mean annual leatherback bycatch as follows: 40 turtles
(with a range of 37 to 44) in the driftnet fishery, with 80 percent
released alive; six turtles (with a range of 3 to 9) in the dolphinfish
longline fishery, all released alive; and 26 turtles (with a range of
24 to 27) in the shark longline fishery, all released alive. Alfaro-
Shigueto et al. (2015) assessed the bycatch of leatherback turtles in
driftnet vessels in northern Peru (through at-sea monitoring) and
central Peru (shore-based monitoring). From December 2013 to November
2014, 31 leatherback turtles were captured, of which 13 died.
Interactions occurred primarily with juveniles and subadults (mean CCL
was 125.1 14.8). Nearshore driftnets from San Jose
(northern Peru) captured 20 leatherback turtles (five dead). At least
one animal was butchered, indicating that even animals caught alive may
be killed, despite Peruvian laws restricting such practices.
Approximately 3,000 net vessels fish along the coast of Peru, but only
a fraction were included in this study (Alfaro-Shigueto et al. 2015).
Efforts are being made to patrol nets to reduce bycatch, conduct
extensive education and outreach, and increase regulation and
enforcement (Alfaro-Shigueto et al. 2015). A review of information
collected from official statistics, literature, and surveys of beaches
and dumpsites revealed that the size of captured leatherback turtles
declined over the years. In 1987, the mean CCL of captured leatherback
turtles was 117 10.65 cm, while in 2005, the mean CCL was
109.27 14.4, possibly indicating overexploitation due to
systematic and sustained harvests, particularly during El Ni[ntilde]o
years (Campos et al. 2009). Greater captures of all sea turtles,
including leatherback turtles, occurred during periods of El
Ni[ntilde]o, when turtles are more likely to be found in more coastal
waters (where there is increased artisanal fishery activity) due to
environmental variability and availability of jellyfish in those areas
(Campos et al. 2009).
In Chile, a commercial fishery was established in 2001 that
permitted longlining for swordfish (shallow-set) with the condition
that all vessels were required to take an observer on board to collect
information on bycatch. Between 2001 and 2005, over 10 million hooks
were observed, and leatherback turtles were the most common species
caught (n = 284), with the majority (n = 282) released alive.
Leatherback turtles were caught primarily between 24[deg] S and 38[deg]
S (furthest south was 38[deg]39' S and 84[deg]15' W) in less than 4
percent of the sets with an overall mean of 0.0268 turtles per one
thousand hooks. Size estimates revealed both juveniles and adults.
Fishermen were trained to use the best practices for de-hooking,
disentangling, and releasing sea turtles, which likely increased the
survival rate of leatherback turtles (Donoso and Dutton 2010).
Researchers recently presented information on the incidental capture of
sea turtles in industrial and artisanal longlines, gillnets and
artisanal espinel (i.e., small-scale handline or longline) fisheries
all targeting swordfish off Chile (Z[aacute]rate et al. 2019). Over an
8-year period (2006-2014), 182 leatherbacks were documented as bycatch
(mortality of bycaught turtles was not reported). Over this study
period, 44 percent of turtles were caught in industrial longline, 28
percent in artisanal espinel, 17 percent in gillnets and 11 percent in
artisanal longline (with sea turtle species undefined). Researchers
noted that while observer coverage in the industrial longline fleet has
been generally high (>70 percent of total fishing trips), the
monitoring coverage of artisanal espinel and gillnets is very low (<3
percent). Thus, these estimates of bycatch can be considered minimal.
While the number of industrial and artisanal vessels has declined (from
12 vessels in 2001 to 3 vessels in 2014, the number of artisanal
espinel and gillnet vessels has not declined, remaining around 90
vessels (Z[aacute]rate et al. 2019).
We conclude that juvenile and adult life stages of the East Pacific
DPS are exposed to high fishing effort throughout their foraging range
and in coastal waters near nesting beaches. Mortality is also high in
some fisheries, with reported mortality rates of up to 58 percent due
in part to the use of gillnets and as well as consumption of bycaught
turtles in Peru. As noted above, there have been efforts by individual
nations and regional fishery management organizations to mitigate and
reduce the threat of bycatch, but those efforts have not been
successful at ameliorating the risks. We conclude that fisheries
bycatch remains a major threat to the East Pacific DPS.
Pollution
Pollution is a threat to the East Pacific DPS. Pollution includes
contaminants, marine debris, and ghost fishing gear. The South Pacific
Garbage Patch, discovered in 2011 and confirmed in
[[Page 48408]]
2017, contains an area of elevated levels of marine debris and plastic
particle pollution, most of which is concentrated within the ocean's
pelagic zone and in area where leatherback turtles forage for many
years of their life. The area is located within the South Pacific Gyre,
which spans from waters east of Australia to the South American
continent and as far north as the Equator.
Given the amount of floating debris in the Pacific Ocean (Lebreton
et al. 2018), marine debris has the potential to be a significant
threat to the East Pacific leatherback population. The precise impact
cannot be quantified using the best available data. Leatherback turtles
subsist primarily on jellyfish and other gelatinous zooplankton and may
be prone to ingesting plastics resembling their food source (Mrosovsky
1981; Schuler et al. 2013, 2015). Dead leatherback turtles have been
found choked on plastic bags, and phthalates derived from plastics have
been found in leatherback egg yolk (Lebreton et al. 2018).
Prior to the early 1990s, high seas driftnet fisheries freely
operated in the Pacific Ocean and interacted with thousands of sea
turtles. Researchers estimated that over 1,000 leatherback turtles were
taken by the combined fleets of Japan, Korea, and Taiwan during a one-
year period (Wetherall 1997). However, because genetic analyses of
Pacific leatherback turtles were relatively new at that time, the data
does not indicate the nesting beach origin of those bycaught
leatherback turtles. In 1992, a UN moratorium banned high seas driftnet
fisheries, so that active large scale driftnets no longer pose a threat
to leatherback turtles. However, numerous discarded driftnets continue
to entangle and drown leatherback turtles in a phenomenon known as
``ghost fishing'' (Gilman et al. 2016),
In 2007, the IATTC passed a resolution pertaining to sea turtle
bycatch in purse seine and longline fisheries which primarily target
tuna. In order to address the marine debris and potential interactions
with sea turtles in the eastern tropical Pacific Ocean, fishermen are
required to disentangle sea turtles entangled in fish aggregating
devices, even if the device does not belong to the vessel.
Only a few studies of levels or effects of toxins on leatherback
turtles have examined effects to their health and fitness, as well as
any effects to eggs and hatchlings. Sill et al. (2008) sampled non-
viable leatherback eggs and hatchlings that died in the egg chamber at
Las Baulas National Park. Researchers analyzed the samples for metals
and other toxicants to explore the relationship between pollution and
hatching success for 30 females. Metal levels were highly variable, but
there were no significant differences within and between groups of
females, and none of the pesticides tested were present in the samples
(Sill et al. 2008). Overall, the study found no relationship between
metal concentrations and hatching success. The researchers postulated
that eggs may take up some metals from the nest environment and deposit
other metals in the egg shell, as unhatched eggs contained more nickel,
copper, and cadmium and contained significantly less iron, manganese
and zinc than dead hatchlings (Sill and Paladino 2008).
As with all leatherback turtles, entanglement in and ingestion of
marine debris and plastics is a threat that likely kills several
individuals a year. However, data are not available because most
affected turtles are not observed. Given the amount of pollution
turtles are exposed to throughout their lifetime, this has the
potential to be a significant threat to the East Pacific leatherback
population, although the impact cannot be quantified using the best
available data. We conclude that pollution is a threat to this DPS.
Oceanographic Regime Shifts
The East Pacific DPS is affected by oceanographic regime shifts. In
the eastern equatorial Pacific Ocean, reductions in productivity
parameters are primarily associated with ENSO, during which sex ratios
become biased up to 100 percent female (Santidri[aacute]n Tomillo et
al. 2014). There is also an effect on hatching and emergence success in
North Pacific Costa Rica (Santidri[aacute]n Tomillo et al. 2012):
During El Ni[ntilde]o years, hatching success is very low due to dry
and hot conditions on the nesting beaches and is high during La
Ni[ntilde]a events due to increased precipitation in this area. La
Ni[ntilde]a events are characterized by high phytoplankton
productivity, cooler sea surface temperatures, enhanced precipitation
in northwestern Costa Rica, and cooler air temperatures. These factors
lead to increases in the biomass and distribution of gelatinous
zooplankton, the primary food of leatherback turtles. Foraging success
and the frequency of reproduction are enhanced following such periods
of high primary productivity (Saba et al. 2007). Nesting seasons that
follow the La Ni[ntilde]a events, result in peaks in the number of
nesting females, higher than average hatching success and emergence
rates, and a larger proportion of male hatchlings (Saba et al. 2012).
Saba et al. (2008) found that a shift from 1 [deg]C to -1 [deg]C in the
El Ni[ntilde]o sea surface temperature anomaly resulted in a five-fold
increase in leatherback remigration probabilities at Playa Grande. Such
large-scale regime shifts are likely to affect the entire DPS.
Productivity is positively (La Ni[ntilde]a) or negatively (El
Ni[ntilde]o) impacted. Wallace et al (2006) hypothesize that prey
availability related to ENSO exacerbates the effects of fisheries
bycatch mortality, resulting in declining trends. Because of the small
abundance of the DPS, extended El Ni[ntilde]o events are likely to pose
a threat to the East Pacific DPS.
Climate Change
Climate change is a threat to the East Pacific DPS. The impacts of
climate change include: Increases in temperatures (air, sand, and sea
surface); sea level rise; increased coastal erosion; more frequent and
intense storm events; and changes in oceanographic regimes and
currents.
Climate projections assessed by the IPCC indicate that Central
America is very likely (defined as 90 to 99 percent probability; IPCC
2007) to become warmer and likely (defined as 66 to 90 percent
probability; IPCC 2007) to become drier by 2100 (Saba et al. 2012). In
addition, climate variability is likely to change the strength and
frequency of El Ni[ntilde]o events, although there is less scientific
consensus on the frequency and magnitude of changes to these events. A
climate-forced population dynamics model developed by Saba et al.
(2012) showed sea surface temperatures to be highly correlated with
large phytoplankton productivity throughout a 100-year projection to
the year 2100. Relative to a stable nesting population given mean
surface air temperatures and precipitation from 1975 to 1999, Saba et
al. (2012) estimated that the nesting population at Playa Grande would
decline at a rate of 7 (1) percent per decade over the next
century of climate change under a scenario which considered increasing
emissions from 2000 to 2100 (A2 scenario). Similar declines occurred
for other scenarios (Special Report on Emissions Scenarios 2007). The
nesting population was projected to remain stable up until around 2030
but reduced 75 percent by the year 2100. Hatching success and emergence
rates, which would decrease associated with 2.5 [deg]C warming of the
nesting beaches, served as a primary driver of the decline.
Santidri[aacute]n Tomillo et al. (2012) developed a similar climate
forcing model, which considered projected changes associated with El
Ni[ntilde]o events
[[Page 48409]]
and demonstrated that hatching success would decline from approximately
42 to 18 percent by 2100, while emergence rates would decline between
approximately 76 to 29 percent. The authors concluded that even with
protection at the primary nesting beaches in Costa Rica, with the
general warming of Central America in the near future, the chances of a
new nesting area emerging with more ideal conditions (i.e., cooler and
wetter) is unlikely (Santidri[aacute]n Tomillo et al. 2012).
Increasing sand temperature is an existing threat to the DPS. The
long-term data set on leatherback turtles nesting at Playa Grande,
Costa Rica indicates reduced emergence success, skewed sex ratios, and
increased hatchling mortality as a result of increased sand temperature
(Santidri[aacute]n Tomillo et al. 2015). From 2004 to 2013, primary sex
ratios fluctuated between a minimum sex ratio of 41 percent females
(and the only year with a male-biased hatchling production) to 100
percent females produced during two seasons (Santidri[aacute]n Tomillo
et al. 2014). Low emergence success and low hatchling output (i.e.,
higher mortality as a result of high sand temperatures) were associated
with a strongly biased female ratio, because these resulted from
female-producing high temperatures. Variability in these results occur
during and between nesting seasons, largely due to highly variable
climatic conditions in northwestern Costa Rica, resulting in ``boom-
bust'' cycles in leatherback hatchling production and primary sex
ratios (in Santidri[aacute]n Tomillo et al. 2014). Sand temperatures
are projected to continue to increase, which will likely result in a
further decline in the number of hatchlings produced (Santidri[aacute]n
Tomillo et al. 2014). An increase in the percentage of females could
potentially benefit the productivity of the DPS in the short-term.
However, any such benefits would be tempered by the associated lower
emergence and hatchling success rates. Relocation of sea turtle
clutches that may be ``doomed'' due to high sand temperatures and
inundation is a common conservation practice, particularly at areas
with warming beaches. However, relocation is not always possible and is
also associated with lower emergence and hatchling success rates.
In addition to climate change influencing the nesting beach habitat
of eastern Pacific leatherback turtles, the impacts of a warming ocean
may also affect the environmental variables of their pelagic migratory
and foraging habitat, which may further increase population declines.
As mentioned previously, the preferred foraging habitat of eastern
Pacific is characterized by relatively low sea surface temperatures and
low levels of chlorophyll-a. Using information derived from satellite
tracked leatherback turtles, which established migratory pathways and
core foraging habitat (as summarized in Shillinger et al. 2008), in
combination with generalized additive mixed models, researchers were
able to project that between 2001 and 2100, there would be a net loss
of the core foraging habitat of the DPS. The loss was predicted to be a
15 percent decline over the next century (Willis-Norton et al. 2014).
Depending on whether this population is able to shift their preferred
migratory routes and foraging habitat over time (which is unclear),
remigration intervals may shorten or lengthen, which could influence
reproductive productivity.
Climate change is a threat to the East Pacific DPS that affects
nesting females (e.g., remigration interval and fitness), their progeny
(e.g., hatching success, embryonic development, and feminization of
hatchlings), and foraging subadult and adult leatherback turtles.
Detrimental impacts of increased sand temperatures have already
occurred and are likely to continue or worsen. Foraging areas will also
be impacted via changes in ocean productivity, sea surface
temperatures, and availability of prey.
Conservation Efforts
There are numerous efforts to conserve the leatherback turtle. The
following conservation efforts apply to turtles of the East Pacific DPS
(for a description of each effort, please see the section on
conservation efforts for the overall species): Convention on the
Conservation of Migratory Species of Wild Animals, Convention on
Biological Diversity, Convention on International Trade in Endangered
Species of Wild Fauna and Flora, Convention for the Protection of the
Marine Environment and Coastal Area of the South-East Pacific (Lima
Convention), Convention for the Conservation and Management of Highly
Migratory Fish Stocks in the Western and Central Pacific Ocean (WCPF
Convention), Convention Concerning the Protection of the World Cultural
and Natural Heritage (World Heritage Convention), Eastern Pacific
Leatherback Network, Eastern Tropical Pacific Marine Corridor
Initiative, FAO Technical Consultation on Sea Turtle-Fishery
Interactions, IAC, MARPOL, IUCN, Ramsar Convention on Wetlands, RFMOs,
Secretariat of the Pacific Regional Environment Programme, UNCLOS, and
UN Resolution 44/225 on Large-Scale Pelagic Driftnet Fishing. Although
numerous conservation efforts apply to the turtles of this DPS, they do
not adequately reduce its risk of extinction.
Extinction Risk Analysis
After reviewing the best available information, the Team concluded
that the East Pacific DPS is at high risk of extinction. The DPS
exhibits a total index of nesting female abundance of 755 females at
monitored beaches. Such a limited nesting population size makes this
DPS vulnerable to stochastic or catastrophic events that increase its
extinction risk. This DPS exhibits a decreasing nest trend, which along
with lower than-average productivity metrics, has the potential to
further reduce abundance and increase the risk of extinction. The
nesting range is somewhat limited to the Pacific Central American
coast, with little diversity among sites. Thus, stochastic events could
have catastrophic effects on nesting for the entire DPS, with no
distant subpopulations to buffer losses or provide additional
diversity. Most foraging occurs in the eastern Pacific Ocean, which is
subject to oceanographic regimes shifts that expose the DPS to low-
productivity events. Based on these demographic factors, we find the
DPS to be at risk of extinction as a result of past threats.
Current threats also contribute to the risk of extinction of this
DPS. Fisheries bycatch is the major threat, capturing, and often
killing, turtles throughout their foraging areas, thus reducing
abundance. There are few mechanisms in place, including internationally
through the IATTC or other bilateral or international instruments and
through monitoring and enforcement of coastal fisheries laws, to
mitigate or reduce bycatch. Overutilization is also a major threat.
Historically, harvest of turtles and eggs reduced the once high
abundance of turtles to current low levels. The poaching of eggs
continues, reducing productivity, especially at unprotected beaches,
where egg collection may reach 100 percent and nesting females may also
be at risk of poaching. The effects of climate change, including the
observed and predicted increase in frequency and strength of ENSO
events (i.e., oceanographic regime shifts), are threats to this DPS,
given its restricted foraging range and the vulnerability of nesting
beaches to high sand temperatures and low levels of rainfall, which
affect sex ratios and emergence and hatching success (i.e.,
productivity). Additional threats include: Habitat loss and
modification;
[[Page 48410]]
predation; and pollution. Development modifies nesting habitat.
However, most beaches are protected throughout the nesting range.
Though many regulatory mechanisms are in place, they do not adequately
reduce the impact of these threats. Further, it is important to note
that efforts (e.g., relocation) to protect and mitigate threats from
the harvest of turtles and eggs, predation, and environmental impacts
related to erosion and lethal temperatures are dependent upon the
presence of monitoring or management programs. Some of these are
dependent on funding from the MTCA. Even when undertaken, these efforts
may not be successful.
We determine, consistent with the Team's findings, that the East
Pacific DPS is currently in danger of extinction. Its nesting female
abundance and declining trend make the DPS highly vulnerable to
threats. Though numerous conservation efforts apply to this DPS, they
do not adequately reduce the risk of extinction. We conclude that the
East Pacific DPS is currently in danger of extinction throughout its
range and therefore meets the definition of an endangered species. The
threatened species definition does not apply because the DPS is
currently at risk of extinction (i.e., at present), rather than on a
trajectory to become so within the foreseeable future.
Leatherback Turtle, Overall Species
The petition under review sought specifically to identify the NW
Atlantic population of leatherback sea turtles as a separate DPS and
assign it a different status from the global listing. As explained
throughout this finding, we have determined that seven leatherback
populations would satisfy the tests for recognition under our DPS
Policy (i.e., that they are discrete from one another and significant
to the overall species), and we have referred to these hypothetically,
for purposes of our analysis only, as DPSs. This includes the NW
Atlantic DPS. However, we have also determined that, even if these
populations were formally recognized as DPSs through a listing process
under the Act, each of the DPSs would have the same status as the
overall species, which is currently listed throughout its range
(globally) as endangered. Nothing in the petition or in the best
available information we have reviewed has led us to conclude that
there is any basis to disturb the long-standing global listing, which
remains in effect and is unaffected by this finding. For completeness,
here we present an overview of current information pertaining to the
status of the overall species, including a summary of some of the key
information from the DPS-specific sections as well as an evaluation of
the demographic factors affecting the overall species.
As explained in the Background section, the leatherback turtle was
originally listed as endangered in 1970 under the precursor to the ESA
and was carried forward as an ``endangered species'' when the ESA
became effective. The Services designated the nesting beaches at Sandy
Point, St. Croix (43 FR 43688; September 26, 1978) and surrounding
marine waters (44 FR 17710; March 23, 1979) as critical habitat. NMFS
designated additional marine habitat along 41,914 square miles (108,558
square km) of the U.S. West Coast as critical habitat (77 FR 4170;
January 26, 2012). The Services issued the recovery plans for
leatherback turtles in the U.S. Caribbean, Atlantic, and Gulf of Mexico
(1991) and U.S. Pacific (1998; https://www.fisheries.noaa.gov/action/recovery-plans-leatherback-sea-turtle).
The species has the widest distribution of any reptile, with a
global range extending from 71[deg] N, based on an at-sea capture off
Norway (Carriol and Vader 2002) to 47[deg] S, based on an at-sea
sighting off New Zealand (Eggleston 1971; Eckert et al. 2012). The
species has several thermoregulatory adaptations to allow such a large
latitudinal range, maintain its core temperature while foraging, and
avoid overheating during nesting. These include its large size, low
metabolic rates, countercurrent heat exchange at the base of its limbs,
and peripheral insulation (Frair et al. 1972; Greer et al. 1973;
Paladino et al. 1990; Fossette et al. 2009; Bostrom et al. 2010; Eckert
et al. 2012; Casey et al. 2014; reviewed in Wallace and Jones 2015).
Nesting is restricted to mainly tropical or subtropical beaches.
However, nesting also occurs on temperate beaches of the SW Indian
Ocean (Pritchard and Mortimer 1999). Nesting usually occurs on high-
energy beaches (Pritchard 1976), resulting in high rates of natural
erosion. The primary factors influencing shoreline suitability for
nesting appear to be a lack of abrasive substrate material, a deep-
water approach to minimize energy expenditure needed to reach nesting
sites, and proximity to oceanic currents that can facilitate hatchling
dispersal (Eckert et al. 2012). Leatherback turtles appear to prefer
wide, long beaches with a steep slope, deep rock-free sand, and an
unobstructed deep water or soft-bottom approach (Pritchard and Mortimer
1999; Eckert et al. 2015). As a result, it has been proposed that the
choice of nesting location is based on site characteristics within a
geographic location (MacKay et al. 2014).
Foraging areas are generally characterized by zones of upwelling,
including off the edges of continents, where major currents converge,
and in deep-water eddies (Saba 2013). Important foraging areas include
but are not limited to: upwelling off the west coasts of North and
South America (Benson et al. 2011; Roe et al. 2014); Benguela Current
Marine Ecosystem (Honig et al. 2007); and Canadian waters on the
Scotian Shelf (James et al. 2005a, 2006b, 2007b).
Abundance
Adding together the total indices of nesting female abundance for
all DPSs, the total index of nesting female abundance for the species
is 32,174 females. This number, however, should be considered as a
compilation of seven populations ranging in size from 27 to 20,659
nesting females because nesting female exchange does not occur between
DPSs.
Comparisons with historical accounts of nesting female abundance
are complicated by the discovery of new nesting beaches over time,
changes in remigration intervals and/or clutch frequency, and modified
observational effort. Abundance estimates for even large nesting
beaches were not available prior to 1950 (Rivalan et al. 2006), several
large nesting beaches were not discovered until the 1960s or later
(NMFS and USFWS 2013), and monitoring efforts were variable over time.
Pritchard's 1971 global estimate of 29,000 to 40,000 nesting females
included a maximum estimate (i.e., 40,000 nesting females) based on the
assumption that large nesting aggregations had yet to be discovered
(Pritchard 1971); this estimate did not include large nesting female
abundances from the East Pacific and SE Atlantic Oceans. At that time,
the nesting aggregation at Terengganu, Malaysia nesting population was
thought to be one of the largest; however it has since been extirpated
(Chan and Liew 1996). In 1982, Pritchard revised his initial global
estimate to 115,000 nesting females, based largely on the nesting
beaches in Pacific Mexico (n = 75,000; Pritchard 1982). However, the
1982 estimate was extrapolated from a brief aerial survey and may have
been an overestimate (Pritchard 1996). When the Mexico nesting
population collapsed, Spotila (1996) estimated the total global
estimate to be 34,500 nesting females, with a range of 26,200 to 42,900
nesting females. However, this estimate did not include the nesting
aggregation in Gabon, which in 2002 was identified as
[[Page 48411]]
the largest in the world at that time, with tens of thousands of
nesting females (Witt et al. 2009). Recent data indicate less than
9,000 nesting females in Gabon (Formia in progress). Thus, we find that
leatherback nesting female abundance has declined rapidly in several
populations. Our total index of nesting female abundance for the
species, which does include the largest nesting aggregations from all
DPSs, is lower than previous estimates by at least 10,000 females.
Species go extinct through the loss of populations. Therefore, the
loss of any of these populations (which we refer to in this finding
hypothetically as DPSs) would increase the extinction risk of the
species. Most of the DPSs exhibit total indices of nesting female
abundances that place them at risk for environmental variation, genetic
complications, demographic stochasticity, negative ecological feedback,
and catastrophes (McElhany et al. 2000; NMFS 2017). The current total
index of nesting female abundance for the species reflects the impact
of threats that have affected the species to this point. This reduced
abundance renders it particularly vulnerable to threats and contributes
to its extinction risk.
Productivity
Nest trends are decreasing across the species, except at the least
abundant nesting aggregation in Brazil (i.e., the SE Atlantic DPS),
with a total index of 27 nesting females, which is increasing by 4.8
percent annually. Current nest trends are declining at rates ranging
from -0.3 percent (within the SW Indian DPS) to -9.3 percent (the
overall decline for the NW Atlantic DPS). Historical declines are even
larger. Aerial surveys of nesting beaches in Mexico detected declines
from over 70,000 nesting females in 1982 to fewer than 250 in 1998,
with an annual mortality rate of 22.7 percent (Spotila 2000) and an
overall decline of 97.4 percent in three generations (Wallace et al.
2013). The Terengganu, Malaysia nesting aggregation has declined by
17.9 percent annually from 1967 to 2010. It was been reduced to less
than one percent of its original size between the 1950s and 1995 (Chan
and Liew 1996) and is now considered functionally extirpated.
Significant declines in nesting have been documented for other
populations (Benson et al. 2015). Declining nesting trends reflect the
impact of threats that have been operating on the species, and these
trends increase the extinction risk of the species.
Spatial Distribution
The species occurs over a broad spatial range, in tropical and
temperate waters worldwide, from 71[deg] N to 47[deg] S (Goff and Lien
1988; Carriol and Vader 2002; McMahon and Hayes 2006; Shillinger et al.
2008; Wallace et al. 2010; Benson et al. 2011; Eckert et al. 2012). It
nests and forages across a wide spatial range, which provides some
degree of resilience against local impacts to nesting and foraging
areas. The DPSs are reproductively isolated with little to no gene flow
connecting them. However, within some DPSs there is fine-scale
population structure (Dutton et al. 1999; Dutton et al. 2003; Dutton et
al. 2013; Molfetti et al. 2013). These subpopulations exhibit
metapopulation dynamics, which make a DPS more resilient to stochastic
and environmental changes. It is likely that all DPSs once exhibited
such dynamics, given the ephemeral, high-energy beaches where they nest
and their regional, but not necessarily beach-specific, philopatry
(Dutton et al. 1999; Dutton et al. 2013). However, the reduction of
nesting aggregations within a DPS has likely reduced or removed this
structure, and the associated resilience, in some DPSs and in the
overall species.
Diversity
Relative to other sea turtle species, the leatherback turtle has
low genetic diversity and shallow mtDNA coalescence (Dutton et al.
1999), reflecting its recent global radiation, i.e., Post-Pleistocene
expansion from a refugium in the Indian Ocean (Dutton et al. 1999). As
a species, it uses diverse and widely distributed nesting and forage
areas. Differences in size at maturity, remigration rate, clutch
frequency, and clutch size likely reflect environmental variability
among DPSs (Saba et al. 2008; Saba et al. 2015). The age of the species
and its flexible use of multiple foraging and nesting areas indicate
that the species has some resilience to stochastic and environmental
changes.
Present or Threatened Destruction, Modification, or Curtailment of
Habitat or Range
The destruction or modification of nesting habitat is a threat to
most leatherback turtles, and in some areas, this threat is major, as a
result of development, erosion, or obstruction from logs. By the year
2025, the UN Educational, Scientific and Cultural Organization (2001)
forecasts that human population growth and migration will result in 75
percent of people living within 60 km of the sea. This will place
significant additional pressure on coastal habitats.
Coastal development and associated activities cause accelerated
erosion rates and interruption of natural shoreline migration (National
Research Council 1990). Numerous beaches are eroding due to both
natural (e.g., storms, sea level changes, waves, shoreline geology) and
anthropogenic (e.g., development and expansion, construction of
armoring structures, groins, jetties, marinas, coastal development,
inlet dredging) factors. Such shoreline erosion has led and will
continue to lead to a loss of nesting habitat for leatherback turtles
and potential loss of nests from inundation. Erosion or inundation and
accretion of sand above incubating nests appear to be the principal
abiotic factors that negatively affect incubating egg clutches in some
areas (Dow et al. 2007; USFWS 1999; NMFS and USFWS 2013). Shoreline
structuring can also physically prevent females from reaching suitable
nesting habitat or prevent them from returning to sea (Witherington et
al. 2011).
Low hatching success, relative to other sea turtle species, is
characteristic of many leatherback populations despite high fertility
rates (reviewed by Bell et al. 2003; Eckert et al. 2012). Nest
relocation is undertaken as a conservation measure in some locations
when erosion (or poaching and predation) threaten the viability of a
nest. However, studies have found that hatching success of nests in
hatcheries or nests relocated to another area of a beach is lower than
in situ nests (reviewed in Hern[aacute]ndez et al. 2007; Eckert et al.
2012). In addition, nest relocation results in altered sand
temperatures, which influences the sex ratio of hatchlings produced
(Sieg et al. 2011).
Coastal development and expansion also contributes to habitat
degradation via artificial lighting (i.e., light pollution). The
presence of artificial lighting on or adjacent to nesting beaches
alters the behavior of nesting females (often deterring nesting) and is
often fatal to post-nesting females and emerging hatchlings, when they
are attracted to terrestrial light sources and drawn away from the
water (Witherington 1992; Sella et al. 2006; Witherington et al. 2014).
As hatchlings head toward lights or meander along the beach, their
exposure to predators and likelihood of desiccation are greatly
increased. Artificial lighting may also affect hatchlings that
successfully find the water, causing them to be misoriented after
entering the surf zone or while in nearshore waters.
[[Page 48412]]
The modification of nesting habitat generally results in loss of
productivity for the species, as a result of reductions in nest and
hatching success. In addition, several DPSs experience nesting beach
habitat modifications (e.g., artificial lighting, logs, and other
obstructions) that result in the death of nesting females and
hatchlings. Therefore, abundance is also reduced, posing an even
greater threat to the continued existence of the turtles of the DPS.
The loss and modification of nesting habitat poses a major threat to
the species.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
Historically, the harvest of turtles and eggs was the primary
threat to the species, leading to the loss of severe depletion of many
nesting aggregations worldwide (Spotila et al. 1996). At one point in
time, egg harvest was ubiquitous with all nests taken at many beaches
(Chan and Liew 1996; Sarti et al. 2007; reviewed by Eckert et al.
2012). For the NW Atlantic, NE Indian, and West Pacific DPSs, legal
harvest of turtle and/or eggs continues. Despite laws in many
countries, the poaching of eggs continues at most nesting beaches,
ranging in severity from minor at monitored or protected beaches to
near 100 percent harvest at unmonitored beaches. Nesting females, and
turtles caught at sea, continue to be poached for their meat, eggs, and
fat in many locations (Eckert et al. 2012). As described in detail in
the prior sections evaluating the status of each individual DPS, the
harvest of eggs and turtles is a threat to each and to the species
overall, and for the NE Indian and West Pacific DPSs, it is a primary
threat. The legal and illegal harvest of turtles and eggs poses a
threat to the species.
Disease or Predation
We do not have adequate information on disease to assess its impact
on the species. However, we have enough information to conclude that
predation is clearly a threat. Numerous species prey on leatherback
eggs and hatchlings. Eckert et al. (2012) provide an exhaustive list of
the documented predators for each life stage and area. For eggs, common
predators include ants, ghost crabs, monitor lizards, crows, mongoose,
domestic and feral dogs, and feral pigs (Eckert et al. 2012). For
hatchlings, common predators include the terrestrial predators listed
above as well as numerous species of carnivorous fish, including
sharks. Sharks and killer whales, and in some areas jaguars and
crocodiles, prey on subadult and adult turtles. Predation on eggs and
hatchlings is common and reduces productivity of the species; predation
on subadults and adults is less prevalent but reduces abundance when it
occurs. Predation is a threat to the species, and for some DPSs, it is
a major threat.
Inadequacy of Existing Regulatory Mechanisms
Numerous regulatory mechanisms provide certain protections to sea
turtles at the international, regional, national, and local levels. For
example, the harvest of sea turtles and their eggs is prohibited by
regional conventions and national laws. Fisheries bycatch is also
addressed, although not comprehensively, by several international and
national instruments and/or governing bodies. However, as we detail
below and has been discussed in prior sections reviewing each
individual DPS, these measures are often poorly implemented or
enforced, resulting in inadequate protections against the threats they
are designed to ameliorate.
In some nations (e.g., South Africa) sea turtles were among the
first species to receive legal protections and have been the focus of
concentrated conservation efforts. However, current regulatory
mechanisms often fall short of preventing further population declines
and ensuring persistence (Eckert et al. 2012). For many nations the
regulations in place are inadequate (usually due to lack of enforcement
and implementation) to address the impacts of a wide range of
anthropogenic activities that directly injure and kill turtles, disturb
eggs, disrupt necessary behaviors, and alter terrestrial and marine
habitats used by the species. In many areas, regulations for the
harvest of turtles and eggs are inadequate due to a lack of
enforcement. In some areas, the regulation of fisheries bycatch do not
adequately reduce associated mortality. Fishery observer coverage is
often inadequate to accurately estimate leatherback bycatch.
Due in part to their worldwide distribution and highly migratory
nature, combined with nesting site fidelity, leatherback turtles
require international, national, regional, and local protection. Hykle
(2002) and Tiwari (2002) reviewed the value of some international
instruments and concluded that they vary in their effectiveness. Often,
international treaties do not realize their full potential because:
They do not include all key nations; do not specifically address sea
turtle conservation; are handicapped by the lack of a sovereign
authority to promote enforcement; and/or lack of legally-binding
requirements. Lack of implementation or enforcement by some nations may
make them less effective than if they were implemented in a more
consistent manner across the target region. A thorough discussion of
this topic is available in the 2002 special issue of the Journal of
International Wildlife Law and Policy: International Instruments and
Marine Turtle Conservation (Hykle 2002). Additional information on
national, regional, and local protection is provided in the prior
sections of this finding relating to each individual DPS.
In summary, numerous regulatory mechanisms protect leatherback
turtles, eggs, and nesting habitat throughout the range of the species.
Although the regulatory mechanisms provide some protection, many do not
adequately reduce the threat that they were designed to address,
generally as a result of limited implementation or enforcement. As a
result, bycatch, incomplete nesting habitat protection, and poaching
remain threats to the species. We conclude that the inadequacy of the
regulatory mechanisms is a threat to the leatherback turtle.
Fisheries Bycatch
Fisheries bycatch is the primary threat to leatherback turtles
(Crowder 2000; Spotila et al. 2000; Lewison et al. 2004; Wallace et al.
2011; Wallace et al. 2013; Angel et al. 2014). It is a primary threat
to all DPSs. Leatherback turtles are susceptible to bycatch in a wide
range of fisheries, from large scale commercial to artisanal. Gear
types that affect leatherbacks include: longlines, purse seines,
driftnets, gillnets, trawls, pots/traps, and pound nets (Gray and Diaz
2017). Turtles often drown after becoming entangled in nets and other
gear or become injured and possibly die as a result of hooking or
interactions with the gear. While bycatch in pelagic shallow-set
swordfish longline fisheries has received the most attention to date,
small-scale coastal fisheries occur worldwide, employing over 99
percent of the world's 51 million fishers (FAO 2011).
Bycatch data are most commonly collected by trained observers on
fishing vessels or via surveys or interviews (Lewison et al. 2015).
Though often the best available data on bycatch, observer data
generally cover less than five percent of fisheries' total effort
(Finkbeiner et al. 2011) and are rarely available for small-scale
fisheries (Wallace et al. 2013; Lewison et al. 2015). The use of
different metrics also makes the data difficult to compare
[[Page 48413]]
among fisheries, gear types, and regions (Lewison et al. 2015).
Therefore, estimates of bycatch and resulting mortality often
underestimate the magnitude of this threat.
Furthermore, IUU fishing is a significant yet unquantified threat
to sea turtles worldwide. In addition to killing and injuring turtles,
it undermines national and regional efforts to estimate fisheries
bycatch. IUU fishing represents up to 26 million tonnes of fish caught
annually (https://www.fao.org/iuu-fishing/en/). We have no estimates of
the impacts to leatherback turtles from IUU fishing, though interaction
and mortality rates are likely high because of the magnitude of this
additional fishing pressure and because it is unregulated.
Generally, leatherback turtles do not attempt to consume the bait
associated with fishing gear, as other sea turtles do, but become
entangled in fishing gear (Lewison et al. 2015). Longline fisheries
involve the deployment of a horizontal main line and vertical
branchlines with baited hooks, which may entangle leatherback turtles.
Bycatch reduction measures include using circle hooks, finfish bait,
minimizing soak times, and limiting mainline length (Angel et al. 2014;
https://www.fisheries.noaa.gov/national/bycatch/fishing-gear-pelagic-longlines#risks-to-sea-turtles). Purse seines capture schools of fish
in a vertical wall of netting that can be closed at the bottom (https://www.fisheries.noaa.gov/national/bycatch/fishing-gear-purse-seines);
bycatch rates are generally much lower than longline bycatch rates
(Angel et al. 2014). Leatherback turtles also become entangled and
drowned in drift or set gillnets (https://www.fisheries.noaa.gov/national/bycatch/fishing-gear-gillnets). Gillnets can be devastating to
leatherback populations when set near nesting beaches and represent the
primary threat to leatherback turtles in some areas (e.g., Trinidad;
Eckert and Eckert 2005). Trawl fisheries drag nets along the substrate
or through the water column and can capture and drown sea turtles.
Although TEDs may mitigate this threat, they are not always required or
used in all areas. Vertical lines extending and/or connecting pot and
trap gear with surface buoys commonly entangle and can kill leatherback
turtles.
Longline and net fisheries are often the greatest threats to
leatherback turtles. In a global study of sea turtle bycatch, Wallace
et al. (2013) compiled data (n = 239 records) published between 1990
and 2011 to compare gear types (longline, net, and trawl) and their
impacts to leatherback RMUs, which are similar to the DPSs discussed in
this rule, though their exact boundaries differ. Wallace et al. (2013)
defined high bycatch impact as follows: A weighted median bycatch per
unit effort (BPUE) greater than or equal to one; median mortality rate
greater than or equal to 0.5; and affecting adult or subadult turtles.
They found that longline bycatch had a high impact on SW Atlantic, SE
Atlantic, and SW Indian RMUs and that net bycatch had a high impact on
the NW Atlantic and East Pacific RMUs (Wallace et al. 2013).
Integrating catch data from over 40 nations and bycatch data from
13 international observer programs, Lewison et al. (2004) estimated the
numbers of leatherback turtles taken by pelagic longliners to be more
than 50,000 leatherback turtles in just one year (2000). With over half
of the total fishing effort (targeting tuna and swordfish) occurring in
the Pacific Ocean, an estimated 20,000 leatherback turtles interacted
with longline fishing gear, with 1,000 to 3,200 mortalities in 2000
(Lewison et al. 2004). However, Beverly and Chapman (2007) estimated
sea turtle longline bycatch mortality to be approximately 20 percent of
that estimated by Lewison et al. (2004), or approximately 200 to 640
leatherback turtle mortalities annually. We consider the estimate of
Beverly and Chapman (2007) to be more realistic, considering the low
nesting females abundance of Pacific leatherback turtles, and because
Beverly and Chapman (2007) combined the effort data from Lewison et al.
(2004) with bycatch data from Molony (2005) that differentiated between
deep-set and shallow-set fisheries (which have different interaction
rates).
In the Pacific Ocean, Roe et al. (2014) predicted leatherback
turtle bycatch hotspots by comparing the satellite tracks of 135 adult
turtles with longline fishing effort. The greatest bycatch risk
occurred adjacent to primary nesting beaches of the West Pacific DPS.
Bycatch risk was also high in the South Pacific Gyre, where the East
Pacific DPS forages. Expanding on this study, a study of observer data
from 34 swordfish-targeting shallow-set longline fleets found there
were 331 leatherback turtle interactions between 1989 and 2015 (Clarke
2017). Clarke (2017) identified two bycatch hotspot areas: Central
North Pacific Ocean and eastern Australia (Clarke 2017).
In the Atlantic Ocean, Fossette et al. (2014) compared leatherback
telemetry data to longline fishing effort data from ICCAT to identify
nine areas in which leatherback turtles are exposed to bycatch
associated with high longline fishery pressure. The high pressure
fishing areas include foraging areas in the North and South Atlantic
Ocean and in waters off Brazil and western Africa. These high pressure
fishing areas are not comparable to those identified by Roe et al.
(2014), who used a different methodology, but both studies identify
high risk areas within each ocean basin.
Additional bycatch information that we have set out in prior
sections specific to each DPS applies to our consideration of the risk
to the overall species. In summary, fisheries bycatch is a threat that
is encountered by numerous juvenile and adult leatherback turtles.
Mortality rates are often high, and individuals that are released may
experience injuries or sublethal effects associated with entanglement,
submergence, or handling. Fisheries bycatch reduces abundance, and when
it prevents nesting females from returning to nesting beaches, reduces
productivity as well. Fisheries bycatch is the primary threat to the
leatherback species.
Vessel Strikes
Vessel strikes pose a threat to the species throughout its range.
As mature individuals move from oceanic foraging areas into coastal
waters to reproduce, they are exposed to a greater concentration of
vessels. Vessel strikes off nesting beaches may injure or kill these
individuals, reducing the abundance and productivity of the DPS. Most
vessel strikes likely go unnoticed or unreported, making this threat
potentially much more significant that documented occurrences would
suggest. Vessel strikes are a threat to the leatherback species.
Pollution
We define pollution as including contaminants, marine debris, and
ghost or derelict fishing gear. Such interactions are likely to go
unnoticed and unreported and thus likely present a more significant
impact than documented occurrences would suggest. Leatherback turtles
of all life stages are vulnerable to oil spills, on land and at sea,
where exposure to oil and dispersants occurs via contact (i.e.,
physical fouling), inhalation, or ingestion (reviewed by Stacy et al.
in press).
Marine debris is ubiquitous throughout the range of the species.
Marine debris includes plastics (including plastic bags),
microplastics, derelict fishing gear (e.g., ghost nets and other
discarded or lost gear), and other man-made materials. Leatherback
turtles may directly consume floating plastics, mistaking it for their
gelatinous prey or accidentally ingest plastics while foraging. In
particular, plastic bags appear similar to jellyfish in the marine
[[Page 48414]]
environment, inappropriately triggering the sensory cue to feed
(Schuyler et al. 2014; Nelms et al. 2016). Plastic bags have been found
during necropsy of stranded leatherback turtles, and phthalates derived
from plastics have been found in leatherback egg yolk (Lebreton et al.
2018). Mrosovsky et al. (2009) reviewed 408 necropsy records from 1885
to 2007 and found evidence of plastic in the gastrointestinal tract of
34 percent of leatherback turtles, including some cases in which the
plastic obstructed the passage of food through the gut. The most
commonly identified items were plastic bags, fishing lines, twine, and
fragments of mylar balloons. Ghost or derelict fishing gear include
discarded or lost nets, line, and other gear. Ghost fishing gear can
drift in the ocean and fish unattended for decades and kill numerous
individuals (Wilcox et al. 2013). The main sources of ghost fishing
gear are gillnet, purse seine, and trawl fisheries (Stelfox et al.
2016). Marine debris affects leatherback turtles via ingestion or
entanglement and can reduce food intake and digestive capacity, cause
distress and/or drowning, expose turtles to contaminants, and in some
cases cause direct mortality (Mrosovsky et al. 2009; NMFS and USFWS
2013). In terms of microplastics, all samples analyzed from all species
(including leatherbacks) had microplastics evident in their gastro-
intestinal tracts (Duncan et al. 2018). Given the increase of pollution
entering the marine environment over the past 30 years or approximately
5.2 to 19.3 million tonnes per year (Lebreton et al. 2018), we conclude
that pollution is a threat to the species.
Natural Disasters and Oceanographic Regime Shifts
Leatherback turtles are susceptible to the impacts of natural
disasters and oceanographic regime shifts as a result of their nesting
and foraging preferences. Nesting usually occurs on high-energy beaches
that are inherently unstable (Pritchard 1976) and which are susceptible
to natural erosion. The primary factors influencing shoreline
suitability for nesting appear to be a lack of abrasive substrate
material, a deep-water approach to minimize energy expenditure needed
to reach nesting sites, and proximity to oceanic currents that can
facilitate hatchling dispersal (Eckert et al. 2012). Leatherback
turtles nest lower on the beach than other species, exposing their
nests to erosion and inundation. Storm events, King Tides, tsunamis,
and hurricanes can destroy or modify preferred nesting beaches of some
DPSs.
Gelatinous prey have relatively low energy content, requiring
leatherback turtles to consume large quantities to meet metabolic
demands (Heaslip et al. 2012; Jones et al. 2012). Leatherback turtles
likely maximize their caloric intake by aligning their foraging
behavior to prey distribution abundance. Foraging areas are generally
characterized by zones of upwelling, including off the edges of
continents, where major currents converge, and in deep-water eddies
(Saba 2013). Some of these areas experience oceanographic regime shifts
that alter water temperature, downwelling, Ekman upwelling, sea surface
height, chlorophyll-a concentration, and mesoscale eddies (Bailey et
al. 2013; Benson et al. 2011). These shifts alter prey availability,
and thus productivity parameters (e.g., remigration rates, clutch size,
and clutch frequency), for leatherback turtles. Some DPSs are not
affected by such shifts because they have access to diverse foraging
areas, such as: coastal and pelagic waters; subtropical, temperate, and
boreal waters; and ephemeral eddies (Neeman et al. 2015). Such
flexibility allows the leatherback turtle to consume large amounts of
prey at various locations throughout the year.
We conclude that natural disasters and oceanographic regime shifts
are threats to the species, affecting some but not all populations,
depending on the location of nesting and foraging areas. These threats
reduce productivity by reducing nesting, nesting habitat, and nest and
hatching success.
Climate Change
Climate change is a threat that affects leatherback turtles of all
life stages and within all DPSs. A warming climate and rising sea
levels can impact leatherback turtles through changes in beach
morphology, increased sand temperatures leading to a greater incidence
of lethal incubation temperatures, changes in hatchling sex ratios, and
the loss of nests or nesting habitat due to beach erosion (Benson et
al. 2013).
Impacts from climate change, especially due to global warming, are
already being observed and are likely to become more apparent in future
years (IPCC 2007a). In its Fifth Assessment Report, the IPCC (2014)
stated that the globally averaged combined land and ocean surface
temperature data has shown a warming of 0.85 [deg]C from 1880 to 2012.
The mean rate of globally averaged sea level rise was 1.7 millimeters
annually between 1901 and 2010, 2.0 millimeters annually between 1971
and 2010, and 3.2 millimeters annually between 1993 and 2010. Climate
model projections exhibit a wide range of plausible scenarios for both
temperature and precipitation over the next several decades. The global
mean surface temperature change for the period 2016 to 2035 relative to
1986 to 2005 will likely be in the range of 0.3 [deg] to 0.7 [deg]C
(medium confidence; IPCC 2014). The global ocean temperature will
continue to warm, and increases in seasonal and annual mean surface
temperatures are expected to be larger in the tropics and Northern
Hemisphere subtropics (i.e., where leatherback turtles nest; IPCC
2014). Under Representative Concentration Pathway 8.5, the change in
global mean sea level rise for the mid- and late 21st century relative
to the reference period of 1986 to 2005 is projected to be 0.30 meters
higher from 2046 to 2065 and 0.63 meters higher from 2081 to 2100, with
a rate of sea level rise during 2081 to 2100 of 8 to 16 millimeters
annually (medium confidence; IPCC 2014).
For all sea turtles, including leatherback turtles, a warming
climate and rising sea levels are likely to result in changes in beach
morphology, increased sand temperatures leading to a greater incidence
of lethal incubation temperatures, changes in hatchling sex ratios, and
the loss of nests and nesting habitat due to beach erosion (Benson et
al. 2015; Hamann et al. 2013). Leatherback turtles are most likely to
be affected by climate change at nesting beaches due to warming
temperatures, sea level rise, and storm events and due to oceanic
changes that are likely to alter foraging and migration. Warming
temperatures and increased precipitation at nesting beaches affect
reproductive output including hatching success, hatchling emergence
rate, and hatchling sex ratios (e.g., Hawkes et al. 2009). Sea level
rise results in a reduction or shift in available nesting beach
habitat, an increased risk of erosion and nest inundation (e.g., Boyes
et al. 2010), and reduced nest success (Fish et al. 2005; Fuentes et
al. 2010; Fonseca et al. 2013). Increased frequency and severity of
storm events impact nests and nesting habitat, thus reducing nesting
and hatching success (e.g., Van Houtan and Bass 2007; Fuentes and Abbs
2010). Changes in productivity affect the abundance and distribution of
forage species, resulting in changes in the foraging behavior and
distribution of leatherback turtles (e.g., Saba et al. 2008, 2012) as
well as changes in leatherback fitness and growth. Changes in water
temperature lead to a shift in range and changes in phenology (timing
of nesting seasons,
[[Page 48415]]
timing of migrations) and different threat exposure (e.g., Saba et al.
2008, 2012).
Increasing sand temperatures will alter the thermal regime of
incubating nests, resulting in altered sex ratios and reduced hatching
output (Hawkes et al. 2009). Leatherback turtles exhibit temperature-
dependent sex determination (reviewed by Binckley and Spotila 2015),
whereby phenotypic sex is determined by temperatures experienced during
the thermosensitive period of egg incubation. A 1:1 sex ratio is
produced when this pivotal temperature lies between 29.2 and 30.4
[deg]C for leatherback turtles in Malaysia, 29.2 and 29.8 [deg]C in
French Guiana/Suriname, and 29.2 and 29.5 [deg]C in Pacific Costa Rica
(Binckley and Spotila 2015). Warmer temperatures produce more female
embryos (Mrosovsky et al. 1984; Hawkes et al. 2007), but temperatures
over 32 [deg]C are likely to result in death. As temperatures continue
to increase, emergence rates decrease (Santidri[aacute]n Tomillo et al.
2015), removing any advantage of increased female production.
Santidri[aacute]n Tomillo et al. (2015) conclude that leatherback
turtles may not survive if temperatures rise as projected by current
climate change models. Increases in precipitation might temporarily
reduce the temperatures at some nesting beaches thereby mitigating some
impacts relative to increasing sand temperatures.
Beach erosion and nest inundation already threaten leatherback
nesting habitat globally. Sea level rise is likely to increase the
number of nests lost to erosion and inundation. Such loss of nests is
especially problematic in areas prone to storm events, which are likely
to increase in intensity and duration, and in areas where coastal
development impedes natural shoreline migration.
Climate change is also likely to alter the productivity in some
marine environments, which could affect leatherback prey availability.
With reports on the increasing incidence of jellyfish blooms in some
locations, there is the perception that jellyfish abundance is
increasing globally (Condon et al. 2012), which could result in more
prey for leatherback turtles (Hawkes et al. 2009). However, after
analyzing all available long-term datasets on jellyfish abundance,
Condon et al. (2012) found that there is no robust evidence for a
global increase in jellyfish. Rather, jellyfish populations undergo
larger, worldwide oscillations with an approximate 20-year periodicity
(Condon et al. 2012). Additional monitoring is needed to determine
whether the weak linear trend in jellyfish abundance since 1970
represents an actual increase or is a phase of an oscillation (Condon
et al. 2012). Therefore, the effects of climate change on productivity
are uncertain.
As described in prior sections with respect to each individual
population, some impacts from climate change have already been
observed. At several nesting beaches, increased erosion occurs, and sex
ratios are severely skewed toward females. Beach erosion reduces
productivity. Although the skew toward females could increase
productivity in the short-term, it is often correlated with low
hatching success. For these reasons, climate change is a threat to the
species.
Conservation Efforts
The ESA requires the Services to make their listing determinations
solely on the basis of the best scientific and commercial data
available, after conducting a status review, and after taking into
account those efforts, if any, being made by any State or foreign
nation to protect the species, whether by predatory control, protection
of habitat and food supply, or other conservation practices, within any
area under its jurisdiction, or on the high seas (16 U.S.C. 1533
(b)(1)(A)). In addition, the Services published a policy for the
evaluation of domestic conservation efforts which have yet to be
implemented or to show effectiveness (68 FR 15100; March 28, 2003). We
did not identify any conservation efforts that required such evaluation
for leatherbacks (i.e., the conservation efforts reviewed are
international in nature or have already been implemented to a
sufficient degree that they have a track record of being effective or
not being effective). Several conservation efforts have been previously
discussed in prior sections evaluating regulatory mechanisms with
respect to each DPS. Therefore, the list below describes only those
conservation efforts that have not been previously discussed and that
apply generally to the leatherback species rather than being clearly
associated with a particular population. We considered these efforts
prior to making our listing determination. After reviewing these
efforts, we concluded that they have been somewhat effective, in that
they have prevented this endangered species from going extinct.
However, these efforts have not reduced the threats to a level at which
protections under the ESA are no longer necessary.
African Convention on the Conservation of Nature and Natural
Resources (Algiers Convention): Adopted in September 1968, the
contracted states were ``to undertake to adopt the measures necessary
to ensure conservation, utilization and development of soil, water,
floral and faunal resources in accordance with scientific principles
and with due regard to the best interests of the people.'' The Algiers
Convention recently has undergone revision, and its objectives are to
enhance environmental protection, foster conservation and sustainable
use of natural resources, and harmonize and coordinate policies in
these fields with a view to achieving ecologically rational,
economically sound, and socially acceptable development policies and
programs. Additional information is available at https://www.unep.ch/regionalseas/legal/afr.htm.
Atlantic Sea Turtle Network (ASO): Created in 2003 to foster
greater collaboration in southern Brazil, Uruguay, and Argentina for
the protection of sea turtles and their habitats. ASO represents dozens
of local and regional NGOs and government agencies as well as hundreds
of community members. ASO and its partners have significantly advanced
policies to protect sea turtles from fisheries interactions, which is
one of the most severe threats in the region. Brazil plays a major role
in South American (and global) sea turtle conservation and research,
and it serves as an example to other countries. Projeto TAMAR, a
partnership of the Centro TAMAR/ICMBio, government agencies, and
Fundac[atilde]o Pr[oacute] TAMAR, has been active since 1980. Today,
the group carries out sea turtle research and conservation from 22
stations on the coast and the offshore islands of Brazil. Another NGO
based in the southern Brazilian state of Rio Grande do Sul, called NEMA
has been collecting systematic sea turtle stranding data since 1990.
Those data have been instrumental to conservation efforts in Brazil and
have shown that southern Brazil has the highest stranding rates for
loggerheads in the western Atlantic Ocean.
Association of Southeast Asian Nations (The ASEAN) Ministers on
Agriculture and Forestry (AMAF): A Memorandum of Understanding (MoU) on
ASEAN sea turtle conservation was created in 1999. From this, a Sea
Turtle Conservation and Protection Program and Work plan has developed;
research and monitoring activities have also been produced regionally
(Kadir 2000). The objectives of this Memorandum of Understanding,
initiated by ASEAN, are to promote the protection, conservation,
replenishing, and recovery of sea turtles and their habitats based on
the best
[[Page 48416]]
available scientific evidence, taking into account the environmental,
socio-economic and cultural characteristics of the Parties. It
currently has nine signatory states in the South East Asian Region
(https://document.seafdec.or.th/projects/2012/seaturtles.php).
Andaman and Nicobar Island Environmental Team (ANET): A division of
the Centre for Herpetology/Madras Crocodile Bank Trust has been
conducting surveys and monitoring since 1991. Over the last few years,
conservation and monitoring of sea turtles in these islands has been
carried by Dakshin Foundation and Indian Institute of Science in
collaboration with ANET, centered around a leatherback monitoring
program on Little Andaman Island. A multi- institution stakeholder
platform for marine conservation, including government and non-
governmental agencies, was established by these groups to facilitate
the conservation of marine turtles and other endangered species
(Tripathy et al. 2012). The Trust, along with the Wildlife Institute of
India and Ministry of Environment and Forests, produced a series of
manuals on sea turtle conservation, management and research to help
forest officers, conservationists, NGOs and wildlife enthusiasts
conduct sea turtle conservation and research programs (ANET, 2003 as
cited in Shanker and Andrews 2004). A consolidated manual has been
produced to achieve these goals by Dakshin Foundation and the Trust
(Tripathy et al. 2012).
Central American Regional Network: This collaborative effort
created the national sea turtle network in each country of the region,
as well as the development of first hand tools, such as a regional
diagnosis, a 10-year strategic plan, a manual of best practices, and
four regional training and information workshops for people in the
region (e.g., Chac[oacute]n and Arauz, 2001). This initiative is
managed by stakeholders in various sectors (private, non-governmental
and governmental) across the region.
Convention on the Conservation of Migratory Species of Wild Animals
(CMS): This Convention, also known as the Bonn Convention or CMS, is an
international treaty that focuses on the conservation of migratory
species and their habitats. As of December 2018, the Convention had 127
Parties, including Parties from Africa, Central and South America,
Asia, Europe, and Oceania. While the Convention has successfully
brought together about half the countries of the world with a direct
interest in sea turtles, it has yet to realize its full potential
(Hykle 2002). Its membership does not include a number of key
countries, including Canada, China, Indonesia, Japan, Mexico, Oman, and
the United States. Under the CMS, two Memoranda of Understanding (MOUs)
apply to leatherback turtles: The MOU concerning Conservation Measures
for Marine Turtles of the Atlantic Coast of Africa and the MOU on the
Conservation and Management of Marine Turtles and their Habitats of the
Indian Ocean and South-East Asia. Additional information is available
at https://www.cms.int.
Convention on Biological Diversity (CBD): The primary objectives of
this international treaty are: (1) The conservation of biological
diversity, (2) the sustainable use of its components, and (3) the fair
and equitable sharing of the benefits arising out of the utilization of
genetic resources. This Convention has been in force since 1993 and had
193 Parties as of March 2013. While the Convention provides a framework
within which are broad conservation objectives, it does not
specifically address sea turtle conservation (Hykle 2002). Additional
information is available at https://www.cbd.int.
Convention on International Trade in Endangered Species of Wild
Fauna and Flora (CITES): Known as CITES, this Convention was designed
to regulate international trade in a wide range of wild animals and
plants. CITES was implemented in 1975 and currently has 183 Parties.
Although CITES has been effective at minimizing the international trade
of sea turtle products, it does not limit legal harvest within
countries, nor does it regulate intra-country commerce of sea turtle
products (Hykle, 2002). The leatherback turtle is included (since 1977)
in CITES Appendix I, which bans trade, including individuals and
products, except as permitted for exceptional circumstances, not to
include commercial purposes (Lyster 1985). Additional information is
available at https://www.cites.org.
Convention on the Conservation of European Wildlife and Natural
Habitats: Also known as the Bern Convention, the goals of this
instrument are to conserve wild flora and fauna and their natural
habitats, especially those species and habitats whose conservation
requires the cooperation of several States, and to promote such
cooperation. The Convention was enacted in 1982 and currently includes
51 European and African States and the European Union. Additional
information is available at https://www.coe.int/t/dg4/cultureheritage/nature/bern/default_en.asp.
Convention for the Co-operation in the Protection and Development
of the Marine and Coastal Environment of the West and Central African
Region (Abidjan Convention): The Abidjan Convention covers the marine
environment, coastal zones, and related inland waters from Mauritania
to Namibia. The Abidjan Convention countries are Angola, Benin,
Cameroon, Cape Verde, Congo, Cote d'Ivoire, Democratic Republic of
Congo, Equatorial Guinea, Gabon, Gambia, Ghana, Guinea, Guinea-Bissau,
Liberia, Mauritania, Namibia, Nigeria, Sao Tome and Principe, Senegal,
Sierra Leone, and Togo. The Abidjan Convention is an agreement for the
protection and management of the marine and coastal areas that
highlights sources of pollution, including pollution from ships,
dumping, land-based sources, exploration and exploitation of the sea-
bed, and pollution from or through the atmosphere. The Convention also
identifies where co-operative environmental management efforts are
needed. These areas of concern include coastal erosion, specially
protected areas, combating pollution in cases of emergency and
environmental impact assessment.
Convention for the Protection Management and Development of the
Marine and Coastal Environment of the Eastern African Region (Nairobi
Convention): The Nairobi Convention was signed in 1985 and came into
force in 1996. This instrument ``provides a mechanism for regional
cooperation, coordination and collaborative actions, and enables the
Contracting Parties to harness resources and expertise from a wide
range of stakeholders and interest groups towards solving interlinked
problems of the coastal and marine environment.'' Parties are
responsible for ``the conservation and wise management of the sea
turtle populations frequenting their waters and shores [and] agree to
work closely together to improve the conservation status of the sea
turtles and the habitats upon which they depend.'' The Western Indian
Ocean-Marine Turtle Task Force, which was created under the Nairobi
Convention and the IOSEA, plays a role in sea turtle conservation. This
is a technical, non-political working group comprised of specialists
from eleven countries: Comoros, France (La R[eacute]union), Kenya,
Madagascar, Mauritius, Mozambique, Seychelles, Somalia, South Africa,
United Kingdom and Tanzania, as well as representatives from inter-
governmental organizations, academic, and non-governmental
organizations within the region. Additional information is available at
https://www.unep.org/NairobiConvention.
[[Page 48417]]
Convention for the Protection of the Marine Environment of the
North-East Atlantic: Also called the OSPAR Convention, this 1992
instrument combines and updates the 1972 Oslo Convention against
dumping waste in the marine environment and the 1974 Paris Convention
addressing marine pollution stemming from land-based sources. The
convention is managed by the OSPAR Commission, which is comprised of
representatives from 15 signatory nations (Belgium, Denmark, Finland,
France, Germany, Iceland, Ireland, Luxembourg, The Netherlands, Norway,
Portugal, Spain, Sweden, Switzerland, and United Kingdom), as well as
the European Commission, representing the European Community. The
mission of the OSPAR Convention ``. . . is to conserve marine
ecosystems and safeguard human health in the North-East Atlantic by
preventing and eliminating pollution; by protecting the marine
environment from the adverse effects of human activities; and by
contributing to the sustainable use of the seas.'' Leatherback turtles
are included on the OSPAR List of Threatened and/or Declining Species
and Habitats, used by the OSPAR Commission for setting priorities for
work on the conservation and protection of marine biodiversity.
Additional information is available at https://www.ospar.org.
Convention for the Protection and Development of the Marine
Environment of the Wider Caribbean Region: Also called the Cartagena
Convention, this instrument that benefits turtles of the Northwest
Atlantic leatherback DPS, has been in place since 1986 and currently
has 38 member states and territories. Under this Convention, the
component that relates to leatherback turtles is the Protocol
Concerning Specially Protected Areas and Wildlife (SPAW) that has been
in place since 2000. The goals are to encourage Parties ``to take all
appropriate measures to protect and preserve rare or fragile
ecosystems, as well as the habitat of depleted, threatened or
endangered species, in the Convention area.'' The SPAW protocol has
partnered with WIDECAST to develop a program of work on sea turtle
conservation, which has helped many of the Caribbean nations to
identify and prioritize their conservation actions through Sea Turtle
Recovery Action Plans. Each recovery action plan summarizes the known
distribution of sea turtles, discusses major causes of mortality,
evaluates the effectiveness of existing conservation laws, and
prioritizes implementing measures for stock recovery. The objective of
the recovery action plan series is not only to assist Caribbean
governments in the discharge of their obligations under the SPAW
Protocol, but also to promote a regional capability to implement
science-based sea turtle management and conservation programs.
Additional information is available at https://www.cep.unep.org/about-cep/spaw.
Convention on Nature Protection and Wildlife Preservation in the
Western Hemisphere (Washington or Western Hemisphere Convention):
Elements of the Convention include the protection of species from
human-induced extinction, the establishment of protected areas, the
regulation of international trade in wildlife, special measures for
migratory birds and stressing the need for co-operation in scientific
research and other fields are all elements of wildlife conservation.
Additional information is available at https://www.oas.org/juridico/english/treaties/c-8.html.
Convention for the Protection of the Marine Environment and Coastal
Area of the South-East Pacific (Lima Convention): This Convention's
signatories include all countries along the Pacific Rim of South
America from Panama to Chile. Among other resource management
components, this Convention established protocol for the conservation
and management of protected marine resources. Stemming from this
Convention is the Commision Permanente del Pacifico Sur (CPPS) that has
developed a Marine Turtle Action Plan for the Southeast Pacific that
outlines a strategy for protecting and recovering marine turtles in
this region. Convention for the Protection of the Natural Resources and
Environment of the South Pacific Region (Noumea Convention): This
Convention has been in force since 1990 and currently includes 26
Parties. The purpose of the Convention is to protect the marine
environment and coastal zones of the South-East Pacific within the 200-
mile area of maritime sovereignty and jurisdiction of the Parties and,
beyond that area, the high seas up to a distance within which pollution
of the high seas may affect that area. Additional information is
available at https://www.unep.org/regionalseas/programmes/nonunep/pacific/instruments/default.asp.
Convention Concerning the Protection of the World Cultural and
Natural Heritage (World Heritage Convention): The World Heritage
Convention was signed in 1972 and, as of November 2007, 185 states were
parties to the Convention. The instrument requires parties to take
effective and active measures to protect and conserve habitat of
threatened species of animals and plants of scientific or aesthetic
value. The World Heritage Convention currently includes 31 marine
sites. Additional information is available at https://whc.unesco.org/en/conventiontext.
Convention for the Conservation and Management of Highly Migratory
Fish Stocks in the Western and Central Pacific Ocean (WCPF Convention):
The convention entered into force on 19 June 2004. The WCPF Convention
draws on many of the provisions of the UN Fish Stocks Agreement [UNFSA]
while, at the same time, reflecting the special political, socio-
economic, geographical and environmental characteristics of the western
and central Pacific Ocean (WCPO) region. The WCPFC Convention seeks to
address problems in the management of high seas fisheries resulting
from unregulated fishing, over-capitalization, excessive fleet
capacity, vessel re-flagging to escape controls, insufficiently
selective gear, unreliable databases and insufficient multilateral
cooperation in respect to conservation and management of highly
migratory fish stocks.
Convention for the Prohibition of Fishing with Long Driftnets in
the South Pacific: This regional convention, also known as the
Wellington Convention, was adopted in 1989 in Wellington, New Zealand,
and entered into force in 1991. The objective of the Convention is ``to
restrict and prohibit the use of drift nets in the South Pacific region
in order to conserve marine living resources.'' Additional information
is available at https://www.mfat.govt.nz/Treaties-and-International-Law/01-Treaties-for-which-NZ-is-Depositary/0-Prohibition-of-Fishing.php.
Eastern Pacific Leatherback Network: Also known as La Red de la
Tortuga La[uacute]d del Oc[eacute]ano Pacifico (La[uacute]d OPO)
(www.savepacificleatherbacks.org) was formed to address the critical
need for regional coordination of East Pacific leatherback conservation
actions necessary to track conservation priorities and progress at the
population level. Led by Fauna & Flora International, this network has
brought together conservationists, researchers, practitioners and
government representatives from 22 institutions across nine East
Pacific countries with varying priorities, capacities and historical
experiences in leatherback research and conservation to contribute to
shared activities, projects, and goals. Through these efforts,
La[uacute]d now has mutually-agreed upon mechanisms for sharing
information and data, as well as
[[Page 48418]]
standardized protocols for nesting beach monitoring and bycatch
assessments/fishing practices.
The Eastern Tropical Pacific Marine Corridor (CMAR) is a regional
and cross-border initiative for the conservation and sustainable use of
the region's marine and coastal resources. Its objective is to
sustainably manage biodiversity through ecosystem based management and
the development of regional intergovernmental strategies with support
of non-governmental organizations and international cooperation
agencies.
United Nations' Food and Agricultural Organization (FAO) Technical
Consultation on Sea Turtle-Fishery Interactions: While not a true
international instrument for conservation, the 2004 FAO of the UN's
technical consultation on sea turtle-fishery interactions was
groundbreaking in that it solidified the commitment of the lead UN
agency for fisheries to reduce sea turtle bycatch in marine fisheries
operations. Recommendations from the technical consultation were
endorsed by the FAO Committee on Fisheries (COFI) and called for the
immediate implementation by member nations and Regional Fishery
Management Organizations (RFMOs) of guidelines to reduce sea turtle
mortality in fishing operations, developed as part of the technical
consultation. Currently, all five of the tuna RFMOs call on their
members and cooperating non-members to adhere to the 2009 FAO
``Guidelines to Reduce Sea Turtle Mortality in Fishing Operations,''
which describes all the gear types sea turtles could interact with and
the latest mitigation options. The Western and Central Pacific
Fisheries Commission (https://www.wcpfc.int) has the most protective
measures (CMM 2008-03), which follow the FAO guidelines and ensure safe
handling of all captured sea turtles. Fisheries deploying purse seines,
to the extent practicable, must avoid encircling sea turtles and
release entangled turtles from fish aggregating devices. Longline
fishermen must carry line cutters and use dehookers to release sea
turtles caught on a line. Longliners must either use large circle
hooks, whole finfish bait, or mitigation measures approved by the
Scientific Committee and the Technical and Compliance Committee.
Inter-American Tropical Tuna Convention (IATTC) has enacted a
resolution to mitigate the impact of tuna fishing vessels on sea
turtles by reducing bycatch, injury, and mortality of sea turtles. The
IATTC has also developed a memorandum of understanding with the IAC.
For more information, see https://www.iattc.org/PDFFiles/Resolutions/IATTC/_English/C-07-03-Active_Sea%20turtles.pdf.
The International Commission for the Conservation of Atlantic Tunas
(ICCAT) has adopted a resolution for the reduction of sea turtle
mortality (Resolution 03-11), encouraging States to submit data on sea
turtle interactions, release sea turtles alive wherever possible, and
conduct research on mitigation measures. It calls for implementing the
FAO Guidelines for sea turtles, avoiding encirclement of sea turtles by
purse seiners, safely handling and releasing sea turtles, and reporting
on interactions. The Commission does not have any specific gear
requirements applicable to longline fisheries. ICCAT is currently
undertaking an ecological risk assessment to better understand the
impact of its fisheries on sea turtle populations. For more information
see https://www.iattc.org/. Other international fisheries organizations
that may influence leatherback turtle recovery include the Southeast
Atlantic Fisheries Organization (https://www.seafo.org) and the North
Atlantic Fisheries Organization (https://nafo.int). These organizations
regulate trawl fisheries in their respective Convention areas. Given
that sea turtles can be incidentally captured in these fisheries, both
organizations have sea turtle resolutions calling on their Parties to
implement the FAO Guidelines on sea turtles as well as to report data
on sea turtle interactions.
The Indian Ocean Tuna Commission (IOTC) is playing an increased
role in turtle conservation. Resolution 05/08, superseded by Resolution
09/06 on Sea Turtles, sets out reporting requirements related to
interactions with sea turtles and accordingly provides an executive
summary per species for adoption at the Working Party on Ecosystem and
By-catch and then subsequently at the Scientific Committee. In 2011,
IOTC developed a ``Sea Turtle Identification Card'' to be distributed
to all longliners operating in the Indian Ocean (www.iotc.com). In
2012, the Indian Ocean Tuna Commission (IOTC) began requiring its 31
contracting Parties to report sea turtle bycatch and to use safe
handling and release techniques for sea turtles on longline vessels.
Indian Ocean--South-East Asian Marine Turtle Memorandum of
Understanding (IOSEA): Under the auspices of the Convention of
Migratory Species, the IOSEA memorandum of understanding provides a
mechanism for States of the Indian Ocean and South-East Asian region,
as well as other concerned States, to work together to conserve and
replenish depleted marine turtle populations. This collaboration is
achieved through the collective implementation of an associated
Conservation and Management Plan. Currently, there are 33 Signatory
States. The United States became a signatory in 2001. The IOSEA has an
active sub-regional group for the Western Indian Ocean, which has
improved collaboration amongst sea turtle conservationists in the
region. Further, the IOSEA website provides reference materials,
satellite tracks, on-line reporting of compliance with the Convention,
and information on all international mechanisms currently in place for
the conservation of sea turtles. Finally, at the 2012 Sixth Signatory
of States meeting in Bangkok, Thailand, the Signatory States agreed to
procedures to establish a network of sites of importance for sea
turtles in the IOSEA region (https://www.ioseaturtles.org).
Inter-American Convention for the Protection and Conservation of
Sea Turtles (IAC): This Convention is the only legally binding
international treaty dedicated exclusively to sea turtles and sets
standards for the conservation of these endangered animals and their
habitats with a large emphasis on bycatch reduction. The Convention
area is the Pacific and the Atlantic waters of the Americas. Currently,
there are 15 Parties. The United States became a Party in 1999. The IAC
has worked to adopt fisheries bycatch resolutions, carried out
workshops on Caribbean sea turtle conservation, and established
collaboration with other agreements such as the Convention for the
Protection and Development of the Marine Environment of the Wider
Caribbean Region and the International Commission for the Conservation
of Atlantic Tunas. Additional information is available at https://www.iacseaturtle.org.
International Convention for the Prevention of Pollution from Ships
(MARPOL): The MARPOL Convention is a combination of two treaties
adopted in 1973 and 1978 to prevent pollution of the marine environment
by ships from operational or accidental causes. The 1973 treaty covered
pollution by oil, chemicals, and harmful substances in packaged form,
sewage and garbage. The 1978 MARPOL Protocol was adopted at a
Conference on Tanker Safety and Pollution Prevention which included
standards for tanker design and operation. The 1978 Protocol
incorporated the 1973 Convention as it had not yet been in force and is
known as the International Convention for the Prevention of Marine
Pollution from Ships, 1973, as modified by the Protocol
[[Page 48419]]
of 1978 relating thereto (MARPOL 73/78). The 1978 Convention went into
force in 1983 (Annexes I and II). The Convention includes regulations
aimed at preventing and minimizing accidental and routine operations
pollution from ships. Amendments passed since have updated the
convention.
International Union for Conservation of Nature (IUCN): The IUCN
Species Programme assesses the conservation status of species on a
global scale. This assessment provides objective, scientific
information on the current status of threatened species. The IUCN Red
List of Threatened Species provides taxonomic, conservation status and
distribution information on plants and animals that have been globally
evaluated using the IUCN Red List Categories and Criteria. This system
is designed to determine the relative risk of extinction, and the main
purpose of the IUCN Red List is to catalogue and highlight those plants
and animals that are facing a higher risk of global extinction (i.e.,
those listed as Critically Endangered, Endangered and Vulnerable).
Additional information is available at https://www.iucnRed List.org/about.
Marine Turtle Conservation Act (MTCA): The MTCA is a key element of
sea turtle protection in the United States and internationally. This
Act authorizes a dedicated fund to support marine turtle conservation
projects in foreign countries, with emphasis on protecting nesting
populations and nesting habitat. Additional information is available at
https://www.fws.gov/international/wildlife-without-borders/marine-turtle-conservation-fund.html.
Memorandum of Agreement between the Government of the Republic of
the Philippines and the Government of Malaysia on the Establishment of
the Turtle Island Heritage Protected Area: Through a bilateral
agreement, the Governments of the Philippines and Malaysia established
The Turtle Island Heritage Protected Area (TIHPA), made up of nine
islands (6 in the Philippines and 3 in Malaysia). The following
priority activities were identified: management-oriented research, the
establishment of a centralized database and information network,
appropriate information awareness programs, a marine turtle resource
management and protection program, and an appropriate ecotourism
program (Bache and Frazier 2006).
Memorandum of Understanding of a Tri-National Partnership between
the Government of the Republic of Indonesia, the Independent State of
Papua New Guinea and the Government of Solomon Islands: This agreement
promotes the conservation and management of Western Pacific leatherback
turtles at nesting sites, feeding areas and migratory routes in
Indonesia, Papua New Guinea and Solomon Islands. This is done through
the systematic exchange of information and data on research, population
and migratory routes monitoring, nesting sites and feeding areas
management activities for Western Pacific leatherback turtles and by
enhancing public awareness of the importance of conserving these
turtles and their critical habitats. https://awsassets.wwf.or.id/downloads/mou_trinationalpartneshipagreement_clean.pdf.
Memorandum of Understanding Concerning Conservation Measures for
Marine Turtles of the Atlantic Coast of Africa (Abidjan Memorandum):
This MOU was concluded under the auspices of the Convention on the
Conservation of Migratory Species of Wild Animals (CMS) and became
effective in 1999. The MOU area covers 26 Range States along the
Atlantic coast of Africa extending approximately 14,000 km from Morocco
to South Africa. The goal of this MOU is to improve the conservation
status of marine turtles along the Atlantic Coast of Africa. It aims at
safeguarding six marine turtle species--including the leatherback
turtle--that are estimated to have rapidly declined in numbers during
recent years due to excessive exploitation (both direct and incidental)
and the degradation of essential habitats. This includes the protection
of the life stages from hatchlings through adults with particular
attention paid to the impacts of fishery bycatch and the need to
include local communities in the development and implementation of
conservation activities. However, despite this agreement, killing of
adult turtles and harvesting of eggs remains rampant in many areas
along the Atlantic African coast. Additional information is available
at https://www.cms.int/species/africa_turtle/AFRICAturtle_bkgd.htm.
National Sea Turtle Conservation Project in India: Launched in 1998
with the aim of protecting Lepidochelys olivacea, but it also has
conservation and protection strategies for all the other turtle species
nesting in the country. This project was undertaken by the Indian
government to oversee: Surveys, monitoring programs, fisheries
interactions, community and NGO participation, awareness raising and
education, research support, and other support for regional and
international co-operation and collaboration for sea turtles
conservation (Choudhury et al., 2001).
North American Agreement for Environmental Cooperation: As mandated
by the 1994 North American Agreement for Environmental Cooperation, the
Commission for Environmental Cooperation (CEC) encourages Canada, the
United States, and Mexico to adopt a continental approach to the
conservation of flora and fauna. In 2003, this mandate was strengthened
as the three North American countries launched the Strategic Plan for
North American Cooperation in the Conservation of Biodiversity. The
North American Conservation Action Plan (NACAP) initiative began as an
effort promoted by the three countries, through the CEC, to facilitate
the conservation of marine and terrestrial species of common concern.
In 2005, the CEC supported the development of a NACAP for Pacific
leatherbacks by Canada, the United States, and Mexico. Identified
actions in the plan addressed three main objectives: (1) protection and
management of nesting beaches and females; (2) mortality reduction from
bycatch throughout the Pacific Basin; and (3) waste management, control
of pollution, and disposal of debris at sea.
Ramsar Convention on Wetlands: The Convention on Wetlands, signed
in Ramsar, Iran, in 1971, is an intergovernmental treaty, which
provides the framework for national action and international
cooperation for the conservation and wise use of wetlands and their
resources. Currently, there are 158 parties to the convention, with
1,752 wetland sites, including important marine turtle habitat.
Additional information is available at https://www.ramsar.org.
Secretariat of the Pacific Regional Environment Programme (SPREP):
SPREP's turtle conservation program seeks to improve knowledge about
sea turtles in the Pacific through an active tagging program, as well
as maintaining a database to collate information about sea turtle tags
in the Pacific. SPREP supports capacity building throughout the central
and southwest Pacific. SPREP established an action plan for the Pacific
Islands (https://www.sprep.org/).
South-East Atlantic Fisheries Organization (SEAFO): SEAFO manages
fisheries activities in the Southeast Atlantic high seas area,
excluding tunas and billfish. SEAFO adopted Resolution 01/06, ``to
Reduce Sea Turtle Mortality in Fishing Operations,'' in 2006. The
Resolution requires Members to: (1) Implement the FAO Guidelines; and
(2) establish on-board observer programs to collect information on sea
turtle
[[Page 48420]]
interactions in SEAFO-managed fisheries. This Resolution is not legally
binding. Additional information is available at https://www.seafo.org.
South Atlantic Association: In the southwest Atlantic, the South
Atlantic Association is a multinational group that includes
representatives from Brazil, Uruguay, and Argentina and meets bi-
annually to share information and develop regional action plans to
address threats including bycatch (https://www.tortugasaso.org/). At the
national level, Brazil has developed a national plan for sea turtle
bycatch reduction that was initiated in 2001 (Marcovaldi et al. 2002).
This national plan includes various activities to mitigate bycatch,
including time-area restrictions of fisheries, use of bycatch reduction
devices, and working with fishermen to successfully release live-
captured turtles. In Uruguay, all sea turtles are protected from human
impacts, including fisheries bycatch, by presidential decree (Decreto
Presidencial 144/98).
United Nations Convention on the Law of the Sea (UNCLOS): To date,
155 countries, including most mainland countries lining the western
Pacific, and the European Community have joined in the convention. The
United States has signed the treaty and abides by some provisions, but
the Senate has not ratified it. Aside from its provisions defining
ocean boundaries, the convention establishes general obligations for
safeguarding the marine environment through mandating sustainable
fishing practices and protecting freedom of scientific research on the
high seas. Additional information is available at https://www.un.org/Depts/los/index.htm.
United Nations' Food and Agricultural Organization (FAO): The FAO
published guidelines for sea turtle protection, entitled Technical
Consultation on Sea Turtle-Fishery Interactions (FAO 2005). The UN 1995
Code of Conduct for Responsible Fisheries (FAO 2004) provides
guidelines for the development and implementation of national fisheries
policies, including gear modification (e.g., circle hooks, fish bait,
deeper sets, and reduced soak time), new technologies, and management
of areas where fishery and sea turtle interactions are more severe. The
guidelines stress the need for mitigation measures, data on all
fisheries, fishing industry involvement, and education for fishers,
observers, managers, and compliance officers (FAO 2004).
United Nations Resolution 44/225 on Large-Scale Pelagic Driftnet
Fishing: In 1989, the UN called, in a unanimous resolution, for the
elimination of all high seas driftnets by 1992. Additional information
is available at https://www.un.org/documents/ga/res/44/a44r225.htm.
Although numerous conservation efforts apply to the species, they
do not adequately reduce its risk of extinction for the reasons
discussed previously.
Extinction Risk Analysis
The best available information is consistent with the species'
current ``endangered'' listing. The species exhibits a global total
index of nesting female abundance of 32,060 females at monitored
beaches. This number is lower than historical estimates of nesting
female abundance (n = 115,000, Pritchard 1982; and n = 34,500, Spotila
1996), which did not include the large, but then unknown, Gabon nesting
aggregation. Limited nesting female abundance is a major source of
concern for most DPSs, whose small population sizes place them in
danger of stochastic or catastrophic events that increase extinction
risk. The limited nesting female abundance increases the extinction
risk of the species.
The species also exhibits declining nesting trends for all but one
of the DPSs. With the exception of the DPS with the smallest index of
nesting female abundance (i.e., SW Atlantic DPS, with 27 nesting
females), the DPSs are declining at rates of 0.3 to 9.3 percent
annually. Even low levels of decline are a threat for DPSs with limited
nesting female abundance, and nesting declines of approximately nine
percent (i.e., NW and SE Atlantic DPSs) are unsustainable. Total
declines of 97 and 99 percent have occurred within the East Pacific and
NE Indian DPSs, respectively, since nesting was first identified and
quantified for these populations. The declining trends in nesting
increase the extinction risk of the species.
The species exhibits broad nesting and foraging ranges. However,
metapopulation dynamics have likely been reduced, with reductions in
abundance and the loss of some nesting aggregations. The species also
demonstrates little genetic diversity, relative to other sea turtle
species. Although the species demonstrates some resilience to threats,
overall we find it to be at risk of extinction, due to limited
abundance and declining nesting trends, which reflect the cumulative
impacts of threats that have acted on the species in the past (and in
many cases continue to act on the species).
Current threats continue to place the species in danger of
extinction. The primary threat to the species is bycatch in commercial
and artisanal, pelagic and coastal, fisheries. Fisheries bycatch
reduces abundance by removing individuals from the population. Because
several fisheries operate near nesting beaches, productivity is also
reduced when nesting females are prevented from returning to nesting
beaches. The harvest of eggs and turtles is also a major threat to the
species. Illegal poaching occurs throughout the range of the species,
and harvest is legal but poorly documented in some nations. The loss
and modification of nesting habitat is another major threat, reducing
productivity and, in some instances, abundance, when nesting females
die as a result of artificial lighting or obstructions preventing them
from returning to sea. Predation results in the loss of eggs and
hatchlings, reducing productivity of the species. Additional threats
that occur throughout the range of the species include vessel strikes,
pollution, marine debris, oil and gas exploration, and climate change.
Natural disasters and oceanographic regime shifts are threats in some
areas. Though many regulatory mechanisms are in place, they do not
adequately reduce the impact of these threats.
Based on our review of the best available scientific and commercial
data, we find nothing that is inconsistent with the leatherback
species' current listing as an endangered species. In sum, the best
available information is consistent with the current listing status of
the leatherback sea turtle as an endangered species throughout its
range. The threatened species definition does not apply because the
species is currently in danger of extinction (i.e., at present), rather
than on a trajectory to become so within the foreseeable future.
Final Determination
The Services determined that the best available scientific and
commercial information would support recognizing seven populations as
DPSs (including the NW Atlantic) because they meet the discreteness and
significance criteria for DPSs. However, we found that--even were they
to be recognized and listed separately--all DPSs meet the definition of
an endangered species because they are in danger of extinction
throughout all of their ranges. The leatherback turtle is currently
listed throughout its range as an endangered species. Replacing this
listing with seven endangered DPSs would not be consistent with
Congressional guidance to use the authority to list DPSs ``sparingly''
while encouraging the conservation of genetic diversity (see Senate
Report 151, 96th
[[Page 48421]]
Congress, 1st Session). Such guidance clearly indicates that the
Services have some discretion to determine whether or not to recognize
DPSs that would require disaggregating an existing listing even where
those populations can be shown to meet the discreteness and
significance tests of the DPS Policy.
After determining that all seven populations would have the same
status as the overall species, we next considered whether there was any
reason to nevertheless replace the global (range-wide) listing with
individual listings for the seven DPSs. We conclude that disaggregating
the global listing is not warranted. It would be inconsistent with
Congressional guidance and run counter to the conservation purposes of
the Act to disaggregate the current listing into DPSs, because those
DPSs would have the same listing status as the whole currently.
Disaggregating this listing would bring about significant complications
and possible public confusion without any meaningful corresponding
conservation benefit. Replacing the range-wide listing with seven DPSs
having the same status would not provide leatherback turtles with an
overriding conservation benefit, as all members are currently protected
to the fullest extent under the ESA as an endangered species. Section 7
consultations already consider the effects of an action on individuals
and populations to determine whether a Federal agency has insured that
its action is not likely to jeopardize the continued existence of the
species. Even if the species were disaggregated into DPSs, this change
would not be expected to result in different substantive outcomes in
consultations. In addition, focused conservation efforts have been, and
will continue to be, applied at scales smaller than the species-level.
For example, FWS' Marine Turtle Conservation Fund provides funding to
partners in foreign nations to protect leatherback turtles and their
nesting habitats; projects include efforts to monitor and protect
leatherback turtles in Indonesia and Gabon (https://www.fws.gov/international/wildlife-without-borders/marine-turtle-conservation-fund.html). Similarly, Pacific leatherback turtles are highlighted
under NMFS' Species in the Spotlight: Survive to Thrive initiative,
which directs attention and resources to highly-at-risk species
(https://www.fisheries.noaa.gov/topic/endangered-species-conservation#species-in-the-spotlight).
For these reasons, the Services have determined that replacing the
existing global listing with separate listings for individual DPSs is
not warranted. Although the best available data indicates that the
populations meet the criteria for significance and discreteness, we
find that it would not further the purposes of the Act to recognize and
list seven DPSs separately as endangered under the ESA. The current
global listing of the species remains in effect.
We conclude that the petitioned actions, to identify the NW
Atlantic population as a DPS and list it as a threatened species under
the ESA, are not warranted. This is a final action, and, therefore, we
are not soliciting public comments.
Peer Review
In December 2004, the Office of Management and Budget (OMB) issued
a Final Information Quality Bulletin for Peer Review, establishing
minimum peer review standards, a transparent process for public
disclosure of peer review planning, and opportunities for public
participation. The OMB Bulletin, implemented under the Information
Quality Act (Pub. L. 106-554), is intended to enhance the quality and
credibility of the Federal government's scientific information and
applies to influential or highly influential scientific information
disseminated on or after June 16, 2005. To satisfy our requirements
under the OMB Bulletin, we obtained independent peer review of the
Status Review Report by independent scientists with expertise in
leatherback turtle biology, endangered species listing policy, and
related fields. All peer reviewer comments were addressed prior to the
publication of the Status Review Report and this finding.
References Cited
A complete list of references is available upon request to the NMFS
Office of Protected Resources (see ADDRESSES).
Authority
The authority for this action is the Endangered Species Act of
1973, as amended (16 U.S.C. 1531 et seq.).
Samuel D. Rauch III,
Deputy Assistant Administrator for Regulatory Programs, National Marine
Fisheries Service.
Aurelia Skipwith,
Director, U.S. Fish and Wildlife Service.
[FR Doc. 2020-16277 Filed 8-7-20; 8:45 am]
BILLING CODE 3510-22-P