Endangered and Threatened Species; Determination on the Designation of Critical Habitat for Giant Manta Ray, 66652-66664 [2019-26265]
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Federal Register / Vol. 84, No. 234 / Thursday, December 5, 2019 / Notices
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Dated: December 2, 2019.
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[FR Doc. 2019–26272 Filed 12–4–19; 8:45 am]
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DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
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[Docket No. 191127–0095; RTID 0648–
XR030]
Endangered and Threatened Species;
Determination on the Designation of
Critical Habitat for Giant Manta Ray
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
AGENCY:
Notice of critical habitat
determination.
ACTION:
We, NMFS, have determined
that a designation of critical habitat is
not prudent at this time. Based on a
comprehensive review of the best
scientific data available, we find that
there are no identifiable physical or
biological features that are essential to
the conservation of the giant manta ray
within areas under U.S. jurisdiction. We
also find that there are no areas outside
of the geographical area occupied by the
species under U.S. jurisdiction that are
essential to its conservation. As such,
we find that there are no areas within
the jurisdiction of the United States that
meet the definition of critical habitat for
the giant manta ray.
SUMMARY:
This finding is made on
December 5, 2019.
DATES:
Electronic copies of the
determination, list of references, and
supporting documents prepared for this
action are available from the NMFS
Office of Protected Resources website at
https://www.fisheries.noaa.gov/species/
giant-manta-ray.
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ADDRESSES:
FOR FURTHER INFORMATION CONTACT:
Maggie Miller, NMFS, Office of
Protected Resources, (301) 427–8403.
SUPPLEMENTARY INFORMATION:
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Background
On January 22, 2018, we published a
final rule to list the giant manta ray
(Manta birostris) as a threatened species
under the Endangered Species Act
(ESA) (83 FR 2916). Section 4(b)(6)(C) of
the ESA requires the Secretary of
Commerce (Secretary) to designate
critical habitat concurrently with
making a determination to list a species
as threatened or endangered unless it is
not determinable at that time, in which
case the Secretary may extend the
deadline for this designation by 1 year.
At the time of listing, we concluded that
critical habitat was not determinable
because sufficient information was not
available to: (1) Identify the physical
and biological features essential to the
conservation of the species at an
appropriate level of specificity,
particularly given the uncertainty
regarding habitats required to support
its life history (e.g., pupping and
nursery grounds were unknown) and
migratory movements, (2) determine the
specific geographical areas that contain
the physical and biological features
essential to conservation of the species,
particularly given the global range of the
species, and (3) assess the impacts of the
designation. We requested relevant
information from interested persons to
help us identify and describe the
physical and biological features
essential to the conservation of the giant
manta ray, and assess the economic
consequences of designating critical
habitat for the species. We solicited
input from the public, other concerned
government agencies, the scientific
community, industry and any other
interested party on features and areas
that may meet the definition of critical
habitat for the giant manta ray within
U.S. waters. We received information
regarding giant manta ray occurrence in
the Flower Garden Banks National
Marine Sanctuary (Stewart et al. 2018b)
as well as off the coast of Florida. We
reviewed this information and
considered it along with other available
information we compiled. Together, this
information comprises the best available
scientific data for use in the
identification of critical habitat for the
giant manta ray. However, as discussed
below, based on these data we find that
there are no identifiable physical or
biological features that are essential to
the conservation of the giant manta ray
within areas under U.S. jurisdiction, or
unoccupied areas under U.S.
jurisdiction that are essential to the
conservation of the species. Therefore,
at this time we find no areas within U.S.
jurisdiction that meet the definition of
critical habitat for the giant manta ray.
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This finding describes information on
the biology, distribution, and habitat use
of the giant manta ray and the methods
used to identify areas that may meet the
definition of critical habitat. In this
determination, we focus on information
directly relevant to the designation of
critical habitat for giant manta rays.
Giant Manta Ray Biology and Status
The following discussion of the life
history and status of giant manta ray is
based on the best scientific data
available, including the ‘‘Endangered
Species Act Status Review Report: Giant
Manta Ray (Manta birostris) and Reef
Manta Ray (Manta alfredi)’’ (Miller and
Klimovich 2017).
Manta rays are large bodied,
planktivorous rays, considered part of
the Mobulidae subfamily. Manta species
are distinguished from other Mobula
rays in that they tend to be larger, with
a terminal mouth, and have long
cephalic fins (Evgeny 2010); however,
misidentifications are common both
between Manta species (i.e., between M.
alfredi and M. birostris) as well as
between Manta and Mobula rays. In
addition, recent taxonomic studies have
suggested that Manta birostris and
Manta alfredi may actually be closely
related to the giant devil ray (Mobula
mobular) (White et al. 2017), with
genetic analyses that demonstrate
support for nesting these species under
the genus Mobula rather than Manta
(White et al. 2017; Hosegood et al.
2019). The studies still recognize both
manta rays as distinct species, but refer
to them as Mobula birostris and Mobula
alfredi.
The giant manta ray, M. birostris, can
be found in all ocean basins, while the
reef manta ray, M. alfredi, is currently
only observed in the Indian Ocean and
the western and south Pacific.
Additionally, we note that a third,
putative manta ray species has been
identified (referred to here as M. cf.
birostris), with its range extending along
the Atlantic coast, Gulf of Mexico, and
Caribbean, based on research conducted
in the western Atlantic (A. Marshall,
MMF, pers. comm. to M. Miller, NMFS
OPR, 2019). A manuscript identifying
this third species is expected in the near
future; however, according to Dr.
Andrea Marshall, this newly identified
manta species is highly abundant off the
U.S. east coast, with a large population
also found off the Yucata´n peninsula (A.
Marshall, MMF, pers. comm. to M.
Miller, NMFS OPR, 2019). This new
species looks very similar to M.
birostris, with only a few diagnostic
features that could potentially
distinguish the two (mainly small
morphological and meristic ones; A.
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Marshall, MMF, pers. comm. to M.
Miller, NMFS OPR, 2019). Without
genetic testing, species identification
cannot be completely validated
(Hinojosa-Alvarez et al. 2016; Kashiwagi
et al. 2017; Hosegood et al. 2019).
Therefore, for purposes of this critical
habitat determination, we will consider
any records of manta rays in the
Atlantic to be M. birostris (even though
an unknown proportion may comprise
M. cf. birostris) and will continue to
recognize Manta birostris as a species
under the genus Manta.
The genus Manta has a complex
taxonomic history due partially to the
difficulty of preserving such large
specimens and also the conflicting
historical reports of taxonomic
characteristics (Couturier et al. 2012;
Kitchen-Wheeler 2013). Prior to 2009,
most manta rays were categorized as
Manta birostris, but Marshall et al.
(2009) presented new data that
supported the splitting of the Manta
genus into two species: M. birostris and
M. alfredi.
Both Manta species have diamondshaped bodies with wing-like pectoral
fins; the distance over this wingspan is
termed disc width (DW). There are two
distinct color types in both species:
Chevron and black (melanistic). Most of
the chevron variants have a black dorsal
surface and a white ventral surface with
distinct patterns on the underside that
can be used to identify individuals
(Marshall et al. 2008; Kitchen-Wheeler
2010; Deakos et al. 2011). The black
color variants of both species are
entirely black on the dorsal side and
almost completely black on the ventral
side, except for areas between the gillslits and the abdominal area below the
gill-slits (Kitchen-Wheeler 2013).
Giant manta rays inhabit tropical,
subtropical, and temperate bodies of
water and are commonly found offshore,
in oceanic waters, and near productive
coastlines. It is thought to be a generally
long-lived species (>28 years) (Stewart
et al. 2018a) with low reproductive
output. Manta rays, like all
chondrichthyans, reproduce via internal
fertilization (Wourms 1981), and the
sexes can be differentiated by the
presence of myxopterigia, or claspers,
on the inner margin of the pelvic fins in
males, whereas females lack these
structures. Sexual maturity in males can
be easily determined by examining the
level of calcification in these
intromittent organs. In their
examination of mobulids taken as
bycatch in the Indonesian drift net
fishery, White et al. (2006) found that
male M. birostris greater than 3,800 mm
DW possessed fully calcified claspers
and were, therefore, mature, while those
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less than 3,800 mm DW possessed either
non-calcified or partially calcified
claspers. In the same study, White et al.
(2006) found that females 2,732 to 3,774
mm DW were immature and females
measuring 4,126 mm DW and greater
were mature. White and Last (2016)
report similar ranges, with males
maturing between approximately 3,750
and 4,000 mm DW and females
maturing between approximately 4,100
and 4,700 mm DW. In the Flower
Gardens Banks National Marine
Sanctuary (FGBNMS), Stewart et al.
(2018b) observed a mature male M.
birostris with an estimated size of 3,600
mm. The age that M. birostris matures is
not known, but it may be similar to that
of reef mantas, with males maturing at
3–6 years and females at 8–10 years
(Stewart et al. 2018a).
Gestation time is also not known for
this species, and parturition has only
been witnessed once and under
unnatural conditions (Coles 1916). It is
suspected that gestation would be
similar to that observed in M. alfredi,
which is generally accepted to be 12 to
13 months (Kitchen-Wheeler 2013). In
addition to the Coles (1916) observation
of a single embryo aborted during
capture, the limited investigations of
pregnant females with embryos intact
have all indicated the presence of a
single embryo per pregnancy (Muller
and Henle 1838–1841; Beebe and TeeVan 1941). Similarly, reports of reef
manta ray births and dissections have
also all revealed only a single embryo
(Homma et al. 1999; Uchida et al. 2008).
Size at birth has remained elusive for M.
birostris. The embryos examined in the
previous studies had sizes of 1,140 mm
and 1,270 mm DW (Muller and Henle
1838–1841; Beebe and Tee-Van 1941),
while the smallest free swimming
individuals reported by Stewart et al.
(2018b) were approximately 1,000 mm
DW (however, these individuals may
have been M. cf. birostris).
Rambahiniarison et al. (2018) recently
estimated size at birth of M. birostris to
be 2,000 mm DW based on the DW of
the largest fetus and the smallest freeliving specimen captured in the
Philippines mobulid fishery.
Very little is known about the early
life stages or habitat needs or
requirements of M. birostris because,
until fairly recently, juveniles have
rarely been observed in the wild.
However, large numbers of juvenile M.
birostris have been caught in Sri Lanka
in offshore pelagic habitats by the gillnet fisheries, landed by fisherman in
Brazil and Indonesia, and also observed
in oceanic habitats off Mexico (Stewart
et al. 2016a; Stewart et al. 2018b).
Stewart et al. (2016a) suggests that adult
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and juvenile giant mantas may use
similar offshore pelagic habitats, but
that the juveniles may avoid cleaning
stations and other near-shore areas
where adults are more commonly
observed to reduce predation risk. In
fact, results from stable isotope analyses
of muscle tissues collected from both
adult and juvenile M. birostris off Peru,
Sri Lanka, and the Philippines appear to
provide further confirmation that the
species may not undergo an ontogenetic
shift in feeding behavior or trophic
level, with both adults and juveniles
sharing the same habitats and targeting
the same prey (Stewart et al. 2017).
In terms of prey, giant manta rays
primarily feed on planktonic organisms
such as euphausiids, copepods, mysids,
decapod larvae, and shrimp, with some
studies noting their consumption of
small and moderate sized fishes as well
(Bigelow and Schroeder 1953; Carpenter
and Niem 2001; Graham et al. 2012;
Stewart et al. 2016b; Burgess 2017;
Rohner et al. 2017). They feed by
swimming with their mouths open,
continuously filtering zooplankton.
Their gill rakers filter out water, leaving
behind food particles that are then
directed to the esophagus through crossflow (Paig-Tran 2012). This filter
mechanism allows mantas to retain prey
of various sizes, even if they are smaller
than the filter pores, which means they
can effectively feed on mixed
zooplankton assemblages where prey
range in size from small calanoid
copepods to larger mysids and
euphausiids (Stewart et al. 2016b).
Given the feeding habits of the giant
manta ray, it can be considered a
generalist carnivore, with a trophic
position of approximately 3.4 (Burgess
et al. 2016; Burgess 2017).
With regards to movement, the giant
manta ray is considered to be a
migratory species, with satellite tracking
studies measuring straight line distances
of up to 1,500 km (Hearn et al. 2014).
Some giant manta rays appear to migrate
seasonally, possibly due to the seasonal
fluctuations in food sources (Wilson et
al. 2001; Luiz et al. 2009; Graham et al.
2012; Sobral and Afonso 2014; De Boer
et al. 2015; Girondot et al. 2015; Stewart
et al. 2016a; Hacohen-Domene´ et al.
2017). However, in some portions of its
range, the species may actually exist as
well-structured subpopulations with a
high degree of residency (Stewart et al.
2016a).
As discussed in the proposed rule (82
FR 3694, January 12, 2017) and final
rule (83 FR 2916, January 22, 2018) to
list the giant manta ray, the most
significant threat to the species is
overutilization for commercial
purposes. Giant manta rays are both
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targeted and caught as bycatch in a
number of fisheries throughout their
range, and are most susceptible to
industrial purse-seine and artisanal
gillnet fisheries. With the expansion of
the international mobulid gill raker
market and increasing demand for
manta ray products, estimated take of
giant manta rays, particularly in many
portions of the Indo-Pacific, frequently
exceeds numbers of identified
individuals in those areas. Observations
from these areas also indicate declines
in sightings and landings of the species.
Given the extremely low reproductive
output and overall productivity of the
giant manta ray, it is inherently
vulnerable to threats that would deplete
its abundance, with a low likelihood of
recovery. So, while there is considerable
uncertainty regarding the current
abundance of M. birostris throughout its
entire range, the best available
information indicates that the species is
likely to become an endangered species
within the foreseeable future throughout
a significant portion of its range (the
Indo-Pacific and eastern Pacific portion)
due to overutilization.
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Critical Habitat Identification and
Designation
Critical habitat is defined by section
3 of the ESA as: ‘‘(i) the specific areas
within the geographical area occupied
by the species, at the time it is listed
. . . , on which are found those physical
or biological features (I) essential to the
conservation of the species and (II)
which may require special management
considerations or protection; and (ii)
specific areas outside the geographical
area occupied by the species at the time
it is listed . . . upon a determination by
the Secretary that such areas are
essential for the conservation of the
species.’’ This definition provides a
step-wise approach to identifying areas
that may qualify as critical habitat for
the giant manta ray: (1) Determine the
geographical area occupied by the
species at the time of listing; (2) identify
physical or biological habitat features
essential to the conservation of the
species; (3) delineate specific areas
within the geographical area occupied
by the species on which are found the
physical or biological features; (4)
determine whether the features in a
specific area may require special
management considerations or
protection; and (5) determine whether
any unoccupied areas are essential for
conservation. Our evaluation and
conclusions as we worked through this
step-wise process are described in detail
in the following sections.
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Geographical Area Occupied by the
Species
The ‘‘geographical area occupied by
the species’’ is defined in our
regulations as ‘‘an area that may
generally be delineated around species’
occurrences, as determined by the
Secretary (i.e., range). Such areas may
include those areas used throughout all
or part of the species’ life cycle, even if
not used on a regular basis (e.g.,
migratory corridors, seasonal habitats,
and habitats used periodically, but not
solely by vagrant individuals).’’ (50 CFR
424.02). Further, our regulations at 50
CFR 424.12(g) state: ‘‘The Secretary will
not designate critical habitat within
foreign countries or in other areas
outside of the jurisdiction of the United
States.’’ As such, we focus the following
discussion on the range of the species
within waters under U.S. jurisdiction.
In the Atlantic, giant manta rays have
been confirmed as far north as Long
Island, New York (offshore around the
Hudson Canyon region) (Normandeau
Associates and APEM Ltd 2017);
however, as will be discussed later, we
note that they are generally rare north of
Cape Hatteras, North Carolina. To the
south, giant manta rays occur off the
coast of North Carolina, South Carolina,
Georgia, and Florida (Marshall et al.
2011). Giant manta rays can also be
found throughout the U.S. Gulf of
Mexico and within the U.S. Caribbean,
including off Puerto Rico and the U.S.
Virgin Islands (Marshall et al. 2011). In
the central Pacific, giant manta rays are
found off Hawaii (Clark 2010) and Jarvis
Island (K. Lino unpublished data).
While there have been no confirmed
sightings of giant manta rays in waters
of the other Pacific Remote Island Areas,
Northern Mariana Islands (Kashiwagi et
al. 2011), Guam (Kashiwagi et al. 2011),
or American Samoa, based on confirmed
observations of the species elsewhere
throughout the Pacific (e.g., Ogasawara
Islands, Japan (Kashiwagi et al. 2010);
Philippines (Verdote and Ponzo 2014);
French Polynesia (Mourier 2012); Jarvis
Island (K. Lino unpublished data);
Hawaii (Clark 2010)) and coupled with
the migratory and pelagic nature of giant
manta rays, their ability to exploit
significant depths, and tolerance of
tropical to temperate water
temperatures, we find no known
barriers to their movement that may
prevent them from occurring at these
locations.
In the eastern U.S. Pacific, while there
is documentation of a giant manta off
the west coast (i.e., San Clemente Island,
California), this sighting was of a single
individual in 2014 (Warneke 2014) and
there have been no documented
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sightings since (or prior to) this time.
Given the amount of fishing effort, as
well as the human population density in
these regions, it is highly unlikely that
substantial concentrations of giant
manta rays would have passed
unnoticed. As such, we consider this
individual to be a vagrant of the species
(an individual that occurs outside of the
species’ normal range). Therefore, as the
occurrence of giant manta rays in waters
off the U.S. west coast is extremely
uncommon, we do not consider this
geographical area to be part of the
species’ occupied range at the time of
listing.
Conclusion
Based on the above information and
analysis, we define the geographical
area occupied by the giant manta ray at
the time of listing as all U.S. waters off
the east coast, from Florida to Long
Island, New York, the entire Gulf coast,
the U.S. Virgin Islands and Puerto Rico
in the Caribbean, and Hawaii, the
Pacific Remote Islands Areas, Guam,
American Samoa, and the Northern
Mariana Islands in the Pacific.
Physical or Biological Features
Essential for Conservation
Within the geographical area
occupied by an endangered or
threatened species at the time of listing,
critical habitat consists of specific areas
upon which are found those physical or
biological features essential to the
conservation of the species and that may
require special management
considerations or protection. The ESA
does not specifically define physical or
biological features; however, court
decisions and joint NMFS-USFWS
regulations at 50 CFR 424.02 provide
guidance on how physical or biological
features are expressed. Specifically,
these regulations state that the physical
and biological features are those that are
essential to support the life-history
needs of the species, including but not
limited to, water characteristics, soil
type, geological features, sites, prey,
vegetation, symbiotic species, or other
features. A feature may be a single
habitat characteristic, or a more
complex combination of habitat
characteristics. Features may include
habitat characteristics that support
ephemeral or dynamic habitat
conditions. Features may also be
expressed in terms relating to principles
of conservation biology, such as patch
size, distribution distances, and
connectivity. (50 CFR 424.02).
Section 3 of the ESA (16 U.S.C.
1532(3)) defines the terms ‘‘conserve,’’
‘‘conserving,’’ and ‘‘conservation’’ to
mean: ‘‘to use and the use of all
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methods and procedures which are
necessary to bring any endangered
species or threatened species to the
point at which the measures provided
pursuant to this chapter are no longer
necessary.’’ For giant manta rays, we
consider conservation to include the use
of all methods and procedures necessary
to bring giant manta rays to the point at
which factors related to population
ecology and vital rates indicate that the
species is recovered in accordance with
the definition of recovery in 50 CFR
402.02. Important factors related to
population ecology and vital rates
include population size and trends,
range, distribution, age structure, gender
ratios, age-specific survival, age-specific
reproduction, and lifetime reproductive
success. Based on the available
knowledge of giant manta ray
population ecology and life history, we
have identified four biological behaviors
that are critical to the goal of increasing
survival and population growth: (1)
Foraging, (2) pupping, (3) migration,
and (4) breeding. In the following
section, we evaluate whether there are
physical and biological features of the
habitat areas known or thought to be
used for these behaviors that are
essential to the species’ conservation
because they facilitate or are intimately
tied to these behaviors and, hence,
support the life-history needs of the
species. Because these behaviors are
essential to the species’ conservation,
facilitating or protecting each one is
considered a key conservation objective
for any critical habitat designation for
this species.
Analysis of the Physical and Biological
Features of Foraging Habitat That Are
Essential to the Conservation of the
Species
Giant manta rays are filter-feeders and
generalist carnivores that feed on a
variety of planktonic organisms,
including euphausiids, copepods,
mysids, decapod larvae and shrimp, as
well as small fishes. Prey needs to be of
sufficient density and quality to support
the energy requirements for the giant
manta rays, particularly as they conduct
long-distance migrations across open
oceans. Sustained decreases in prey
quantity, quality, availability, or
accessibility can decrease foraging
success of giant manta rays and
eventually lead to reduced individual
growth, reproduction, and development.
Therefore, using the best available data,
we examined the diet and energy needs
of giant manta rays, including foraging
behavior, to determine whether we
could identify physical or biological
features of habitat that facilitate
successful giant manta ray feeding and,
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thus, are essential for the conservation
of the species.
As mentioned above, planktonic
organisms comprise the majority of the
diet for giant manta rays. While it was
previously assumed that manta rays
obtain most of their energy needs from
surface zooplankton, results from recent
studies indicate that these feeding
events may not be the primary source of
the dietary intake (Burgess et al. 2016;
Stewart et al. 2016b). For example, for
giant manta rays off Ecuador, Burgess et
al. (2016) estimated that, on average,
mesopelagic food sources contribute 73
percent to the giant manta ray’s diet
compared to 27 percent for surface
zooplankton. In the Mexican Pacific,
Stewart et al. (2016b) interpreted dive
profiles and submersible video data of
M. birostris to suggest that giant manta
rays frequently forage on vertically
migrating zooplankton and zooplankton
in the epipelagic scattering layers in
addition to surface zooplankton.
Analysis of stomach contents and
collection of zooplankton during
observed giant manta ray feeding events
reveal a varied diet, with no targeting of
a specific species or size of prey
(Graham et al. 2012; Armstrong et al.
2016; Stewart et al. 2016b; Burgess
2017; Rohner et al. 2017). Rather,
density of the prey appears to be the
driving factor that triggers giant manta
ray feeding behavior. However, the
levels necessary to attract giant manta
rays remain unknown. For example, a
study conducted by Burgess (2017)
found that giant manta ray aggregations
off the northwest side of Isla de la Plata,
Ecuador, were unlikely associated with
foraging opportunities as observations of
feeding events were rare. Specifically,
Burgess (2017) collected surface
zooplankton during feeding events (n=5)
and during non-feeding events (n=79)
and calculated that the dry zooplankton
biomass was 1.9 mg m¥3 during the rare
M. birostris feeding events and 1.4 mg
m¥3 during non-feeding events.
Although comparable data are
unavailable for M. birostris elsewhere
throughout its range, these figures are
substantially lower than what has been
reported for the closely related reef
manta ray, M. alfredi, in eastern
Australia during regular active feeding
(19.1 mg m¥3) and non-feeding (9.3 mg
m¥3) events (Armstrong et al. 2016). In
fact, Armstrong et al. (2016) determined
that the critical prey density threshold
for M. alfredi feeding was 11.2 mg
m 3. If M. birostris has similar prey
density thresholds, these data lend
support to Burgess (2017)’s finding that
the aggregative behavior of giant manta
rays at Isla de la Plata is unlikely related
to feeding. Furthermore, the data
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suggest that for habitat to be
characterized as providing necessary
foraging opportunities, it likely requires
substantially higher levels of
zooplankton biomass than what was
found off Isla de la Plata.
In terms of energy needs, the only
available data that provides insight for
M. birostris is from a study that
examined the stomach contents of giant
manta rays collected within the Bohol
Sea (Philippines) in 2015 (Rohner et al.
2017). Using adiabatic bomb
calorimetry, Rohner et al. (2017)
calculated that krill (Euphausia
diomedeae), the dominant prey species
for M. birostris in this particular area,
contributed 24,572 kJ (±20,451 kJ s.d.)
per 100 g of stomach content in M.
birostris. When scaled up based on the
total number of euphausiids per
stomach, the authors estimated that E.
diomedeae contributed up to 631,167
kcal in the giant manta ray diet (Rohner
et al. 2017). This energetic contribution
is significantly greater than what has
been found for reef manta rays in
captivity. Rohner et al. (2017), citing a
personal communication, reports that in
aquaria, a 350 cm DW M. alfredi is fed
3,500 kcal per day and a 450 cm DW M.
alfredi is fed 6,100 kcal per day, with
captive reef manta rays consuming 12.7
percent of their body weight in
euphausiids weekly (Homma et al.
1999). Although energy requirements
and caloric intake for captive manta rays
will likely be different than those found
in the wild, Rohner et al. (2017)
proposes that the significant calorific
value of the M. birostris stomach
contents suggests that giant manta rays
partake in numerous feeding events over
several days or, alternatively, engage in
a few, sporadic, opportunistic feeding
events on large aggregations of prey that
can be used to sustain them until their
next meal. Burgess (2017) tends to agree
with the latter. The author cites the
particularly large capacity of the M.
birostris stomach, as well as the
branchial filter pad and filtration
mechanism used by manta rays (which
allows for the capture of numerous
macroscopic zooplankton and small
fishes of varying sizes) to support the
assumption that manta rays likely
exploit large patches of zooplankton for
a high net energy gain in a short period
of time (Burgess 2017). However, with
only one study that has examined the
energy contents of a particular prey item
of M. birostris in a specific area, it is
difficult to make any conclusions as to
the general energy needs or
requirements for the species throughout
its range.
With the lack of available data
regarding prey density thresholds or
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caloric value requirements, we next
looked at areas where manta rays have
been observed or assumed to be feeding
to determine whether we could identify
any physical or biological features of
these habitats that are tied to foraging
behavior. In many portions of the
species’ range, it is the presence of
seasonal upwelling events, which
concentrate plankton and create patches
of high productivity, that appear to
drive the occurrence of giant manta rays
in areas, presumably for foraging. For
example, off the northern Yucata´n
peninsula, Hacohen-Domene´ et al.
(2017) found a higher probability of M.
birostris occurrence from July through
September, with the main difference
being the increase in primary
productivity during this time of year
(with particularly high probability of
occurrence when primary productivity
was at 4,500 mg C·m¥2·day¥1). Other
features associated with a greater
probability of giant manta ray presence
in this area included sea surface
temperatures (SST) warmer than 27 °C,
shallow (<10 m depths) and nearshore
waters (<50 km from shore), with a
bottom slope of <0.5° (Hacohen-Domene´
et al. 2017). However, the authors note
that most of the manta rays observed in
the study were not foraging but rather
swimming alone or in pairs. While
Hacohen-Domene´ et al. (2017) did not
observe or analyze feeding habits in
their study, Hinojosa-Alvarez et al.
(2016) confirmed foraging behavior in
this area (specifically between
21°46.020′ N and 87°01.200′ W and
21°30.00′ and 86°4100), with videos of
Yucata´n manta rays feeding in surface
waters from May through August (the
same period as the seasonal upwelling).
Seasonal occurrence of manta rays
was also observed off the continental
shelf of French Guiana. Specifically,
Girondot et al. (2015) observed a peak
in the presence of manta rays between
July and December in the river-ocean
transition zone off French Guiana.
While specific features of the habitat
where giant manta rays were observed
was not provided, the authors did note
that phytoplankton biomass and
primary productivity is generally
highest during the months of manta ray
presence, with a biomass of over 25 mg
Chl-a m¥3 and productivity of over 8 g
C°m¥2*day¥1 (Girondot et al. 2015).
Similarly, in southeastern Brazil,
giant manta rays are most frequently
sighted in Laje de Santos Marine State
Park (24° S) during seasonal upwelling,
from June to August (Luiz et al. 2009).
During this time, the warm Brazil
Current weakens and coastal waters
change direction and move northward,
bringing waters from the southern
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Falklands Current to areas of
southeastern Brazil (Luiz et al. 2009).
This current displaces a low salinity
front (generated by discharge from the
La Plata River) from the mouth of the La
Plata River during the summer to areas
north in the winter (Luiz et al. 2009). It
is thought that this coastal front, which
accumulates plankton, may attract giant
manta rays at Laje de Santos Marine
State Park in the winter months (Luiz et
al. 2009). However, besides the greater
presence of manta rays in this region
during the seasonal upwelling event
(based on diver photos), no information
was provided regarding foraging
activities or the essential physical or
biological features of the habitat that are
necessary to support this behavior.
Off the coast of Suriname, De Boer et
al. (2015) found that the presence of M.
birostris coincided with the region’s two
rainy seasons. As the outflows of
nutrient-rich waters from the Amazon
and Suriname rivers lead to a low
salinity front during the rainy seasons,
the authors suggest that giant manta rays
are visiting the coastal waters of
Suriname for feeding purposes (De Boer
et al. 2015). Although only a few
observations of manta rays were
recorded during the survey period, the
authors found the behavior was likely
indicative of foraging (i.e., swimming
just below the surface with pectoral fins
curled) (De Boer et al. 2015); however,
again, no physical or biological features
of the foraging habitat were identified.
While upwelling events appear to be
the main environmental factor driving
manta ray foraging behavior, we note
that Graham et al. (2012) also observed
a giant manta ray feeding in oligotrophic
waters during a seasonal fish spawning
event. The giant manta ray was initially
tagged off the northern Yucata´n
peninsula in eutrophic waters and
observed feeding on copepeds (Graham
et al. 2012). However, 57 days later, it
was re-sighted in oligotrophic waters
foraging on fish eggs released during a
seasonal spawning event of little tunny
(Euthynnus alletteratus), suggesting that
giant manta rays are also able to exploit
different habitats when conditions arise
that are suitable for foraging (Graham et
al. 2012).
Overall, based on the foregoing
information regarding known or
presumed foraging areas for giant manta
rays, the general and consistent physical
oceanographic feature that appear to be
associated with foraging habitat is high
primary productivity from upwelling
events, which favors the potential
accumulation of zooplankton. Yet the
levels of primary productivity necessary
to produce suitable foraging habitat are
unknown, and this feature is relatively
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ubiquitous throughout the global range
of the species, with not all areas of high
primary productivity providing
meaningful foraging habitat for giant
manta rays. Furthermore, given that the
characteristics of habitat necessary to
produce areas of high primary
productivity varies by region and site
(e.g., seasonal upwelling events due to
increased river discharge or winddriven fronts), we proceeded to focus
our examination on whether we could
identify any physical and biological
features of giant manta ray foraging
areas within U.S. waters that are
essential to the conservation of the
species.
In general, very little published
literature exists on giant manta ray
occurrence and behavior in U.S. waters.
Adams and Amesbury (1998)
documented the presence of three giant
manta rays in the estuarine waters of the
Indian River Lagoon system and in Port
Canaveral, Florida. Foraging behavior
was not observed and the authors
proposed that individuals likely enter
the estuary sporadically and stay for
only short durations. Freedman and Roy
(2012) used Ocean Biogeographic
Information System (OBIS) data on giant
manta ray observations to examine the
spatial distribution of the species along
the U.S. east coast. They found a higher
number of observations near the
continental shelf edge and bordering the
Gulf Stream, and suggested a seasonal
distribution of the species driven
mainly by temperature, with giant
manta rays primarily observed in waters
from 19 °C to 22 °C (Freedman and Roy
2012). Manta rays are also known to
visit the east coast of Florida, more often
in the spring and summer months,
moving north as water temperatures rise
above 20 °C (Levesque 2019). However,
while it is known that giant manta rays
prefer warmer waters, there is no
evidence that this is a physical or
biological feature that is essential to the
conservation of the species or related to
foraging activity. In fact, as noted in the
literature, giant manta rays can be found
in waters anywhere from 18 °C to 30 °C
(Yano et al. 1999; Freedman and Roy
2012; Graham et al. 2012; Burgess 2017;
Hacohen-Domene´ et al. 2017).
Additionally, the OBIS data, upon
which Freedman and Roy (2012) based
their conclusions, also has inherent
flaws as it is an open-access database
where any member can submit
observations of marine species without
validation. As will be discussed below,
there are significant misidentification
issues associated with M. birostris
observations and conclusions drawn
from this type of sightings data should
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be made with caution as there are
significant uncertainties and limitations
to the data.
In the FGBNMS, Stewart et al. (2018b)
documented high numbers of giant
manta rays but specifically noted that
foraging behavior was rare. Citing a
personal observation (E. Hickerson),
Stewart et al. (2018b) stated that mantas
were only rarely seen exhibiting barrel
rolling behavior (3 of 88 observations),
indicative of feeding, at the banks. In his
study of the Flower Garden Banks and
surrounding banks, Childs (2001)
documented M. birostris feeding
behavior in February and March of 2000
through the use of a remotely operated
vehicle. He noted that M. birostris
generally fed along escarpments and
within the water column over the reef
crest; however, no other details were
provided regarding these events.
In our own examination of the
available data, we compiled manta ray
sightings data (NMFS unpublished data)
from a number of available surveys
(Table 1), photo databases, individual
observations, and social media websites
(e.g., YouTube and Facebook), and
plotted the information to assess
whether we could determine ‘‘hot
spots’’ of giant manta rays, or areas
where manta rays appear to be visiting
consistently over time. We initially
made the main assumption that
sightings of the species were correlated
with areas of high prey (as tends to be
the case with observations of giant
manta rays in other portions of its
range). In other words, when a manta
ray was spotted, we assumed it was
likely because that animal was foraging
in the area, but we also looked for
behavioral (e.g., barrel rolling, mouth
open, cephalic lobes unfurled) or
environmental data (e.g., high plankton
biomass) that could support this
assumption as foraging may not be the
only reason for manta ray presence.
Because most manta sightings within
surveys are opportunistic in surveys
designed for other species, there are
some misidentification issues and gaps
in the time series. Many of the sightings
data were obtained from aerial surveys
aimed at collecting information on the
distribution and abundance of marine
mammals (for example, the Atlantic
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Marine Assessment Program for
Protected Species (AMAPPS) and North
Atlantic Right Whale Consortium data).
This presents a problem as observers on
these surveys are usually not trained in
identifying mobulid rays to the species
level. In discussions with biologist Todd
Pusser, a contract observer for NOAA in
the southeast region during the 1990s
and early 2000s who was then
contracted through the NOAA Northeast
Fisheries Science Center (NEFSC) at
Woods Hole and participated in these
marine mammal surveys from Canada to
Cape Hatteras, North Carolina, he
confirmed that in both the NOAA aerial
and ship surveys along the Atlantic
coast, mobulid sightings were simply
logged as ‘‘manta ray’’ or ‘‘manta spp,’’
thus greatly inflating the sightings data
for M. birostris (T. Pusser, pers. comm.
to C. Jones, NMFS SEFSC, 2018). In fact,
when photos were available from
accompanying ship and aerial surveys,
the majority of the sightings logged as
M. birostris in the northeast Atlantic
were Mobula tarapacana or M. mobular
(T. Pusser, pers. comm. to C. Jones,
NMFS SEFSC, 2018).
TABLE 1—AVAILABLE SURVEY DATASETS WITH REPORTED MANTA SIGHTINGS
Survey name
Year(s)
Survey location
Digital Aerial Baseline Survey—NYSERDA .................................
AMAPPS (aerial) ...........................................................................
North Atlantic Right Whale Consortium database (various surveys).
SEFSC Mid-Atlantic Tursiops Survey (aerial) ..............................
SEFSC Southeast Cetacean Aerial Survey .................................
Florida Manta Project (boat & aerial; directed manta ray survey)
GA Aquarium (boat & aerial; directed manta ray survey) ............
SEFSC Platform Calibration Survey (aerial) ................................
Gulf of Mexico Marine Mammal Assessment Aerial Surveys—
NRDA.
GoMAPPS (aerial) ........................................................................
GulfCet (aerial) ..............................................................................
SEFSC GoMex (aerial) .................................................................
NOAA Coral Reef Ecosystems Program (towed diver survey) ....
2016, 2017 ..................................
2010 through 2018 ......................
1986 through 2017 ......................
Atlantic (38.45° N to 41.08° N).
Atlantic (26.03° N to 45.32° N).
Atlantic (25° N to 41° N).
1994, 1995 ..................................
1992, 1995 ..................................
2016, 2017, 2018 ........................
2010 through 2017 ......................
1991 ............................................
2010, 2011, 2012 ........................
Atlantic (24.5° N to 40.50° N).
Atlantic (26.21° N to 35.19° N).
Atlantic (26.5° N to 27° N).
Atlantic (29.5° N to 29.9° N).
Atlantic (35.8° N to 39.3° N).
Gulf of Mexico (98° W to 80.5° W).
2017,
1992,
1992,
2006,
Gulf of Mexico (97° W to 81° W).
Gulf of Mexico (96.5° W to 84° W).
Gulf of Mexico (96.3° W to 82° W).
Pacific Islands (160° W; Jarvis Island).
2018 ..................................
1993, 1994, 1996,1997 .....
1993, 1994, 1996 ..............
2010 ..................................
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Note: Survey locations are given as geographic regions: Atlantic, Gulf of Mexico, Pacific Islands. For Atlantic locations, the latitude range over
which the surveys were conducted is given. For Gulf of Mexico and Pacific Island locations, the longitude range over which the surveys were
conducted is given.
We similarly found this to be the case
with another available dataset from the
northeast Atlantic that documented 504
sightings of ‘‘Giant Manta Ray’’
(Normandeau Associates and APEM Ltd
2017). This aerial survey, conducted in
2016 and 2017 and supported by the
New York State Energy Research and
Development Authority (NYSERDA),
encompassed the waters of the New
York Bight from Long Island southeast
to the continental shelf break. This
dataset also had accompanying photos
of each animal observation, which a
NMFS species expert was able to review
and confirm that only 6 of the 504
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‘‘giant manta ray’’ sightings were
actually Manta birostris (C. Horn, NMFS
SERO, pers. comm. to M. Miller, NMFS
OPR, 2018). Similarly, in 2015, the
NMFS Northeast Fisheries Observer
Program database underwent a species
verification review whereby NMFS
scientists conducted a detailed review
of observer photo records with the
assistance of manta and devil ray
experts (i.e., Dr. Giuseppe Notarbartolo
di Sciara, Dr. Andrea Marshall, and Guy
Stevens). From 2009 to 2015, there were
25 manta and mobula species records
with photos in the database (J. Hare,
memo, addressed to R.E. Crabtree,
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February 1, 2019). Most of the mobula
bycatch consisted of Mobula
tarapacana, with only two confirmed
records of Manta birostris. These
individuals were observed caught off
the coast of North Carolina. This
observer data appears to further confirm
the rare occurrence of M. birostris in the
U.S. mid-Atlantic and northeast, and
supports the advice provided by species
experts that all M. birostris sightings
north of Cape Hatteras should be
questioned if there are no corresponding
photos.
There may also be occasional
misidentifications of M. birostris south
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of Cape Hatteras as both Mobula
tarapacana and M. mobular are also
common in this portion of the species’
range within the Atlantic (Stevens et al.
2018a, C. Jones unpublished data).
Additionally, M. tarapacana co-occurs
with Manta birostris in the Gulf of
Mexico and Caribbean (Childs 2001),
potentially confounding those aerial
sighting records as well. Thus, while the
presence of M. birostris south of Cape
Hatteras is much more likely (based on
photographic evidence), the proportion
of M. birostris in these datasets to the
other two commonly misidentified
mobula rays is presently unknown,
significantly increasing the uncertainty
of the accuracy of the available sightings
data.
In addition to misidentification rates,
we found other inherent problems with
the sightings data during our analysis,
including the uncertainty regarding
unique sightings and the large gaps in
time between surveys. For aerial
surveys, planes are generally flown
following designated transect lines.
Depending on the transect distance and
timing, there is potential for doublecounting the same animal if the animal
is also moving. Without being able to
view the ventral side of the animal, it is
difficult for aerial observers to identify
whether the manta ray they are spotting
is the same individual from a previous
observation. Aerial surveys are also
subject to availability bias (i.e., the
percentage of time a manta would be
near enough to the surface to be viewed
by an aerial observer) and perception
bias (i.e., the probability of an observer
viewing the animal when it is available).
While it is possible to control for some
of this uncertainty using distanceweighted sampling techniques for
perception bias combined with data
from satellite tags for availability bias,
we do not have the data or information
that would be necessary in order to
conduct this type of analysis at this
time, nor are we aware of any available
studies that have accounted for this
uncertainty in reporting and analyzing
manta ray sightings.
Furthermore, as some of the aerial
surveys were not regularly conducted
on an annual or seasonal basis, but
rather for specific research purposes
that were unrelated to manta ray
distribution or abundance, the resulting
data was skewed in terms of effort in
specific locations and over certain time
periods and could not be used to
identify potential areas used routinely
or repeatedly by giant manta rays. For
example, along the east coast, the
SEFSC Mid Atlantic Tursiops Surveys
(MATS), for which we have manta ray
sightings information, were conducted
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in February of 1994 and July and August
of 1995 to examine the distribution and
estimate an index of relative abundance
for Atlantic bottlenose dolphins
inhabiting nearshore coastal waters in
the mid and southern Atlantic bight. We
also have data from the SEFSC
Southeast Cetacean Aerial Survey,
SECAS, from February to March in 1992
and March of 1995, a survey that was
conducted to estimate cetacean
abundance. The Gulf of Mexico Marine
Mammal Assessment Aerial Surveys—
Natural Resource Damage and
Assessment surveys were only
conducted during the spring and
summer of 2010 and seasonally during
2011 to 2012 to assess the abundance
and spatial distribution of marine
mammals and sea turtles within the
region impacted by the Deepwater
Horizon oil spill. The Atlantic Marine
Assessment Program for Protected
Species (AMAPPS), which conducted
annual aerial surveys from 2010–2017,
had as its main objective assessing the
abundance, distribution, ecology, and
behavior of marine mammals, sea
turtles, and seabirds throughout the U.S.
Atlantic. However, again, these surveys,
as well as others that were analyzed (see
Table 1), varied with respect to the
geographical coverage, years and even
months in which they were conducted.
Currently there are no available analyses
of datasets or studies that control for
spatial and temporal variation in
sampling effort, perception and
availability bias, and potential
misidentification rates to distinguish
areas of high giant manta ray
abundance.
Recently, we became aware of an
ongoing dedicated manta ray aerial
survey, conducted by the Georgia
Aquarium, which has documented
manta ray presence off the east coast of
Florida since 2010. The manta aerial
surveys are conducted in spring and
summer (March/April to June/July) and
follow general track lines 0 to 2.5
nautical miles (0 to 4.63 km) from the
beach that run parallel to the shore,
from St. Augustine Beach Pier (29°52′ N)
to Flagler Beach Pier (29°29′ N). The
number of mantas are counted and,
occasionally, dorsal photos of mantas
are collected during these surveys.
However, due to the murkiness of the
water, photos are rather hard to obtain
if the mantas are too deep in the water
column, and no ventral photos are
available (H. Webb, GA Aquarium, pers.
comm. to M. Miller, NMFS OPR, 2019),
preventing the identification of
individual manta rays or analysis of
potential site fidelity over the course of
multiple years. Overall, the sightings
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data indicate the seasonal visitation of
manta rays to Florida’s inshore waters;
however, the specific physical or
biological features that attract giant
manta rays to this particular area are
poorly understood. The numbers,
location, and peak timing of the manta
rays to this area varies by year, but with
a notable decline in manta rays
observed in the study area since 2015
(H. Webb unpublished data). While sea
surface temperatures are thought to play
a role in the initial migration of manta
rays to the study site, preliminary
analysis suggests that the within-season
temperatures are not strongly correlated
with manta ray distribution or
abundance within the area (H. Webb,
GA Aquarium, pers. comm. to M. Miller,
NMFS OPR, 2019). Although foraging
has been anecdotally observed during
these surveys (H. Webb, GA Aquarium,
pers. comm. to M. Miller, NMFS OPR,
2019) and mentioned in a few online
fishing articles (Roberts 2016; Levesque
2019), we are unaware of any research
that has determined the driving factor of
manta ray occurrence in this area and/
or investigated the physical or biological
features of this area that may be
essential to support the life history
needs of the species. Without
information on specific habitat
characteristics or the relationship
between environmental variables and
manta ray abundance or distribution,
the available sightings data do not allow
us to identify important foraging areas at
this time. A manuscript summarizing
findings from the Georgia Aquarium
sightings dataset is forthcoming (H.
Webb, GA Aquarium, pers. comm. to M.
Miller, NMFS OPR, 2019), and we
intend to review any new information
that becomes available regarding manta
ray use of this area off Florida.
Overall, the best available information
indicates that giant manta rays will feed
on a variety of planktonic organisms
and are not limited by the required
presence of a specific prey species for
successful foraging to occur. Areas of
high primary productivity (e.g.,
upwelling) are generally regarded as
habitat that could potentially support
giant manta ray foraging events;
however, the physical and biological
characteristics of high productivity
areas can vary depending on the
location and season. Additionally, the
presence of these areas does not
necessarily indicate giant manta ray
foraging will occur as the available data
suggest some unknown prey density
threshold may be necessary to facilitate
manta ray foraging or aggregations. In
U.S. waters, foraging has been
anecdotally observed, but the available
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data do not indicate any specific
physical and biological features of these
areas that are essential for facilitating
foraging events or specific sites that are
used consistently for foraging purposes.
For the foregoing reasons, it is not
possible to identify any physical or
biological features related to foraging
that are essential to the conservation of
the species, nor any specific areas that
are essential to support the foraging
needs of the species within waters
under U.S. jurisdiction.
Analysis of the Physical and Biological
Features of Pupping Habitat That Are
Essential to the Conservation of the
Species
Giant manta rays likely give birth to
only one pup per pregnancy after a long
gestation time (12–13 months). This
very low reproductive output for the
species means that the success of
pupping events is essential for the
conservation of the species. Identifying
and protecting important pupping
habitat throughout the species’ range
will be necessary to support recruitment
of young individuals to the recovering
population. Without sufficient nursery
habitat, the population is unlikely to
increase to a level associated with low
extinction risk and delisting. Protection
of the species’ nurseries is crucial
because the rebuilding of the population
cannot occur without protecting the
source (juvenile) population and its
associated habitats. Therefore, using the
best available data, we attempted to
identify potential nursery habitats and
determine whether we could identify
physical or biological features of the
habitat that facilitate successful giant
manta ray pupping and, thus, are
essential for the conservation of the
species.
For the purposes of identifying
potential nursery habitat, we considered
giant manta rays that were less than
4,000 mm DW to be immature, with a
size at birth of ∼2,000 mm DW. As
mentioned previously, juvenile giant
manta rays are rarely observed in the
wild but are present in the fishery
landings data from many countries,
including Sri Lanka, Brazil, Indonesia,
and the Philippines. While this
indicates that fishermen are accessing
potentially important juvenile habitat
and possibly nursery areas, we have no
data on these fishing grounds that could
provide insight into important physical
or biological features of these areas.
However, recent manta ray research in
U.S. waters has documented the
presence of juvenile giant manta rays in
the FGBNMS in the U.S. Gulf of Mexico
as well as off the east coast of Florida,
suggesting the existence of juvenile and
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potential manta ray nursery habitat,
which we discuss below.
For the FGBNMS, both Childs (2001)
and Stewart et al. (2018b) suggested this
area may contain potential nursery
grounds for the species. Although
juveniles are rarely observed globally, a
high number of juveniles were sighted
at several locations in the FGBNMS over
multiple years. Based on an analysis of
NOAA diver logs (from various coral
reef and fish surveys), approximately
171 individual manta rays have been
sighted within the FGBNMS since 1994
(C. Jones unpublished data). Of these,
114 have approximate recorded sizes.
Around 97 percent of the individuals
sighted were less than 4 m DW (i.e.,
immature), and around 50 percent were
2 m DW (i.e., estimated size at birth of
M. birostris) or less. However, M. cf.
birostris may comprise the majority of
these sightings as Stewart et al. (2018b)
noted that at least 55 percent of the
manta rays identified in their study
likely belong to M. cf. birostris, which
is thought to be closer in size to M.
alfredi (Stevens et al. 2018a) and
potentially explains the observations of
mantas with sizes smaller than the
estimated size at birth for M. birostris.
Using the nursery habitat criteria
proposed by Heupel et al. (2007),
Stewart et al. (2018b) suggested that the
FGBNMS may contain nursery habitat
for giant manta rays because juveniles,
which are generally rare, are found in
this area, remain in the area for a period
of several days to months, and have
been sighted with gaps of more than a
year between re-sightings. The FGBNMS
is a unique area, situated over 100 miles
offshore of the Texas/Louisiana border
and comprised of shallow, underwater
features, called salt domes, upon which
diverse coral reef communities have
developed and thrived. There is
substantial upwelling, distinct
thermoclines, and unique eddies that
form in the area, presumably due to
interactions between currents and the
pronounced benthic features. Stewart et
al. (2018b) proposed that the FGBNMS
may be an optimal nursery ground
because it contains habitat near the edge
of the continental shelf and in proximity
to abundant pelagic food resources.
Important prey for manta rays, like
euphausiids, are abundant in the deep
scattering layers in the basin waters of
the Gulf of Mexico (Stewart et al.
2018b). The authors state that an
additional benefit of the FGBNMS is
that the shallow bottom habitat may
protect juvenile rays from predation
while they rest and recover their body
temperature in the warm mixed layer
after deep foraging dives (Stewart et al.
2018b).
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However, while the FGBNMS
provides habitat for juvenile giant manta
rays, the available data do not indicate
any specific physical and biological
features within the FGBNMS that are
essential for supporting pupping
behavior or necessary for a manta ray
nursery. For example, in examining
specific physical features, like
temperature, we found that the majority
of individuals (∼75 percent) at the
FGBNMS were sighted between July and
September (Stewart et al. 2018b). Sea
surface temperatures during these
sightings ranged from 20 °C to 32 °C,
with ∼75 percent of mantas observed in
28 °C to 31 °C (C. Jones unpublished
data). However, dives during which
observations were collected were
skewed towards summer months (i.e.,
warmer temperatures) and specific sites
and depths (limited to areas above 150
ft (45.7 m)), meaning that the increased
observations of giant manta rays in the
higher temperature range may be a
consequence of the survey methodology
and not a reflection of an essential
feature of the habitat.
Next, we reviewed the available data
regarding behavior to see if we could
identify specific habitat features based
on use of the habitat that are necessary
to support pupping. As stated in Stewart
et al. (2018b) and Childs (2001), the
primary behavior of manta rays
observed in the FGBNMS was mainly
swimming, with manta rays swimming
above reef crest and sand flats, along
escarpments, and in the water column.
Although more juveniles were sighted at
East and West Flower Garden Banks
(hermatypic coral habitat) than at
Stetson Bank (silt/claystone dominated
coral community), acoustic telemetry
tagging has shown that juvenile mantas
move between East, West, Stetson, and
Bright Bank within FGBNMS (R.
Graham, Wildlife Conservation Society,
pers. comm. to C. Horn, NMFS SERO,
2018). Stewart et al. (2018b) suggest the
FGBNMS likely provides ample feeding
opportunities for juveniles, but they
acknowledge that foraging behavior is
only rarely observed. Similarly, Childs
(2001) mentioned that foraging behavior
at the FGBNMS was observed in only
two months (February and March) of his
study despite manta rays occurring in
the area during all months.
While the presence of young giant
manta rays suggest potential pupping in
the vicinity of the area (Childs 2001),
the available data do not allow us to
identify where this pupping is
occurring. Additionally, the available
data do not explain why or how giant
manta rays are using this particular
habitat (e.g., foraging, transiting, resting)
or allow us to identify the essential
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physical or biological features of the
habitat. Therefore, we cannot identify
any pupping areas that meet the
definition of critical habitat.
Research (supported by NMFS and
the National Ocean Service, in
collaboration with the Manta Trust) on
the movements and genetics of giant
manta rays continues in the FGBNMS
and may help provide answers to these
questions in the future. However, at this
time, the available data do not indicate
any physical or biological features of
this habitat that are essential for the
conservation of the species.
Similar to the FGBNMS, juvenile M.
birostris have also been regularly
observed off the east coast of Florida in
the past several years. Since 2016,
researchers with the Marine Megafauna
Foundation have been conducting
annual surveys along a small transect off
Palm Beach, Florida, between Jupiter
Inlet and Boynton Beach Inlet (∼44 km,
24 nautical miles) (J. Pate, MMF, pers.
comm. to M. Miller, NMFS OPR, 2018).
Results from these surveys indicate that
juvenile manta rays are present in these
waters for the majority of the year
(observations span from May to
December), with re-sightings data that
suggest some manta rays may remain in
the area for extended periods of time or
return in subsequent years (J. Pate
unpublished data). For example, one
satellite tagged male has been re-sighted
multiple times in the past 3 years
(Marine Megafauna Foundation 2019).
However, similar to the limitations of
the FGBNMS data and the level of
resolution, it is currently unclear what
physical or biological characteristics of
this habitat are necessary to facilitate
successful pupping behavior or are
essential for nursery habitat. Manta rays
are difficult to detect using boat-based
observation. When an observer spotted
a manta ray, he/she would get into the
water and collect habitat information,
behavioral data, as well as photos of the
manta ray. This type of data collection
has limitations. For example, water
turbidity, depth, and weather conditions
may make manta rays harder to spot
from a boat. As such, the fact that the
majority of manta rays were spotted
over sand is likely due to increased
visibility over this type of habitat
compared to others (such as reef habitat)
(J. Pate, personal communication, 2018)
as opposed to a biological necessity for
this type of habitat. Additionally, the
main behavior observed in the transect
area was swimming, with occasional
observations of foraging behavior near
Jupiter Inlet (J. Pate, MMF, pers. comm.
to M. Miller, NMFS OPR, 2018). In other
words, similar to the FGBNMS, the
available data only indicate juvenile
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manta ray presence in these areas and
does not explain why or how giant
manta rays are using the particular
habitat that would help us identify any
physical or biological features that are
essential for the conservation of the
species. We also note that the majority,
if not all, of these juvenile manta rays
observed off the east coast of Florida are
thought to be M. cf. birostris (J. Pate,
MMF pers. comm. to M. Miller, NMFS
OPR, 2018) and not M. birostris. NMFS
researchers are currently collaborating
with colleagues at the Marine
Megafauna Foundation to tag these
manta rays off the Florida coast and
collect genetic information in order to
inform taxonomy, determine population
structure, and learn more about their
movements to gain a better
understanding of their habitat use in
this region. Anecdotal observations from
some of these recent tagging trips (June
and August 2019) suggest this area may
provide foraging opportunities (N.
Farmer, NMFS SERO, pers. comm. to M.
Miller, NMFS OPR, 2019); however,
further investigation is required as the
available information does not indicate
any specific physical and biological
features of this area that are essential to
support the life-history needs of the
species.
We also obtained anecdotal
observations of juvenile giant manta
rays in the U.S. Caribbean from off
Puerto Rico (n=10; sightings dating back
to 2004) and the U.S. Virgin Islands
(n=16; sightings dating back to 2012),
and in the U.S. Pacific from off Hawaii
and the Pacific Remote Island Areas
(n=24; sightings dating back to 2003)
that indicate the use of these waters by
young giant manta rays (NMFS
unpublished data). However, as stated
before, simply the observation of the
presence of juveniles using these waters
(and further confounded by a lack of
known abundance, duration, movement,
or frequency of occurrence in these
areas) is not enough information to
indicate that these areas contain
physical and biological features that are
essential to the conservation of the
species.
In summary, while we have evidence
of the presence and use of specific areas
by juvenile giant manta rays, the
available information does not allow us
to identify any physical or biological
features within these areas that are
essential to support the life-history
needs of the species. Without
knowledge of the essential features that
create meaningful pupping and nursery
grounds, we cannot identify any areas
that meet the definition of critical
habitat at this time.
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The Physical and Biological Features of
Migratory Habitat That Are Essential to
the Conservation of the Species
Based on the available data, it is
evident that both small and large-scale
migratory movements are a necessary
component in the life-history of the
giant manta ray. Seasonal sightings data
suggests that large-scale movements are
undertaken primarily for foraging
purposes, correlated with the movement
of zooplankton and influenced by
current circulation and tidal patterns,
seasonal upwelling, and seawater
temperature (Luiz et al. 2009; Couturier
et al. 2012; Freedman and Roy 2012;
Graham et al. 2012; Sobral and Afonso
2014; De Boer et al. 2015; Girondot et al.
2015; Armstrong et al. 2016; HacohenDomene´ et al. 2017). Small-scale
movements also appear to be associated
with exploiting local prey patches in
addition to refuging and cleaning
activities (O’Shea et al. 2010; Marshall
et al. 2011; Graham et al. 2012; Rohner
et al. 2013; Stewart et al. 2016a; Stewart
et al. 2016b; Sotelo 2018). However, as
sightings of giant manta rays tend to be
sporadic, with the species more
commonly found offshore and in
oceanic waters, it is difficult to track
small-scale and large-scale migratory
behavior of the species. For logistical
reasons, survey effort tends to be
focused in nearshore habitats. Yet,
through the opportunistic tagging of
giant manta rays with pop-up satellite
archival tags when in these nearshore
areas, researchers have been able to
provide evidence of the migratory
nature of giant manta rays and
demonstrate the species’ ability to make
large-scale migrations. For example,
satellite tracking has registered
movements of the giant manta ray from
Mozambique to South Africa (a distance
of 1,100 km), around Ecuador and its
islands (between the Isla de la Plata,
Bajo Cope, and Isla Santa Clara (El Oro,
Ecuador); around 230 km), and from the
Yucata´n, Mexico into the Gulf of Mexico
(448 km) (Marshall et al. 2011; Guerrero
and Hearn 2017; Sotelo 2018). Off
Mexico’s Yucata´n peninsula, Graham et
al. (2012) calculated a maximum
distance travelled by a giant manta ray
to be 1,151 km (based on a cumulative
straight line distance between locations;
tag period ranged from 2 to 64 days).
Similarly, Hearn et al. (2014) report on
a tagged M. birostris that was tracked
from Isla de la Plata (Ecuador) to west
of Darwin Island (tag was released after
104 days), a straight-line distance of
1,500 km, further confirming that the
species is capable of fairly long distance
migrations.
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For the most part, these larger-scale
migrations appear to be seasonallybased for foraging purposes, as
described previously, with giant manta
rays appearing in areas undergoing
seasonal upwelling events. For example,
through analysis of photographs and
videos of mobulids from 1990 to 2013,
Sobral and Afonso (2014) confirmed the
presence of M. birostris at the Azores
islands and noted that its occasional
presence (several encounters per year) at
these remote islands indicates a strong
seasonal migratory behavior. However,
the origin of these mantas, and the
potential migratory paths that they use
to get to these remote islands, remain
unknown.
Similarly, seasonal sightings of M.
birostris off the Isla de la Plata, Ecuador,
predominantly occur from August to
October, with a peak in early September
(Guerrero and Hearn 2017); however,
from where these mantas originate is
currently under investigation. Recently,
Sotelo (2018) examined the genetic
diversity of these manta rays from 2010
to 2013 and found that it was
moderately high, with an average
expected heterozygosity value (He =
0.679) comparable to similar species
that are known to undertake longdistance migrations. The results also
suggest that the manta rays may migrate
in family groups, but that they may not
always visit the same areas consistently.
For example, Sotelo (2018) found
population structure between the manta
rays sampled in 2013 compared to the
years 2010, 2011, and 2012, with the
2013 manta rays representing a different
population. The authors note that
copepod numbers peaked at the Isla de
la Plata in May of 2013, two months
later than the previous years in the
study (Sotelo 2018). As manta rays
demonstrate high plasticity in terms of
their movements in search of prey,
Sotelo (2018) reasoned that the change
in timing of the copepod peak likely
explains why a different manta ray
population visited the island in 2013
compared to previous years. However,
again, the origin of these mantas, and
the potential migratory routes traveled
by these mantas to the Isla de la Plata
are currently unknown.
While long-distance migratory
information is lacking, scientists have
tagged some of these mantas during
their seasonal visitation to these
nearshore areas, and have gained
additional information on their smallerscale movement patterns around and
from these sites. For example, in Isla de
la Plata, two mantas were tagged from
September 2017 to January 2018 with
tracks that revealed coastal movements
between Ecuador and northern Peru
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(Sotelo 2018). These two mantas
remained within 200 km of the
shoreline and did not move more than
300 km south of Isla de la Plata, where
they were originally tagged. However,
based on the track lines (see Annex C;
Sotelo 2018), there is no clear migratory
corridor that they appear to use, with
movements traversing throughout the
entire area.
Off the Yucata´n peninsula, Graham et
al. (2012) tagged 6 giant manta rays (4
females, 1 male, and 1 juvenile) and
tracked their movements for up to 64
days. The tagged manta rays traversed
the frontal zones repeatedly, probably in
search of prey (Graham et al. 2012), with
no clear migratory route. The majority of
manta ray tracks were more than 20 km
offshore, in water depths of less than 50
m, and the animals traveled up to 116
km from their original tagging location
(Graham et al. 2012). The authors also
noted that there were no differences in
movement patterns based on sex, body
size, or ambient water-column
temperature. Their conclusion, based on
the tracking data, was that giant manta
rays forage over large spatial scales
(∼100 km long) that are too far offshore
and wide-ranging to be completely
captured in the existing Marine
Protected Area networks within the
Mexican Exclusive Economic Zone
(Graham et al. 2012). In other words,
there does not appear to be a specific
migratory corridor that dictates these
smaller-scale foraging movements.
Rather, manta rays appear to be
opportunistic feeders, with movements
in and around frontal zones or areas that
are likely to contain prey.
While the available data indicate that
giant manta rays may be capable of longdistance movements, a recent study by
Stewart et al. (2016a) suggests that the
species may not be as highly migratory
as previously thought. Using pop-up
satellite archival tags in combination
with analyses of stable isotope and
genetic data, the authors found evidence
that M. birostris off the Pacific coast of
Mexico may actually exist as wellstructured subpopulations that exhibit a
high degree of residency. For example,
unlike the giant manta ray in the Hearn
et al. (2014) study (that traveled from
Isla de la Plata to the Galapagos Islands),
tagged M. birostris individuals from
locations nearshore to Mexico (Bahia de
Banderas; n=5) and offshore Mexico
(Revillagigedo Islands; n=4) showed no
movements between locations (tag
deployment length ranged from 7 days
to 193 days) (Stewart et al. 2016a). The
stable isotope analysis showed higher
d13C values for the nearshore mantas
compared to those offshore, indicating
these mantas were foraging in their
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respective locations rather than moving
between nearshore and offshore
environments (Stewart et al. 2016a).
Additionally, the genetic analysis
provided evidence of population
structure between the coastal Mexico
and offshore Mexico populations
(Stewart et al. 2016a). While the authors
note that the species may be capable of
occasional long-distance movements,
the results from their study indicate
that, for some populations, these types
of long-distance movements may be rare
and may not contribute to substantial
gene flow or inter-population mixing of
individuals (Stewart et al. 2016a).
Overall, the available data indicate
that giant manta rays undergo both
short- and long-distance migrations;
however, the space or any specific
migratory corridor used by the species
during these migrations remains
unknown. In addition, we have no
information on any potential migratory
corridors that may exist within waters
under U.S. jurisdiction for the giant
manta ray. As mentioned previously, we
are currently supporting and conducting
tracking studies of giant manta rays
within U.S. waters to better understand
the fine-scale movements of the species
off the coast of Florida and within the
FGBNMS. Data from these or similar
studies may reveal potential migratory
corridors preferred by giant manta rays.
Similarly, survey efforts by the Georgia
Aquarium off the coast of St. Augustine,
Florida, may also help elucidate some of
these questions in the future.
As noted previously in this
determination, giant manta rays appear
to have a seasonal pattern of occurrence
along the east coast of Florida, showing
up with greater frequencies (and in
greater numbers) in the spring and
summer months. In fact, sightings of
manta rays in the region signal to
fishermen the start of cobia fishing as
fishermen have found that cobia tend to
closely associate with the manta rays as
they migrate along the east coast of
Florida. Based on information from
recreational cobia fishing articles, manta
rays tend to appear off Florida’s coast
when water temperatures climb above
20 °C to 21 °C; however, Levesque
(2019) notes that it is ‘‘impossible to
predict when they will show up from
one year to the next.’’ Killer (2010)
states that in Florida’s Treasure Coast
waters, mantas may not show up every
year, and it is unclear where they come
from or where they go after they leave
the area. Quoting two charter vessel
captains, Killer (2010) reports that the
mantas have been observed along the
coast moving from south to north as
waters warm, but have also been
observed doing the opposite migration,
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with some potentially moving from
offshore to inshore waters as well
during this time. McNally (2012)
believes that the spring migration of
rays off northeast Florida is occurring
much farther offshore than in the past,
noting that the mantas used to be
observed just off the beach breakers but
are now more than 10 miles offshore.
We also note that during the migratory
season, manta rays tend to be found in
both shallow and deep waters (Killer
2010; Levesque 2019), with no
information to suggest they are
restricted to a certain area off the coast
of Florida.
While the available information
confirms the migratory behavior of the
species in U.S. waters, the data do not
indicate that there are any specific
routes or corridors that are consistently
used by the species during their
migration. In fact, as noted previously,
McNally (2012) suggests that a
dedicated corridor may not exist, or that
some other unknown feature may be
influencing their spatial patterns during
these migrations. Additionally, Roberts
(2016) notes that ‘‘no studies have
shown a correlation of bottom structure
(reef lines, continental shelf, etc.) and
the ray’s migration pattern,’’ nor have
we come across any studies since that
article was published. Therefore, at this
time, and based on the foregoing
information, we cannot identify any
specific essential features that define
migratory habitat for giant manta rays.
The Physical and Biological Features of
Breeding Habitat That Are Essential to
the Conservation of the Species
Little information exists on the
reproductive ecology of the giant manta
ray as mating behavior of M. birostris is
rarely observed in the wild. However,
based primarily on observations of M.
alfredi mating behavior, Stevens et al.
(2018b) identified seven stages of
courtship for manta rays: (1) Initiation,
(2) endurance, (3) evasion, (4) precopulation positioning, (5) copulation,
(6) post-copulation holding, (7)
separation. The initiation stage involves
males shadowing females at normal
cruising speeds. During this stage, males
will often attempt to facilitate female
receptiveness by using the cephalic fins
to gently stroke the females’ dorsal
surface. During the endurance stage,
swimming speeds increase and from 1
to 8 males follow closely behind a single
female. The evasion stage is
characterized by continued close
following at increased speeds with the
female incorporating rapid maneuvers,
somersaults, and flips, with males
attempting to stay right behind her. Precopulation positioning involves the
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male using his cephalic fins to guide
himself down the females’ back along
the leading edge of her pectoral fin.
Once at the fin’s tip, the male grasps it
firmly with his mouth then rotates his
body so that he is underneath the female
and the two are abdomen to abdomen.
Copulation then occurs, usually
initiating near the surface, with the male
continuing to move his fins to maintain
position while the female ceases
movement. The clasper is inserted in
the cloaca and copulation lasts between
30 and 90 seconds, while the pair
slowly sinks (Stevens et al. 2018b).
Only a few instances of courtship
involving giant manta rays have actually
been observed, with only a single
instance resulting in copulation. On two
separate occasions, in early August 1996
at the Ogasawara Islands, Japan, Yano et
al. (1999) witnessed a male M. birostris
chasing closely behind a female at
relatively high speeds (∼10 km/hr). In
both instances, the behavior was
observed for approximately 40 minutes
but did not result in copulation. Stevens
et al. (2018b) also witnessed two
occurrences of this ‘‘endurance’’ stage in
M. birostris, one involving a single
female followed by a single male, and
the other involving a single female
followed by eight males. Both of these
observations were made off of the
remote island of Fuvahmulah in the
Maldives, lasted approximately one
minute, and neither resulted in
observed copulation. The only
observation of successful copulation
was reported by Yano et al. (1999) who
witnessed two males chasing a single
female in a zigzag pattern off the
Ogasawara Islands in early July 1997.
Speeds were similar to those witnessed
during other observations; however,
these chases progressed all the way
through the rest of the stages of
copulatory behavior (Yano et al. 1999).
The chases occurred approximately 30
minutes apart, with both males observed
inserting their claspers into the same
female (Yano et al. 1999).
In terms of habitat characteristics, the
mating behavior in the Maldives
location occurred at a known
aggregation site for the species (Stevens
et al. 2018b). Females were chased along
the reef crest of the atolls in the area
(Stevens et al. 2018b). However, while
the authors noted that most of the
mating behavior for M. alfredi happened
at cleaning stations, for M. birostris, the
mating occurred at locations where
giant manta rays tend to just pass
through (Stevens et al. 2018b). In other
words, the area where the mating
behavior was observed did not appear to
have any other significance for the
species. Off the Ogasawara Islands,
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Japan, Yano et al. (1999) described the
site of the mating behavior as 100–200
m offshore of the east coast of
Chichijima (one of the Ogasawara
Islands), within an area comprised of
rocky reefs in 10–20 m depth. The
authors noted that each copulation
event happened within one meter of the
surface (Yano et al. 1999).
Giant manta ray breeding sites are
also thought to occur off Ecuador and
the Galapagos Islands based on the
presence of pregnant females and recent
mating scars. In fact, some of the first
pregnant females ever seen in the wild
have been sighted in the productive
coastal waters off Isla de la Plata in the
Machalilla National Park, Ecuador.
According to Guerrero and Hearn
(2017), between 2009 and 2015, 8
pregnant giant mantas were observed off
Isla de La Plata, with 7 of these reported
in 2011. Additionally, photographic
records from 2012 to 2015 showing
fresh scars on the pectoral fins of mature
female giant manta rays around Isla de
la Plata and Bajo Cope´ indicate the
likely use of these Ecuadorian
aggregation sites as mating areas
(Guerrero and Hearn 2017). In terms of
habitat characteristics of these areas, the
authors note that the majority of giant
manta rays seen in Isla de la Plata are
off the northwest area of the island, in
Punta El Faro, Roca Honda, and La
Pared (Guerrero and Hearn 2017). These
particular areas are close to deep waters,
with a bottom characterized by coarse
sand and scattered rocks. Calcareous
coral formations can be found between
0 and 14 m depths and soft corals
(gorgonians) can be found in deeper
depths (Guerrero and Hearn 2017). La
Pared, in particular, contains pinnacles
and rocks that extend to the northwest
and create an edge with a steep drop to
52 m depths (Guerrero and Hearn 2017).
The authors state that giant manta rays
do not remain in the area for very long
(usually around a few days to a week),
but may return in multiple years and
hypothesize that their purpose for
visiting the island could be primarily for
cleaning purposes, mating, and/or
feeding as all three behaviors are
observed at this site (Guerrero and
Hearn 2017).
Within U.S. waters, there are very few
observations of mating behavior. In our
collection of manta ray sightings and
videos, there are only 4 records of
‘‘chasing’’ or ‘‘courtship’’ behavior of M.
birostris. Three of the records are from
diver observations off the west coast of
Hawaii (Manta Pacific Research
Foundation 2019), and the fourth is
from an instagram video off Avon
Fishing Pier, North Carolina, taken in
July 2019 (G. Stevens, Manta Trust,
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pers. comm. to C. Horn, NMFS SERO,
2019); however, there is no
corresponding information regarding
habitat features related to these records
(just individual sightings data). Given
that the areas where giant manta ray
mating occurs remain largely unknown,
with only a few, opportunistic
observations of courtship behavior or
evidence of breeding (i.e., mating scars,
pregnant females) in a couple of
locations, there has not been any
systematic evaluation of the particular
physical or biological features that
facilitate or are necessary for mating to
occur. The general habitat
characteristics mentioned above in
relation to the observations of mating
behavior, including presence of rocky
and coral reefs, shallow depths, coarse
sand, and reef crests adjacent to deep
water, are found throughout the species’
range and are commonly associated
with giant manta ray sightings (Yano et
al. 1999; Childs 2001; Kashiwagi et al.
2011; Marshall et al. 2011; Stevens et al.
2018b; Stewart et al. 2018b). However,
not all areas with the above features
provide meaningful mating habitat as,
for example, many of the observations
from the studies previously discussed
(for foraging, pupping, and migratory
habitat) also noted the presence of these
habitat features but did not observe
mating behavior in M. birostris. As such,
at this time, the available information
does not allow us to identify any
physical or biological features within
these areas where mating has been
observed that are essential to support
this behavior.
Unoccupied Areas
Section 3(5)(A)(ii) of the ESA defines
critical habitat to include specific areas
outside the geographical area occupied
by a threatened or endangered species at
the time it is listed if the areas are
determined by the Secretary to be
essential for the conservation of the
species. Regulations at 50 CFR
424.12(b)(2) address designation of
unoccupied area as critical habitat and
the regulations at 50 CFR 424.12(g) state
that critical habitat shall not be
designated within foreign countries or
in other areas outside of United States
jurisdiction.
As discussed previously, the waters
off the U.S. west coast are not
considered part of the geographical area
occupied by giant manta ray at the time
of listing. We also conclude that it is not
an unoccupied area essential to the
species’ conservation given the rare,
errant use of the area by a vagrant giant
manta ray in the past, and no
information to suggest the area is
essential to the conservation of the
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17:38 Dec 04, 2019
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species. The other geographical areas
under U.S. jurisdiction that were not
included in the discussion of occupied
areas by the giant manta ray (i.e., U.S.
waters north of Long Island, New York)
are considered to be out of the species’
livable range and, thus, would not be
essential to the conservation of the
species. As such, we find that there are
no specific areas outside the
geographical areas occupied by M.
birostris that would meet the definition
of critical habitat for the giant manta
ray.
Critical Habitat Determination
Given the best available information
and the above analysis of this
information, we find that there are no
identifiable occupied areas under the
jurisdiction of the United States with
physical or biological features that are
essential to the conservation of the
species or unoccupied areas that are
essential to the conservation of the
species. Therefore, we conclude that
there are no specific areas within the
giant manta ray range and under U.S.
jurisdiction that meet the definition of
critical habitat. Per 50 CFR
424.12(a)(1)(iv), if no areas meet the
definition of ‘‘critical habitat,’’ then we
can conclude that a designation of
critical habitat is not prudent.
Although we have made this ‘‘not
prudent’’ determination, the areas
occupied by giant manta rays under U.S.
jurisdiction will continue to be subject
to conservation actions implemented
under section 7(a)(1) of the ESA, as well
as consultation pursuant to section
7(a)(2) of the ESA for Federal activities
that may affect the giant manta ray, as
determined on the basis of the best
available information at the time of the
action. Through the consultation
process, we will continue to assess
effects of Federal actions on the species
and its habitat.
Additionally, we remain committed to
promoting the recovery of the giant
manta ray through both domestic and
international efforts. As noted in the
proposed and final rules (82 FR 3694,
January 12, 2017; 83 FR 2916, January
22, 2018, respectively), the most
significant threat to the giant manta ray
is overutilization by commercial and
artisanal fisheries operating within the
Indo-Pacific and eastern Pacific portions
of its range, primarily in areas outside
of U.S. jurisdiction. Giant manta rays
are both targeted and caught as bycatch
in a number of fisheries throughout
their range, and while the majority of
these fisheries target manta rays for their
meat, there has been an increasing
demand for manta ray gill plates for use
in Asian medicine, primarily in the
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66663
Indo-West Pacific. Efforts to address
overutilization of the species through
regulatory measures appear inadequate,
with evidence of targeted fishing of the
species despite prohibitions in a
number of countries, and only one
regional fisheries management
organization measure to address bycatch
issues (Miller and Klimovich 2017).
Thus, recovery of the giant manta ray is
highly dependent upon international
conservation efforts. To address this, we
have developed a recovery plan outline
that provides our preliminary strategy
for the conservation of the giant manta
ray. This outline can be found on our
website at: https://
www.fisheries.noaa.gov/species/giantmanta-ray# resources and provides an
interim recovery action plan as well as
preliminary steps we will take towards
the development of a full recovery plan.
Currently, we are actively engaged in
manta ray research to gain a better
understanding of the biology, behavior,
and ecology of this threatened species.
We are presently working on collecting
and assimilating anecdotal and surveyrelated manta sightings and effort data
to support the development of an
ensemble species distribution model for
the southeastern United States. We are
also collaborating with partners to
examine giant manta ray movements in
U.S. waters off Florida and within the
FGBNMS. This data will provide a
better understanding of giant manta ray
movements and habitat use, including
environmental drivers of movement. We
are also supporting research projects
assessing the survivorship of giant
manta rays caught in Peruvian and
Indonesian artisanal gillnet fisheries.
We have developed safe handling and
release guidelines for fishermen
(available at: https://www.fisheries
.noaa.gov/webdam/download/
91927887). In an effort to address
species identification issues during
aerial surveys, we have also developed
an aerial survey mobulid species
identification key that will facilitate
accurate species identification in the
future. We added the giant manta ray to
our Northeast and Southeast Observer
Program capture reports, logbooks, and
manuals/reports, and provided a guide
to the identification of mobulid rays to
observers to gain more accurate
information regarding the species’
distribution and prevalence in U.S.
fisheries. In addition, we have set up a
dedicated email (i.e., manta.ray@
noaa.gov) for the public to report giant
manta ray encounters to help us learn
more about M. birostris movement
patterns, habitat use, and human
interactions in our waters. We will
continue to work towards the
E:\FR\FM\05DEN1.SGM
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Federal Register / Vol. 84, No. 234 / Thursday, December 5, 2019 / Notices
conservation and recovery of giant
manta rays, both on a domestic and
global level, including with our
international partners and within
regional fisheries management
organizations and other international
bodies to promote the adoption of
conservation and management measures
for the threatened giant manta ray.
References
A complete list of all references cited
herein is available upon request (see FOR
FURTHER INFORMATION CONTACT).
Authority
The authority for this action is the
Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: December 2, 2019.
Samuel D. Rauch III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2019–26265 Filed 12–4–19; 8:45 am]
BILLING CODE 3510–22–P
DEPARTMENT OF DEFENSE
Department of the Army, Corps of
Engineers
Occupational Safety and Health
Programs for Federal Employees
AGENCY:
U.S. Army Corps of Engineers,
DoD.
ACTION:
Notice; request for comments.
The U.S. Army Corps of
Engineers (USACE) Safety and Health
Requirements Manual (EM 385–1–1) is
the gold standard for Safety and
Occupational Health regulations. The
manual holds a long history dating back
to 1941 and is designed to facilitate the
standardization of all safety programs.
The EM 385–1–1 prescribes the safety
and health requirements for all Corps of
Engineers activities and operations. The
USACE is soliciting comments on the
proposed revisions to EM 385–1–1.
DATES: Consideration will be given to all
comments received by January 30, 2020.
ADDRESSES:
Electronic: You may submit
comments concerning the EM 385–1–1
electronically by accessing the USACE
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www.usace.army.mil/missions/safetyand-occupational-health/next-genem385-1-1/. Follow the instructions at
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Mail/Hand Delivery/Courier: Due to
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SUMMARY:
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hand delivery or courier. Comments for
considerations will only be accepted by
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FOR FURTHER INFORMATION CONTACT:
Steven Washington, Headquarters, U.S.
Army Corps of Engineers, Safety and
Occupational Health Office, in
Washington, DC at 202–761–7678.
SUPPLEMENTARY INFORMATION: Executive
Order (E.O.) 12196, Occupational Safety
and Health Programs for Federal
Employees, was issued in 1980 and
directed agencies heads to (1) Furnish to
employees places and conditions of
employment that are free from
recognized hazards that are causing or
are likely to cause death or serious
physical harm; (2) Operate an
occupational safety and health program
in accordance with the requirements of
this order and basic program elements
promulgated by the Secretary. DoDI
6055.1 was issued in 2014 (incorporated
changes in 2018) and the DoD policy.
Following issuance of DoD Safety and
Occupational Health (SOH) Program
DODI 6055.01; the AR–385–10, Army
Safety Program implements the
requirements of the Occupational Safety
and Health Act of 1970 as implemented
in E.O. 12196; 29 CFR 1960; DODI
6055.1; DoDI 6055.4; and DoDI 6055.7.
Following the issuance of the AR–385–
10; the EM 385–1–1 U.S. Army Corps of
Engineers Safety and Health
Requirements Manual prescribes the
safety and health requirements for all
Corps of Engineers activities and
operations. The manual applies to
Headquarters, US Army Corps of
Engineers (HQUSACE) elements, major
subordinate commands, districts,
centers, laboratories, and field operating
activities (FOA), as well as USACE
contracts and those administered on
behalf of USACE. Applicability extends
to occupational exposure for missions
under the command of the Chief of
Engineers, whether accomplished by
military, civilian, or contractor
personnel. USACE intends to update the
manual within two years, and
periodically thereafter, to reflect such
public input, experience, and
innovation. The agency will address
significant comments received in the
next revision of this manual. All
comments are welcomed and
encouraged for any section of the EM
385–1–1, Safety and Health
Requirements Manual. All suggestions
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Rewrite PDT for adjudication. A process
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PO 00000
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collected. The goal of the collection
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manual. All input and comments
received will help improve the overall
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Instructions for Providing Comments
USACE is requesting assistance in the
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clarify the policy requirements for
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USACE Publications website https://
www.publications.usace.army.mil/
Portals/76/Publications/
EngineerManuals/EM_385-1-1.pdf).
Additionally, a series of questions has
been provided for reviewers to consider
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USACE welcomes any and all feedback
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Comments and responses based on
the current version of the Safety and
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E:\FR\FM\05DEN1.SGM
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Agencies
[Federal Register Volume 84, Number 234 (Thursday, December 5, 2019)]
[Notices]
[Pages 66652-66664]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2019-26265]
-----------------------------------------------------------------------
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
[Docket No. 191127-0095; RTID 0648-XR030]
Endangered and Threatened Species; Determination on the
Designation of Critical Habitat for Giant Manta Ray
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce.
ACTION: Notice of critical habitat determination.
-----------------------------------------------------------------------
SUMMARY: We, NMFS, have determined that a designation of critical
habitat is not prudent at this time. Based on a comprehensive review of
the best scientific data available, we find that there are no
identifiable physical or biological features that are essential to the
conservation of the giant manta ray within areas under U.S.
jurisdiction. We also find that there are no areas outside of the
geographical area occupied by the species under U.S. jurisdiction that
are essential to its conservation. As such, we find that there are no
areas within the jurisdiction of the United States that meet the
definition of critical habitat for the giant manta ray.
DATES: This finding is made on December 5, 2019.
ADDRESSES: Electronic copies of the determination, list of references,
and supporting documents prepared for this action are available from
the NMFS Office of Protected Resources website at https://www.fisheries.noaa.gov/species/giant-manta-ray.
FOR FURTHER INFORMATION CONTACT: Maggie Miller, NMFS, Office of
Protected Resources, (301) 427-8403.
SUPPLEMENTARY INFORMATION:
Background
On January 22, 2018, we published a final rule to list the giant
manta ray (Manta birostris) as a threatened species under the
Endangered Species Act (ESA) (83 FR 2916). Section 4(b)(6)(C) of the
ESA requires the Secretary of Commerce (Secretary) to designate
critical habitat concurrently with making a determination to list a
species as threatened or endangered unless it is not determinable at
that time, in which case the Secretary may extend the deadline for this
designation by 1 year. At the time of listing, we concluded that
critical habitat was not determinable because sufficient information
was not available to: (1) Identify the physical and biological features
essential to the conservation of the species at an appropriate level of
specificity, particularly given the uncertainty regarding habitats
required to support its life history (e.g., pupping and nursery grounds
were unknown) and migratory movements, (2) determine the specific
geographical areas that contain the physical and biological features
essential to conservation of the species, particularly given the global
range of the species, and (3) assess the impacts of the designation. We
requested relevant information from interested persons to help us
identify and describe the physical and biological features essential to
the conservation of the giant manta ray, and assess the economic
consequences of designating critical habitat for the species. We
solicited input from the public, other concerned government agencies,
the scientific community, industry and any other interested party on
features and areas that may meet the definition of critical habitat for
the giant manta ray within U.S. waters. We received information
regarding giant manta ray occurrence in the Flower Garden Banks
National Marine Sanctuary (Stewart et al. 2018b) as well as off the
coast of Florida. We reviewed this information and considered it along
with other available information we compiled. Together, this
information comprises the best available scientific data for use in the
identification of critical habitat for the giant manta ray. However, as
discussed below, based on these data we find that there are no
identifiable physical or biological features that are essential to the
conservation of the giant manta ray within areas under U.S.
jurisdiction, or unoccupied areas under U.S. jurisdiction that are
essential to the conservation of the species. Therefore, at this time
we find no areas within U.S. jurisdiction that meet the definition of
critical habitat for the giant manta ray.
This finding describes information on the biology, distribution,
and habitat use of the giant manta ray and the methods used to identify
areas that may meet the definition of critical habitat. In this
determination, we focus on information directly relevant to the
designation of critical habitat for giant manta rays.
Giant Manta Ray Biology and Status
The following discussion of the life history and status of giant
manta ray is based on the best scientific data available, including the
``Endangered Species Act Status Review Report: Giant Manta Ray (Manta
birostris) and Reef Manta Ray (Manta alfredi)'' (Miller and Klimovich
2017).
Manta rays are large bodied, planktivorous rays, considered part of
the Mobulidae subfamily. Manta species are distinguished from other
Mobula rays in that they tend to be larger, with a terminal mouth, and
have long cephalic fins (Evgeny 2010); however, misidentifications are
common both between Manta species (i.e., between M. alfredi and M.
birostris) as well as between Manta and Mobula rays. In addition,
recent taxonomic studies have suggested that Manta birostris and Manta
alfredi may actually be closely related to the giant devil ray (Mobula
mobular) (White et al. 2017), with genetic analyses that demonstrate
support for nesting these species under the genus Mobula rather than
Manta (White et al. 2017; Hosegood et al. 2019). The studies still
recognize both manta rays as distinct species, but refer to them as
Mobula birostris and Mobula alfredi.
The giant manta ray, M. birostris, can be found in all ocean
basins, while the reef manta ray, M. alfredi, is currently only
observed in the Indian Ocean and the western and south Pacific.
Additionally, we note that a third, putative manta ray species has been
identified (referred to here as M. cf. birostris), with its range
extending along the Atlantic coast, Gulf of Mexico, and Caribbean,
based on research conducted in the western Atlantic (A. Marshall, MMF,
pers. comm. to M. Miller, NMFS OPR, 2019). A manuscript identifying
this third species is expected in the near future; however, according
to Dr. Andrea Marshall, this newly identified manta species is highly
abundant off the U.S. east coast, with a large population also found
off the Yucat[aacute]n peninsula (A. Marshall, MMF, pers. comm. to M.
Miller, NMFS OPR, 2019). This new species looks very similar to M.
birostris, with only a few diagnostic features that could potentially
distinguish the two (mainly small morphological and meristic ones; A.
[[Page 66653]]
Marshall, MMF, pers. comm. to M. Miller, NMFS OPR, 2019). Without
genetic testing, species identification cannot be completely validated
(Hinojosa-Alvarez et al. 2016; Kashiwagi et al. 2017; Hosegood et al.
2019).
Therefore, for purposes of this critical habitat determination, we
will consider any records of manta rays in the Atlantic to be M.
birostris (even though an unknown proportion may comprise M. cf.
birostris) and will continue to recognize Manta birostris as a species
under the genus Manta.
The genus Manta has a complex taxonomic history due partially to
the difficulty of preserving such large specimens and also the
conflicting historical reports of taxonomic characteristics (Couturier
et al. 2012; Kitchen-Wheeler 2013). Prior to 2009, most manta rays were
categorized as Manta birostris, but Marshall et al. (2009) presented
new data that supported the splitting of the Manta genus into two
species: M. birostris and M. alfredi.
Both Manta species have diamond-shaped bodies with wing-like
pectoral fins; the distance over this wingspan is termed disc width
(DW). There are two distinct color types in both species: Chevron and
black (melanistic). Most of the chevron variants have a black dorsal
surface and a white ventral surface with distinct patterns on the
underside that can be used to identify individuals (Marshall et al.
2008; Kitchen-Wheeler 2010; Deakos et al. 2011). The black color
variants of both species are entirely black on the dorsal side and
almost completely black on the ventral side, except for areas between
the gill-slits and the abdominal area below the gill-slits (Kitchen-
Wheeler 2013).
Giant manta rays inhabit tropical, subtropical, and temperate
bodies of water and are commonly found offshore, in oceanic waters, and
near productive coastlines. It is thought to be a generally long-lived
species (>28 years) (Stewart et al. 2018a) with low reproductive
output. Manta rays, like all chondrichthyans, reproduce via internal
fertilization (Wourms 1981), and the sexes can be differentiated by the
presence of myxopterigia, or claspers, on the inner margin of the
pelvic fins in males, whereas females lack these structures. Sexual
maturity in males can be easily determined by examining the level of
calcification in these intromittent organs. In their examination of
mobulids taken as bycatch in the Indonesian drift net fishery, White et
al. (2006) found that male M. birostris greater than 3,800 mm DW
possessed fully calcified claspers and were, therefore, mature, while
those less than 3,800 mm DW possessed either non-calcified or partially
calcified claspers. In the same study, White et al. (2006) found that
females 2,732 to 3,774 mm DW were immature and females measuring 4,126
mm DW and greater were mature. White and Last (2016) report similar
ranges, with males maturing between approximately 3,750 and 4,000 mm DW
and females maturing between approximately 4,100 and 4,700 mm DW. In
the Flower Gardens Banks National Marine Sanctuary (FGBNMS), Stewart et
al. (2018b) observed a mature male M. birostris with an estimated size
of 3,600 mm. The age that M. birostris matures is not known, but it may
be similar to that of reef mantas, with males maturing at 3-6 years and
females at 8-10 years (Stewart et al. 2018a).
Gestation time is also not known for this species, and parturition
has only been witnessed once and under unnatural conditions (Coles
1916). It is suspected that gestation would be similar to that observed
in M. alfredi, which is generally accepted to be 12 to 13 months
(Kitchen-Wheeler 2013). In addition to the Coles (1916) observation of
a single embryo aborted during capture, the limited investigations of
pregnant females with embryos intact have all indicated the presence of
a single embryo per pregnancy (Muller and Henle 1838-1841; Beebe and
Tee-Van 1941). Similarly, reports of reef manta ray births and
dissections have also all revealed only a single embryo (Homma et al.
1999; Uchida et al. 2008). Size at birth has remained elusive for M.
birostris. The embryos examined in the previous studies had sizes of
1,140 mm and 1,270 mm DW (Muller and Henle 1838-1841; Beebe and Tee-Van
1941), while the smallest free swimming individuals reported by Stewart
et al. (2018b) were approximately 1,000 mm DW (however, these
individuals may have been M. cf. birostris). Rambahiniarison et al.
(2018) recently estimated size at birth of M. birostris to be 2,000 mm
DW based on the DW of the largest fetus and the smallest free-living
specimen captured in the Philippines mobulid fishery.
Very little is known about the early life stages or habitat needs
or requirements of M. birostris because, until fairly recently,
juveniles have rarely been observed in the wild. However, large numbers
of juvenile M. birostris have been caught in Sri Lanka in offshore
pelagic habitats by the gill-net fisheries, landed by fisherman in
Brazil and Indonesia, and also observed in oceanic habitats off Mexico
(Stewart et al. 2016a; Stewart et al. 2018b). Stewart et al. (2016a)
suggests that adult and juvenile giant mantas may use similar offshore
pelagic habitats, but that the juveniles may avoid cleaning stations
and other near-shore areas where adults are more commonly observed to
reduce predation risk. In fact, results from stable isotope analyses of
muscle tissues collected from both adult and juvenile M. birostris off
Peru, Sri Lanka, and the Philippines appear to provide further
confirmation that the species may not undergo an ontogenetic shift in
feeding behavior or trophic level, with both adults and juveniles
sharing the same habitats and targeting the same prey (Stewart et al.
2017).
In terms of prey, giant manta rays primarily feed on planktonic
organisms such as euphausiids, copepods, mysids, decapod larvae, and
shrimp, with some studies noting their consumption of small and
moderate sized fishes as well (Bigelow and Schroeder 1953; Carpenter
and Niem 2001; Graham et al. 2012; Stewart et al. 2016b; Burgess 2017;
Rohner et al. 2017). They feed by swimming with their mouths open,
continuously filtering zooplankton. Their gill rakers filter out water,
leaving behind food particles that are then directed to the esophagus
through cross-flow (Paig-Tran 2012). This filter mechanism allows
mantas to retain prey of various sizes, even if they are smaller than
the filter pores, which means they can effectively feed on mixed
zooplankton assemblages where prey range in size from small calanoid
copepods to larger mysids and euphausiids (Stewart et al. 2016b). Given
the feeding habits of the giant manta ray, it can be considered a
generalist carnivore, with a trophic position of approximately 3.4
(Burgess et al. 2016; Burgess 2017).
With regards to movement, the giant manta ray is considered to be a
migratory species, with satellite tracking studies measuring straight
line distances of up to 1,500 km (Hearn et al. 2014). Some giant manta
rays appear to migrate seasonally, possibly due to the seasonal
fluctuations in food sources (Wilson et al. 2001; Luiz et al. 2009;
Graham et al. 2012; Sobral and Afonso 2014; De Boer et al. 2015;
Girondot et al. 2015; Stewart et al. 2016a; Hacohen-Domen[eacute] et
al. 2017). However, in some portions of its range, the species may
actually exist as well-structured subpopulations with a high degree of
residency (Stewart et al. 2016a).
As discussed in the proposed rule (82 FR 3694, January 12, 2017)
and final rule (83 FR 2916, January 22, 2018) to list the giant manta
ray, the most significant threat to the species is overutilization for
commercial purposes. Giant manta rays are both
[[Page 66654]]
targeted and caught as bycatch in a number of fisheries throughout
their range, and are most susceptible to industrial purse-seine and
artisanal gillnet fisheries. With the expansion of the international
mobulid gill raker market and increasing demand for manta ray products,
estimated take of giant manta rays, particularly in many portions of
the Indo-Pacific, frequently exceeds numbers of identified individuals
in those areas. Observations from these areas also indicate declines in
sightings and landings of the species. Given the extremely low
reproductive output and overall productivity of the giant manta ray, it
is inherently vulnerable to threats that would deplete its abundance,
with a low likelihood of recovery. So, while there is considerable
uncertainty regarding the current abundance of M. birostris throughout
its entire range, the best available information indicates that the
species is likely to become an endangered species within the
foreseeable future throughout a significant portion of its range (the
Indo-Pacific and eastern Pacific portion) due to overutilization.
Critical Habitat Identification and Designation
Critical habitat is defined by section 3 of the ESA as: ``(i) the
specific areas within the geographical area occupied by the species, at
the time it is listed . . . , on which are found those physical or
biological features (I) essential to the conservation of the species
and (II) which may require special management considerations or
protection; and (ii) specific areas outside the geographical area
occupied by the species at the time it is listed . . . upon a
determination by the Secretary that such areas are essential for the
conservation of the species.'' This definition provides a step-wise
approach to identifying areas that may qualify as critical habitat for
the giant manta ray: (1) Determine the geographical area occupied by
the species at the time of listing; (2) identify physical or biological
habitat features essential to the conservation of the species; (3)
delineate specific areas within the geographical area occupied by the
species on which are found the physical or biological features; (4)
determine whether the features in a specific area may require special
management considerations or protection; and (5) determine whether any
unoccupied areas are essential for conservation. Our evaluation and
conclusions as we worked through this step-wise process are described
in detail in the following sections.
Geographical Area Occupied by the Species
The ``geographical area occupied by the species'' is defined in our
regulations as ``an area that may generally be delineated around
species' occurrences, as determined by the Secretary (i.e., range).
Such areas may include those areas used throughout all or part of the
species' life cycle, even if not used on a regular basis (e.g.,
migratory corridors, seasonal habitats, and habitats used periodically,
but not solely by vagrant individuals).'' (50 CFR 424.02). Further, our
regulations at 50 CFR 424.12(g) state: ``The Secretary will not
designate critical habitat within foreign countries or in other areas
outside of the jurisdiction of the United States.'' As such, we focus
the following discussion on the range of the species within waters
under U.S. jurisdiction.
In the Atlantic, giant manta rays have been confirmed as far north
as Long Island, New York (offshore around the Hudson Canyon region)
(Normandeau Associates and APEM Ltd 2017); however, as will be
discussed later, we note that they are generally rare north of Cape
Hatteras, North Carolina. To the south, giant manta rays occur off the
coast of North Carolina, South Carolina, Georgia, and Florida (Marshall
et al. 2011). Giant manta rays can also be found throughout the U.S.
Gulf of Mexico and within the U.S. Caribbean, including off Puerto Rico
and the U.S. Virgin Islands (Marshall et al. 2011). In the central
Pacific, giant manta rays are found off Hawaii (Clark 2010) and Jarvis
Island (K. Lino unpublished data). While there have been no confirmed
sightings of giant manta rays in waters of the other Pacific Remote
Island Areas, Northern Mariana Islands (Kashiwagi et al. 2011), Guam
(Kashiwagi et al. 2011), or American Samoa, based on confirmed
observations of the species elsewhere throughout the Pacific (e.g.,
Ogasawara Islands, Japan (Kashiwagi et al. 2010); Philippines (Verdote
and Ponzo 2014); French Polynesia (Mourier 2012); Jarvis Island (K.
Lino unpublished data); Hawaii (Clark 2010)) and coupled with the
migratory and pelagic nature of giant manta rays, their ability to
exploit significant depths, and tolerance of tropical to temperate
water temperatures, we find no known barriers to their movement that
may prevent them from occurring at these locations.
In the eastern U.S. Pacific, while there is documentation of a
giant manta off the west coast (i.e., San Clemente Island, California),
this sighting was of a single individual in 2014 (Warneke 2014) and
there have been no documented sightings since (or prior to) this time.
Given the amount of fishing effort, as well as the human population
density in these regions, it is highly unlikely that substantial
concentrations of giant manta rays would have passed unnoticed. As
such, we consider this individual to be a vagrant of the species (an
individual that occurs outside of the species' normal range).
Therefore, as the occurrence of giant manta rays in waters off the U.S.
west coast is extremely uncommon, we do not consider this geographical
area to be part of the species' occupied range at the time of listing.
Conclusion
Based on the above information and analysis, we define the
geographical area occupied by the giant manta ray at the time of
listing as all U.S. waters off the east coast, from Florida to Long
Island, New York, the entire Gulf coast, the U.S. Virgin Islands and
Puerto Rico in the Caribbean, and Hawaii, the Pacific Remote Islands
Areas, Guam, American Samoa, and the Northern Mariana Islands in the
Pacific.
Physical or Biological Features Essential for Conservation
Within the geographical area occupied by an endangered or
threatened species at the time of listing, critical habitat consists of
specific areas upon which are found those physical or biological
features essential to the conservation of the species and that may
require special management considerations or protection. The ESA does
not specifically define physical or biological features; however, court
decisions and joint NMFS-USFWS regulations at 50 CFR 424.02 provide
guidance on how physical or biological features are expressed.
Specifically, these regulations state that the physical and biological
features are those that are essential to support the life-history needs
of the species, including but not limited to, water characteristics,
soil type, geological features, sites, prey, vegetation, symbiotic
species, or other features. A feature may be a single habitat
characteristic, or a more complex combination of habitat
characteristics. Features may include habitat characteristics that
support ephemeral or dynamic habitat conditions. Features may also be
expressed in terms relating to principles of conservation biology, such
as patch size, distribution distances, and connectivity. (50 CFR
424.02).
Section 3 of the ESA (16 U.S.C. 1532(3)) defines the terms
``conserve,'' ``conserving,'' and ``conservation'' to mean: ``to use
and the use of all
[[Page 66655]]
methods and procedures which are necessary to bring any endangered
species or threatened species to the point at which the measures
provided pursuant to this chapter are no longer necessary.'' For giant
manta rays, we consider conservation to include the use of all methods
and procedures necessary to bring giant manta rays to the point at
which factors related to population ecology and vital rates indicate
that the species is recovered in accordance with the definition of
recovery in 50 CFR 402.02. Important factors related to population
ecology and vital rates include population size and trends, range,
distribution, age structure, gender ratios, age-specific survival, age-
specific reproduction, and lifetime reproductive success. Based on the
available knowledge of giant manta ray population ecology and life
history, we have identified four biological behaviors that are critical
to the goal of increasing survival and population growth: (1) Foraging,
(2) pupping, (3) migration, and (4) breeding. In the following section,
we evaluate whether there are physical and biological features of the
habitat areas known or thought to be used for these behaviors that are
essential to the species' conservation because they facilitate or are
intimately tied to these behaviors and, hence, support the life-history
needs of the species. Because these behaviors are essential to the
species' conservation, facilitating or protecting each one is
considered a key conservation objective for any critical habitat
designation for this species.
Analysis of the Physical and Biological Features of Foraging Habitat
That Are Essential to the Conservation of the Species
Giant manta rays are filter-feeders and generalist carnivores that
feed on a variety of planktonic organisms, including euphausiids,
copepods, mysids, decapod larvae and shrimp, as well as small fishes.
Prey needs to be of sufficient density and quality to support the
energy requirements for the giant manta rays, particularly as they
conduct long-distance migrations across open oceans. Sustained
decreases in prey quantity, quality, availability, or accessibility can
decrease foraging success of giant manta rays and eventually lead to
reduced individual growth, reproduction, and development. Therefore,
using the best available data, we examined the diet and energy needs of
giant manta rays, including foraging behavior, to determine whether we
could identify physical or biological features of habitat that
facilitate successful giant manta ray feeding and, thus, are essential
for the conservation of the species.
As mentioned above, planktonic organisms comprise the majority of
the diet for giant manta rays. While it was previously assumed that
manta rays obtain most of their energy needs from surface zooplankton,
results from recent studies indicate that these feeding events may not
be the primary source of the dietary intake (Burgess et al. 2016;
Stewart et al. 2016b). For example, for giant manta rays off Ecuador,
Burgess et al. (2016) estimated that, on average, mesopelagic food
sources contribute 73 percent to the giant manta ray's diet compared to
27 percent for surface zooplankton. In the Mexican Pacific, Stewart et
al. (2016b) interpreted dive profiles and submersible video data of M.
birostris to suggest that giant manta rays frequently forage on
vertically migrating zooplankton and zooplankton in the epipelagic
scattering layers in addition to surface zooplankton.
Analysis of stomach contents and collection of zooplankton during
observed giant manta ray feeding events reveal a varied diet, with no
targeting of a specific species or size of prey (Graham et al. 2012;
Armstrong et al. 2016; Stewart et al. 2016b; Burgess 2017; Rohner et
al. 2017). Rather, density of the prey appears to be the driving factor
that triggers giant manta ray feeding behavior. However, the levels
necessary to attract giant manta rays remain unknown. For example, a
study conducted by Burgess (2017) found that giant manta ray
aggregations off the northwest side of Isla de la Plata, Ecuador, were
unlikely associated with foraging opportunities as observations of
feeding events were rare. Specifically, Burgess (2017) collected
surface zooplankton during feeding events (n=5) and during non-feeding
events (n=79) and calculated that the dry zooplankton biomass was 1.9
mg m-3 during the rare M. birostris feeding events and 1.4
mg m-3 during non-feeding events. Although comparable data
are unavailable for M. birostris elsewhere throughout its range, these
figures are substantially lower than what has been reported for the
closely related reef manta ray, M. alfredi, in eastern Australia during
regular active feeding (19.1 mg m-3) and non-feeding (9.3 mg
m-3) events (Armstrong et al. 2016). In fact, Armstrong et
al. (2016) determined that the critical prey density threshold for M.
alfredi feeding was 11.2 mg m\-3\. If M. birostris has similar prey
density thresholds, these data lend support to Burgess (2017)'s finding
that the aggregative behavior of giant manta rays at Isla de la Plata
is unlikely related to feeding. Furthermore, the data suggest that for
habitat to be characterized as providing necessary foraging
opportunities, it likely requires substantially higher levels of
zooplankton biomass than what was found off Isla de la Plata.
In terms of energy needs, the only available data that provides
insight for M. birostris is from a study that examined the stomach
contents of giant manta rays collected within the Bohol Sea
(Philippines) in 2015 (Rohner et al. 2017). Using adiabatic bomb
calorimetry, Rohner et al. (2017) calculated that krill (Euphausia
diomedeae), the dominant prey species for M. birostris in this
particular area, contributed 24,572 kJ (20,451 kJ s.d.) per
100 g of stomach content in M. birostris. When scaled up based on the
total number of euphausiids per stomach, the authors estimated that E.
diomedeae contributed up to 631,167 kcal in the giant manta ray diet
(Rohner et al. 2017). This energetic contribution is significantly
greater than what has been found for reef manta rays in captivity.
Rohner et al. (2017), citing a personal communication, reports that in
aquaria, a 350 cm DW M. alfredi is fed 3,500 kcal per day and a 450 cm
DW M. alfredi is fed 6,100 kcal per day, with captive reef manta rays
consuming 12.7 percent of their body weight in euphausiids weekly
(Homma et al. 1999). Although energy requirements and caloric intake
for captive manta rays will likely be different than those found in the
wild, Rohner et al. (2017) proposes that the significant calorific
value of the M. birostris stomach contents suggests that giant manta
rays partake in numerous feeding events over several days or,
alternatively, engage in a few, sporadic, opportunistic feeding events
on large aggregations of prey that can be used to sustain them until
their next meal. Burgess (2017) tends to agree with the latter. The
author cites the particularly large capacity of the M. birostris
stomach, as well as the branchial filter pad and filtration mechanism
used by manta rays (which allows for the capture of numerous
macroscopic zooplankton and small fishes of varying sizes) to support
the assumption that manta rays likely exploit large patches of
zooplankton for a high net energy gain in a short period of time
(Burgess 2017). However, with only one study that has examined the
energy contents of a particular prey item of M. birostris in a specific
area, it is difficult to make any conclusions as to the general energy
needs or requirements for the species throughout its range.
With the lack of available data regarding prey density thresholds
or
[[Page 66656]]
caloric value requirements, we next looked at areas where manta rays
have been observed or assumed to be feeding to determine whether we
could identify any physical or biological features of these habitats
that are tied to foraging behavior. In many portions of the species'
range, it is the presence of seasonal upwelling events, which
concentrate plankton and create patches of high productivity, that
appear to drive the occurrence of giant manta rays in areas, presumably
for foraging. For example, off the northern Yucat[aacute]n peninsula,
Hacohen-Domen[eacute] et al. (2017) found a higher probability of M.
birostris occurrence from July through September, with the main
difference being the increase in primary productivity during this time
of year (with particularly high probability of occurrence when primary
productivity was at 4,500 mg
C[middot]m-2[middot]day-1). Other features
associated with a greater probability of giant manta ray presence in
this area included sea surface temperatures (SST) warmer than 27
[deg]C, shallow (<10 m depths) and nearshore waters (<50 km from
shore), with a bottom slope of <0.5[deg] (Hacohen-Domen[eacute] et al.
2017). However, the authors note that most of the manta rays observed
in the study were not foraging but rather swimming alone or in pairs.
While Hacohen-Domen[eacute] et al. (2017) did not observe or analyze
feeding habits in their study, Hinojosa-Alvarez et al. (2016) confirmed
foraging behavior in this area (specifically between 21[deg]46.020' N
and 87[deg]01.200' W and 21[deg]30.00' and 86[deg]4100), with videos of
Yucat[aacute]n manta rays feeding in surface waters from May through
August (the same period as the seasonal upwelling).
Seasonal occurrence of manta rays was also observed off the
continental shelf of French Guiana. Specifically, Girondot et al.
(2015) observed a peak in the presence of manta rays between July and
December in the river-ocean transition zone off French Guiana. While
specific features of the habitat where giant manta rays were observed
was not provided, the authors did note that phytoplankton biomass and
primary productivity is generally highest during the months of manta
ray presence, with a biomass of over 25 mg Chl-a m-3 and
productivity of over 8 g C[deg]m-2*day-1
(Girondot et al. 2015).
Similarly, in southeastern Brazil, giant manta rays are most
frequently sighted in Laje de Santos Marine State Park (24[deg] S)
during seasonal upwelling, from June to August (Luiz et al. 2009).
During this time, the warm Brazil Current weakens and coastal waters
change direction and move northward, bringing waters from the southern
Falklands Current to areas of southeastern Brazil (Luiz et al. 2009).
This current displaces a low salinity front (generated by discharge
from the La Plata River) from the mouth of the La Plata River during
the summer to areas north in the winter (Luiz et al. 2009). It is
thought that this coastal front, which accumulates plankton, may
attract giant manta rays at Laje de Santos Marine State Park in the
winter months (Luiz et al. 2009). However, besides the greater presence
of manta rays in this region during the seasonal upwelling event (based
on diver photos), no information was provided regarding foraging
activities or the essential physical or biological features of the
habitat that are necessary to support this behavior.
Off the coast of Suriname, De Boer et al. (2015) found that the
presence of M. birostris coincided with the region's two rainy seasons.
As the outflows of nutrient-rich waters from the Amazon and Suriname
rivers lead to a low salinity front during the rainy seasons, the
authors suggest that giant manta rays are visiting the coastal waters
of Suriname for feeding purposes (De Boer et al. 2015). Although only a
few observations of manta rays were recorded during the survey period,
the authors found the behavior was likely indicative of foraging (i.e.,
swimming just below the surface with pectoral fins curled) (De Boer et
al. 2015); however, again, no physical or biological features of the
foraging habitat were identified.
While upwelling events appear to be the main environmental factor
driving manta ray foraging behavior, we note that Graham et al. (2012)
also observed a giant manta ray feeding in oligotrophic waters during a
seasonal fish spawning event. The giant manta ray was initially tagged
off the northern Yucat[aacute]n peninsula in eutrophic waters and
observed feeding on copepeds (Graham et al. 2012). However, 57 days
later, it was re-sighted in oligotrophic waters foraging on fish eggs
released during a seasonal spawning event of little tunny (Euthynnus
alletteratus), suggesting that giant manta rays are also able to
exploit different habitats when conditions arise that are suitable for
foraging (Graham et al. 2012).
Overall, based on the foregoing information regarding known or
presumed foraging areas for giant manta rays, the general and
consistent physical oceanographic feature that appear to be associated
with foraging habitat is high primary productivity from upwelling
events, which favors the potential accumulation of zooplankton. Yet the
levels of primary productivity necessary to produce suitable foraging
habitat are unknown, and this feature is relatively ubiquitous
throughout the global range of the species, with not all areas of high
primary productivity providing meaningful foraging habitat for giant
manta rays. Furthermore, given that the characteristics of habitat
necessary to produce areas of high primary productivity varies by
region and site (e.g., seasonal upwelling events due to increased river
discharge or wind-driven fronts), we proceeded to focus our examination
on whether we could identify any physical and biological features of
giant manta ray foraging areas within U.S. waters that are essential to
the conservation of the species.
In general, very little published literature exists on giant manta
ray occurrence and behavior in U.S. waters. Adams and Amesbury (1998)
documented the presence of three giant manta rays in the estuarine
waters of the Indian River Lagoon system and in Port Canaveral,
Florida. Foraging behavior was not observed and the authors proposed
that individuals likely enter the estuary sporadically and stay for
only short durations. Freedman and Roy (2012) used Ocean Biogeographic
Information System (OBIS) data on giant manta ray observations to
examine the spatial distribution of the species along the U.S. east
coast. They found a higher number of observations near the continental
shelf edge and bordering the Gulf Stream, and suggested a seasonal
distribution of the species driven mainly by temperature, with giant
manta rays primarily observed in waters from 19 [deg]C to 22 [deg]C
(Freedman and Roy 2012). Manta rays are also known to visit the east
coast of Florida, more often in the spring and summer months, moving
north as water temperatures rise above 20 [deg]C (Levesque 2019).
However, while it is known that giant manta rays prefer warmer waters,
there is no evidence that this is a physical or biological feature that
is essential to the conservation of the species or related to foraging
activity. In fact, as noted in the literature, giant manta rays can be
found in waters anywhere from 18 [deg]C to 30 [deg]C (Yano et al. 1999;
Freedman and Roy 2012; Graham et al. 2012; Burgess 2017; Hacohen-
Domen[eacute] et al. 2017). Additionally, the OBIS data, upon which
Freedman and Roy (2012) based their conclusions, also has inherent
flaws as it is an open-access database where any member can submit
observations of marine species without validation. As will be discussed
below, there are significant misidentification issues associated with
M. birostris observations and conclusions drawn from this type of
sightings data should
[[Page 66657]]
be made with caution as there are significant uncertainties and
limitations to the data.
In the FGBNMS, Stewart et al. (2018b) documented high numbers of
giant manta rays but specifically noted that foraging behavior was
rare. Citing a personal observation (E. Hickerson), Stewart et al.
(2018b) stated that mantas were only rarely seen exhibiting barrel
rolling behavior (3 of 88 observations), indicative of feeding, at the
banks. In his study of the Flower Garden Banks and surrounding banks,
Childs (2001) documented M. birostris feeding behavior in February and
March of 2000 through the use of a remotely operated vehicle. He noted
that M. birostris generally fed along escarpments and within the water
column over the reef crest; however, no other details were provided
regarding these events.
In our own examination of the available data, we compiled manta ray
sightings data (NMFS unpublished data) from a number of available
surveys (Table 1), photo databases, individual observations, and social
media websites (e.g., YouTube and Facebook), and plotted the
information to assess whether we could determine ``hot spots'' of giant
manta rays, or areas where manta rays appear to be visiting
consistently over time. We initially made the main assumption that
sightings of the species were correlated with areas of high prey (as
tends to be the case with observations of giant manta rays in other
portions of its range). In other words, when a manta ray was spotted,
we assumed it was likely because that animal was foraging in the area,
but we also looked for behavioral (e.g., barrel rolling, mouth open,
cephalic lobes unfurled) or environmental data (e.g., high plankton
biomass) that could support this assumption as foraging may not be the
only reason for manta ray presence.
Because most manta sightings within surveys are opportunistic in
surveys designed for other species, there are some misidentification
issues and gaps in the time series. Many of the sightings data were
obtained from aerial surveys aimed at collecting information on the
distribution and abundance of marine mammals (for example, the Atlantic
Marine Assessment Program for Protected Species (AMAPPS) and North
Atlantic Right Whale Consortium data). This presents a problem as
observers on these surveys are usually not trained in identifying
mobulid rays to the species level. In discussions with biologist Todd
Pusser, a contract observer for NOAA in the southeast region during the
1990s and early 2000s who was then contracted through the NOAA
Northeast Fisheries Science Center (NEFSC) at Woods Hole and
participated in these marine mammal surveys from Canada to Cape
Hatteras, North Carolina, he confirmed that in both the NOAA aerial and
ship surveys along the Atlantic coast, mobulid sightings were simply
logged as ``manta ray'' or ``manta spp,'' thus greatly inflating the
sightings data for M. birostris (T. Pusser, pers. comm. to C. Jones,
NMFS SEFSC, 2018). In fact, when photos were available from
accompanying ship and aerial surveys, the majority of the sightings
logged as M. birostris in the northeast Atlantic were Mobula tarapacana
or M. mobular (T. Pusser, pers. comm. to C. Jones, NMFS SEFSC, 2018).
Table 1--Available Survey Datasets With Reported Manta Sightings
----------------------------------------------------------------------------------------------------------------
Survey name Year(s) Survey location
----------------------------------------------------------------------------------------------------------------
Digital Aerial Baseline Survey-- 2016, 2017...................................... Atlantic (38.45[deg] N
NYSERDA. to 41.08[deg] N).
AMAPPS (aerial)...................... 2010 through 2018............................... Atlantic (26.03[deg] N
to 45.32[deg] N).
North Atlantic Right Whale Consortium 1986 through 2017............................... Atlantic (25[deg] N to
database (various surveys). 41[deg] N).
SEFSC Mid-Atlantic Tursiops Survey 1994, 1995...................................... Atlantic (24.5[deg] N
(aerial). to 40.50[deg] N).
SEFSC Southeast Cetacean Aerial 1992, 1995...................................... Atlantic (26.21[deg] N
Survey. to 35.19[deg] N).
Florida Manta Project (boat & aerial; 2016, 2017, 2018................................ Atlantic (26.5[deg] N
directed manta ray survey). to 27[deg] N).
GA Aquarium (boat & aerial; directed 2010 through 2017............................... Atlantic (29.5[deg] N
manta ray survey). to 29.9[deg] N).
SEFSC Platform Calibration Survey 1991............................................ Atlantic (35.8[deg] N
(aerial). to 39.3[deg] N).
Gulf of Mexico Marine Mammal 2010, 2011, 2012................................ Gulf of Mexico (98[deg]
Assessment Aerial Surveys--NRDA. W to 80.5[deg] W).
GoMAPPS (aerial)..................... 2017, 2018...................................... Gulf of Mexico (97[deg]
W to 81[deg] W).
GulfCet (aerial)..................... 1992, 1993, 1994, 1996,1997..................... Gulf of Mexico
(96.5[deg] W to
84[deg] W).
SEFSC GoMex (aerial)................. 1992, 1993, 1994, 1996.......................... Gulf of Mexico
(96.3[deg] W to
82[deg] W).
NOAA Coral Reef Ecosystems Program 2006, 2010...................................... Pacific Islands
(towed diver survey). (160[deg] W; Jarvis
Island).
----------------------------------------------------------------------------------------------------------------
Note: Survey locations are given as geographic regions: Atlantic, Gulf of Mexico, Pacific Islands. For Atlantic
locations, the latitude range over which the surveys were conducted is given. For Gulf of Mexico and Pacific
Island locations, the longitude range over which the surveys were conducted is given.
We similarly found this to be the case with another available
dataset from the northeast Atlantic that documented 504 sightings of
``Giant Manta Ray'' (Normandeau Associates and APEM Ltd 2017). This
aerial survey, conducted in 2016 and 2017 and supported by the New York
State Energy Research and Development Authority (NYSERDA), encompassed
the waters of the New York Bight from Long Island southeast to the
continental shelf break. This dataset also had accompanying photos of
each animal observation, which a NMFS species expert was able to review
and confirm that only 6 of the 504 ``giant manta ray'' sightings were
actually Manta birostris (C. Horn, NMFS SERO, pers. comm. to M. Miller,
NMFS OPR, 2018). Similarly, in 2015, the NMFS Northeast Fisheries
Observer Program database underwent a species verification review
whereby NMFS scientists conducted a detailed review of observer photo
records with the assistance of manta and devil ray experts (i.e., Dr.
Giuseppe Notarbartolo di Sciara, Dr. Andrea Marshall, and Guy Stevens).
From 2009 to 2015, there were 25 manta and mobula species records with
photos in the database (J. Hare, memo, addressed to R.E. Crabtree,
February 1, 2019). Most of the mobula bycatch consisted of Mobula
tarapacana, with only two confirmed records of Manta birostris. These
individuals were observed caught off the coast of North Carolina. This
observer data appears to further confirm the rare occurrence of M.
birostris in the U.S. mid-Atlantic and northeast, and supports the
advice provided by species experts that all M. birostris sightings
north of Cape Hatteras should be questioned if there are no
corresponding photos.
There may also be occasional misidentifications of M. birostris
south
[[Page 66658]]
of Cape Hatteras as both Mobula tarapacana and M. mobular are also
common in this portion of the species' range within the Atlantic
(Stevens et al. 2018a, C. Jones unpublished data). Additionally, M.
tarapacana co-occurs with Manta birostris in the Gulf of Mexico and
Caribbean (Childs 2001), potentially confounding those aerial sighting
records as well. Thus, while the presence of M. birostris south of Cape
Hatteras is much more likely (based on photographic evidence), the
proportion of M. birostris in these datasets to the other two commonly
misidentified mobula rays is presently unknown, significantly
increasing the uncertainty of the accuracy of the available sightings
data.
In addition to misidentification rates, we found other inherent
problems with the sightings data during our analysis, including the
uncertainty regarding unique sightings and the large gaps in time
between surveys. For aerial surveys, planes are generally flown
following designated transect lines. Depending on the transect distance
and timing, there is potential for double-counting the same animal if
the animal is also moving. Without being able to view the ventral side
of the animal, it is difficult for aerial observers to identify whether
the manta ray they are spotting is the same individual from a previous
observation. Aerial surveys are also subject to availability bias
(i.e., the percentage of time a manta would be near enough to the
surface to be viewed by an aerial observer) and perception bias (i.e.,
the probability of an observer viewing the animal when it is
available). While it is possible to control for some of this
uncertainty using distance-weighted sampling techniques for perception
bias combined with data from satellite tags for availability bias, we
do not have the data or information that would be necessary in order to
conduct this type of analysis at this time, nor are we aware of any
available studies that have accounted for this uncertainty in reporting
and analyzing manta ray sightings.
Furthermore, as some of the aerial surveys were not regularly
conducted on an annual or seasonal basis, but rather for specific
research purposes that were unrelated to manta ray distribution or
abundance, the resulting data was skewed in terms of effort in specific
locations and over certain time periods and could not be used to
identify potential areas used routinely or repeatedly by giant manta
rays. For example, along the east coast, the SEFSC Mid Atlantic
Tursiops Surveys (MATS), for which we have manta ray sightings
information, were conducted in February of 1994 and July and August of
1995 to examine the distribution and estimate an index of relative
abundance for Atlantic bottlenose dolphins inhabiting nearshore coastal
waters in the mid and southern Atlantic bight. We also have data from
the SEFSC Southeast Cetacean Aerial Survey, SECAS, from February to
March in 1992 and March of 1995, a survey that was conducted to
estimate cetacean abundance. The Gulf of Mexico Marine Mammal
Assessment Aerial Surveys--Natural Resource Damage and Assessment
surveys were only conducted during the spring and summer of 2010 and
seasonally during 2011 to 2012 to assess the abundance and spatial
distribution of marine mammals and sea turtles within the region
impacted by the Deepwater Horizon oil spill. The Atlantic Marine
Assessment Program for Protected Species (AMAPPS), which conducted
annual aerial surveys from 2010-2017, had as its main objective
assessing the abundance, distribution, ecology, and behavior of marine
mammals, sea turtles, and seabirds throughout the U.S. Atlantic.
However, again, these surveys, as well as others that were analyzed
(see Table 1), varied with respect to the geographical coverage, years
and even months in which they were conducted. Currently there are no
available analyses of datasets or studies that control for spatial and
temporal variation in sampling effort, perception and availability
bias, and potential misidentification rates to distinguish areas of
high giant manta ray abundance.
Recently, we became aware of an ongoing dedicated manta ray aerial
survey, conducted by the Georgia Aquarium, which has documented manta
ray presence off the east coast of Florida since 2010. The manta aerial
surveys are conducted in spring and summer (March/April to June/July)
and follow general track lines 0 to 2.5 nautical miles (0 to 4.63 km)
from the beach that run parallel to the shore, from St. Augustine Beach
Pier (29[deg]52' N) to Flagler Beach Pier (29[deg]29' N). The number of
mantas are counted and, occasionally, dorsal photos of mantas are
collected during these surveys. However, due to the murkiness of the
water, photos are rather hard to obtain if the mantas are too deep in
the water column, and no ventral photos are available (H. Webb, GA
Aquarium, pers. comm. to M. Miller, NMFS OPR, 2019), preventing the
identification of individual manta rays or analysis of potential site
fidelity over the course of multiple years. Overall, the sightings data
indicate the seasonal visitation of manta rays to Florida's inshore
waters; however, the specific physical or biological features that
attract giant manta rays to this particular area are poorly understood.
The numbers, location, and peak timing of the manta rays to this area
varies by year, but with a notable decline in manta rays observed in
the study area since 2015 (H. Webb unpublished data). While sea surface
temperatures are thought to play a role in the initial migration of
manta rays to the study site, preliminary analysis suggests that the
within-season temperatures are not strongly correlated with manta ray
distribution or abundance within the area (H. Webb, GA Aquarium, pers.
comm. to M. Miller, NMFS OPR, 2019). Although foraging has been
anecdotally observed during these surveys (H. Webb, GA Aquarium, pers.
comm. to M. Miller, NMFS OPR, 2019) and mentioned in a few online
fishing articles (Roberts 2016; Levesque 2019), we are unaware of any
research that has determined the driving factor of manta ray occurrence
in this area and/or investigated the physical or biological features of
this area that may be essential to support the life history needs of
the species. Without information on specific habitat characteristics or
the relationship between environmental variables and manta ray
abundance or distribution, the available sightings data do not allow us
to identify important foraging areas at this time. A manuscript
summarizing findings from the Georgia Aquarium sightings dataset is
forthcoming (H. Webb, GA Aquarium, pers. comm. to M. Miller, NMFS OPR,
2019), and we intend to review any new information that becomes
available regarding manta ray use of this area off Florida.
Overall, the best available information indicates that giant manta
rays will feed on a variety of planktonic organisms and are not limited
by the required presence of a specific prey species for successful
foraging to occur. Areas of high primary productivity (e.g., upwelling)
are generally regarded as habitat that could potentially support giant
manta ray foraging events; however, the physical and biological
characteristics of high productivity areas can vary depending on the
location and season. Additionally, the presence of these areas does not
necessarily indicate giant manta ray foraging will occur as the
available data suggest some unknown prey density threshold may be
necessary to facilitate manta ray foraging or aggregations. In U.S.
waters, foraging has been anecdotally observed, but the available
[[Page 66659]]
data do not indicate any specific physical and biological features of
these areas that are essential for facilitating foraging events or
specific sites that are used consistently for foraging purposes. For
the foregoing reasons, it is not possible to identify any physical or
biological features related to foraging that are essential to the
conservation of the species, nor any specific areas that are essential
to support the foraging needs of the species within waters under U.S.
jurisdiction.
Analysis of the Physical and Biological Features of Pupping Habitat
That Are Essential to the Conservation of the Species
Giant manta rays likely give birth to only one pup per pregnancy
after a long gestation time (12-13 months). This very low reproductive
output for the species means that the success of pupping events is
essential for the conservation of the species. Identifying and
protecting important pupping habitat throughout the species' range will
be necessary to support recruitment of young individuals to the
recovering population. Without sufficient nursery habitat, the
population is unlikely to increase to a level associated with low
extinction risk and delisting. Protection of the species' nurseries is
crucial because the rebuilding of the population cannot occur without
protecting the source (juvenile) population and its associated
habitats. Therefore, using the best available data, we attempted to
identify potential nursery habitats and determine whether we could
identify physical or biological features of the habitat that facilitate
successful giant manta ray pupping and, thus, are essential for the
conservation of the species.
For the purposes of identifying potential nursery habitat, we
considered giant manta rays that were less than 4,000 mm DW to be
immature, with a size at birth of ~2,000 mm DW. As mentioned
previously, juvenile giant manta rays are rarely observed in the wild
but are present in the fishery landings data from many countries,
including Sri Lanka, Brazil, Indonesia, and the Philippines. While this
indicates that fishermen are accessing potentially important juvenile
habitat and possibly nursery areas, we have no data on these fishing
grounds that could provide insight into important physical or
biological features of these areas. However, recent manta ray research
in U.S. waters has documented the presence of juvenile giant manta rays
in the FGBNMS in the U.S. Gulf of Mexico as well as off the east coast
of Florida, suggesting the existence of juvenile and potential manta
ray nursery habitat, which we discuss below.
For the FGBNMS, both Childs (2001) and Stewart et al. (2018b)
suggested this area may contain potential nursery grounds for the
species. Although juveniles are rarely observed globally, a high number
of juveniles were sighted at several locations in the FGBNMS over
multiple years. Based on an analysis of NOAA diver logs (from various
coral reef and fish surveys), approximately 171 individual manta rays
have been sighted within the FGBNMS since 1994 (C. Jones unpublished
data). Of these, 114 have approximate recorded sizes. Around 97 percent
of the individuals sighted were less than 4 m DW (i.e., immature), and
around 50 percent were 2 m DW (i.e., estimated size at birth of M.
birostris) or less. However, M. cf. birostris may comprise the majority
of these sightings as Stewart et al. (2018b) noted that at least 55
percent of the manta rays identified in their study likely belong to M.
cf. birostris, which is thought to be closer in size to M. alfredi
(Stevens et al. 2018a) and potentially explains the observations of
mantas with sizes smaller than the estimated size at birth for M.
birostris.
Using the nursery habitat criteria proposed by Heupel et al.
(2007), Stewart et al. (2018b) suggested that the FGBNMS may contain
nursery habitat for giant manta rays because juveniles, which are
generally rare, are found in this area, remain in the area for a period
of several days to months, and have been sighted with gaps of more than
a year between re-sightings. The FGBNMS is a unique area, situated over
100 miles offshore of the Texas/Louisiana border and comprised of
shallow, underwater features, called salt domes, upon which diverse
coral reef communities have developed and thrived. There is substantial
upwelling, distinct thermoclines, and unique eddies that form in the
area, presumably due to interactions between currents and the
pronounced benthic features. Stewart et al. (2018b) proposed that the
FGBNMS may be an optimal nursery ground because it contains habitat
near the edge of the continental shelf and in proximity to abundant
pelagic food resources. Important prey for manta rays, like
euphausiids, are abundant in the deep scattering layers in the basin
waters of the Gulf of Mexico (Stewart et al. 2018b). The authors state
that an additional benefit of the FGBNMS is that the shallow bottom
habitat may protect juvenile rays from predation while they rest and
recover their body temperature in the warm mixed layer after deep
foraging dives (Stewart et al. 2018b).
However, while the FGBNMS provides habitat for juvenile giant manta
rays, the available data do not indicate any specific physical and
biological features within the FGBNMS that are essential for supporting
pupping behavior or necessary for a manta ray nursery. For example, in
examining specific physical features, like temperature, we found that
the majority of individuals (~75 percent) at the FGBNMS were sighted
between July and September (Stewart et al. 2018b). Sea surface
temperatures during these sightings ranged from 20 [deg]C to 32 [deg]C,
with ~75 percent of mantas observed in 28 [deg]C to 31 [deg]C (C. Jones
unpublished data). However, dives during which observations were
collected were skewed towards summer months (i.e., warmer temperatures)
and specific sites and depths (limited to areas above 150 ft (45.7 m)),
meaning that the increased observations of giant manta rays in the
higher temperature range may be a consequence of the survey methodology
and not a reflection of an essential feature of the habitat.
Next, we reviewed the available data regarding behavior to see if
we could identify specific habitat features based on use of the habitat
that are necessary to support pupping. As stated in Stewart et al.
(2018b) and Childs (2001), the primary behavior of manta rays observed
in the FGBNMS was mainly swimming, with manta rays swimming above reef
crest and sand flats, along escarpments, and in the water column.
Although more juveniles were sighted at East and West Flower Garden
Banks (hermatypic coral habitat) than at Stetson Bank (silt/claystone
dominated coral community), acoustic telemetry tagging has shown that
juvenile mantas move between East, West, Stetson, and Bright Bank
within FGBNMS (R. Graham, Wildlife Conservation Society, pers. comm. to
C. Horn, NMFS SERO, 2018). Stewart et al. (2018b) suggest the FGBNMS
likely provides ample feeding opportunities for juveniles, but they
acknowledge that foraging behavior is only rarely observed. Similarly,
Childs (2001) mentioned that foraging behavior at the FGBNMS was
observed in only two months (February and March) of his study despite
manta rays occurring in the area during all months.
While the presence of young giant manta rays suggest potential
pupping in the vicinity of the area (Childs 2001), the available data
do not allow us to identify where this pupping is occurring.
Additionally, the available data do not explain why or how giant manta
rays are using this particular habitat (e.g., foraging, transiting,
resting) or allow us to identify the essential
[[Page 66660]]
physical or biological features of the habitat. Therefore, we cannot
identify any pupping areas that meet the definition of critical
habitat.
Research (supported by NMFS and the National Ocean Service, in
collaboration with the Manta Trust) on the movements and genetics of
giant manta rays continues in the FGBNMS and may help provide answers
to these questions in the future. However, at this time, the available
data do not indicate any physical or biological features of this
habitat that are essential for the conservation of the species.
Similar to the FGBNMS, juvenile M. birostris have also been
regularly observed off the east coast of Florida in the past several
years. Since 2016, researchers with the Marine Megafauna Foundation
have been conducting annual surveys along a small transect off Palm
Beach, Florida, between Jupiter Inlet and Boynton Beach Inlet (~44 km,
24 nautical miles) (J. Pate, MMF, pers. comm. to M. Miller, NMFS OPR,
2018). Results from these surveys indicate that juvenile manta rays are
present in these waters for the majority of the year (observations span
from May to December), with re-sightings data that suggest some manta
rays may remain in the area for extended periods of time or return in
subsequent years (J. Pate unpublished data). For example, one satellite
tagged male has been re-sighted multiple times in the past 3 years
(Marine Megafauna Foundation 2019). However, similar to the limitations
of the FGBNMS data and the level of resolution, it is currently unclear
what physical or biological characteristics of this habitat are
necessary to facilitate successful pupping behavior or are essential
for nursery habitat. Manta rays are difficult to detect using boat-
based observation. When an observer spotted a manta ray, he/she would
get into the water and collect habitat information, behavioral data, as
well as photos of the manta ray. This type of data collection has
limitations. For example, water turbidity, depth, and weather
conditions may make manta rays harder to spot from a boat. As such, the
fact that the majority of manta rays were spotted over sand is likely
due to increased visibility over this type of habitat compared to
others (such as reef habitat) (J. Pate, personal communication, 2018)
as opposed to a biological necessity for this type of habitat.
Additionally, the main behavior observed in the transect area was
swimming, with occasional observations of foraging behavior near
Jupiter Inlet (J. Pate, MMF, pers. comm. to M. Miller, NMFS OPR, 2018).
In other words, similar to the FGBNMS, the available data only indicate
juvenile manta ray presence in these areas and does not explain why or
how giant manta rays are using the particular habitat that would help
us identify any physical or biological features that are essential for
the conservation of the species. We also note that the majority, if not
all, of these juvenile manta rays observed off the east coast of
Florida are thought to be M. cf. birostris (J. Pate, MMF pers. comm. to
M. Miller, NMFS OPR, 2018) and not M. birostris. NMFS researchers are
currently collaborating with colleagues at the Marine Megafauna
Foundation to tag these manta rays off the Florida coast and collect
genetic information in order to inform taxonomy, determine population
structure, and learn more about their movements to gain a better
understanding of their habitat use in this region. Anecdotal
observations from some of these recent tagging trips (June and August
2019) suggest this area may provide foraging opportunities (N. Farmer,
NMFS SERO, pers. comm. to M. Miller, NMFS OPR, 2019); however, further
investigation is required as the available information does not
indicate any specific physical and biological features of this area
that are essential to support the life-history needs of the species.
We also obtained anecdotal observations of juvenile giant manta
rays in the U.S. Caribbean from off Puerto Rico (n=10; sightings dating
back to 2004) and the U.S. Virgin Islands (n=16; sightings dating back
to 2012), and in the U.S. Pacific from off Hawaii and the Pacific
Remote Island Areas (n=24; sightings dating back to 2003) that indicate
the use of these waters by young giant manta rays (NMFS unpublished
data). However, as stated before, simply the observation of the
presence of juveniles using these waters (and further confounded by a
lack of known abundance, duration, movement, or frequency of occurrence
in these areas) is not enough information to indicate that these areas
contain physical and biological features that are essential to the
conservation of the species.
In summary, while we have evidence of the presence and use of
specific areas by juvenile giant manta rays, the available information
does not allow us to identify any physical or biological features
within these areas that are essential to support the life-history needs
of the species. Without knowledge of the essential features that create
meaningful pupping and nursery grounds, we cannot identify any areas
that meet the definition of critical habitat at this time.
The Physical and Biological Features of Migratory Habitat That Are
Essential to the Conservation of the Species
Based on the available data, it is evident that both small and
large-scale migratory movements are a necessary component in the life-
history of the giant manta ray. Seasonal sightings data suggests that
large-scale movements are undertaken primarily for foraging purposes,
correlated with the movement of zooplankton and influenced by current
circulation and tidal patterns, seasonal upwelling, and seawater
temperature (Luiz et al. 2009; Couturier et al. 2012; Freedman and Roy
2012; Graham et al. 2012; Sobral and Afonso 2014; De Boer et al. 2015;
Girondot et al. 2015; Armstrong et al. 2016; Hacohen-Domen[eacute] et
al. 2017). Small-scale movements also appear to be associated with
exploiting local prey patches in addition to refuging and cleaning
activities (O'Shea et al. 2010; Marshall et al. 2011; Graham et al.
2012; Rohner et al. 2013; Stewart et al. 2016a; Stewart et al. 2016b;
Sotelo 2018). However, as sightings of giant manta rays tend to be
sporadic, with the species more commonly found offshore and in oceanic
waters, it is difficult to track small-scale and large-scale migratory
behavior of the species. For logistical reasons, survey effort tends to
be focused in nearshore habitats. Yet, through the opportunistic
tagging of giant manta rays with pop-up satellite archival tags when in
these nearshore areas, researchers have been able to provide evidence
of the migratory nature of giant manta rays and demonstrate the
species' ability to make large-scale migrations. For example, satellite
tracking has registered movements of the giant manta ray from
Mozambique to South Africa (a distance of 1,100 km), around Ecuador and
its islands (between the Isla de la Plata, Bajo Cope, and Isla Santa
Clara (El Oro, Ecuador); around 230 km), and from the Yucat[aacute]n,
Mexico into the Gulf of Mexico (448 km) (Marshall et al. 2011; Guerrero
and Hearn 2017; Sotelo 2018). Off Mexico's Yucat[aacute]n peninsula,
Graham et al. (2012) calculated a maximum distance travelled by a giant
manta ray to be 1,151 km (based on a cumulative straight line distance
between locations; tag period ranged from 2 to 64 days). Similarly,
Hearn et al. (2014) report on a tagged M. birostris that was tracked
from Isla de la Plata (Ecuador) to west of Darwin Island (tag was
released after 104 days), a straight-line distance of 1,500 km, further
confirming that the species is capable of fairly long distance
migrations.
[[Page 66661]]
For the most part, these larger-scale migrations appear to be
seasonally-based for foraging purposes, as described previously, with
giant manta rays appearing in areas undergoing seasonal upwelling
events. For example, through analysis of photographs and videos of
mobulids from 1990 to 2013, Sobral and Afonso (2014) confirmed the
presence of M. birostris at the Azores islands and noted that its
occasional presence (several encounters per year) at these remote
islands indicates a strong seasonal migratory behavior. However, the
origin of these mantas, and the potential migratory paths that they use
to get to these remote islands, remain unknown.
Similarly, seasonal sightings of M. birostris off the Isla de la
Plata, Ecuador, predominantly occur from August to October, with a peak
in early September (Guerrero and Hearn 2017); however, from where these
mantas originate is currently under investigation. Recently, Sotelo
(2018) examined the genetic diversity of these manta rays from 2010 to
2013 and found that it was moderately high, with an average expected
heterozygosity value (He = 0.679) comparable to similar species that
are known to undertake long-distance migrations. The results also
suggest that the manta rays may migrate in family groups, but that they
may not always visit the same areas consistently. For example, Sotelo
(2018) found population structure between the manta rays sampled in
2013 compared to the years 2010, 2011, and 2012, with the 2013 manta
rays representing a different population. The authors note that copepod
numbers peaked at the Isla de la Plata in May of 2013, two months later
than the previous years in the study (Sotelo 2018). As manta rays
demonstrate high plasticity in terms of their movements in search of
prey, Sotelo (2018) reasoned that the change in timing of the copepod
peak likely explains why a different manta ray population visited the
island in 2013 compared to previous years. However, again, the origin
of these mantas, and the potential migratory routes traveled by these
mantas to the Isla de la Plata are currently unknown.
While long-distance migratory information is lacking, scientists
have tagged some of these mantas during their seasonal visitation to
these nearshore areas, and have gained additional information on their
smaller-scale movement patterns around and from these sites. For
example, in Isla de la Plata, two mantas were tagged from September
2017 to January 2018 with tracks that revealed coastal movements
between Ecuador and northern Peru (Sotelo 2018). These two mantas
remained within 200 km of the shoreline and did not move more than 300
km south of Isla de la Plata, where they were originally tagged.
However, based on the track lines (see Annex C; Sotelo 2018), there is
no clear migratory corridor that they appear to use, with movements
traversing throughout the entire area.
Off the Yucat[aacute]n peninsula, Graham et al. (2012) tagged 6
giant manta rays (4 females, 1 male, and 1 juvenile) and tracked their
movements for up to 64 days. The tagged manta rays traversed the
frontal zones repeatedly, probably in search of prey (Graham et al.
2012), with no clear migratory route. The majority of manta ray tracks
were more than 20 km offshore, in water depths of less than 50 m, and
the animals traveled up to 116 km from their original tagging location
(Graham et al. 2012). The authors also noted that there were no
differences in movement patterns based on sex, body size, or ambient
water-column temperature. Their conclusion, based on the tracking data,
was that giant manta rays forage over large spatial scales (~100 km
long) that are too far offshore and wide-ranging to be completely
captured in the existing Marine Protected Area networks within the
Mexican Exclusive Economic Zone (Graham et al. 2012). In other words,
there does not appear to be a specific migratory corridor that dictates
these smaller-scale foraging movements. Rather, manta rays appear to be
opportunistic feeders, with movements in and around frontal zones or
areas that are likely to contain prey.
While the available data indicate that giant manta rays may be
capable of long-distance movements, a recent study by Stewart et al.
(2016a) suggests that the species may not be as highly migratory as
previously thought. Using pop-up satellite archival tags in combination
with analyses of stable isotope and genetic data, the authors found
evidence that M. birostris off the Pacific coast of Mexico may actually
exist as well-structured subpopulations that exhibit a high degree of
residency. For example, unlike the giant manta ray in the Hearn et al.
(2014) study (that traveled from Isla de la Plata to the Galapagos
Islands), tagged M. birostris individuals from locations nearshore to
Mexico (Bahia de Banderas; n=5) and offshore Mexico (Revillagigedo
Islands; n=4) showed no movements between locations (tag deployment
length ranged from 7 days to 193 days) (Stewart et al. 2016a). The
stable isotope analysis showed higher [delta]\13\C values for the
nearshore mantas compared to those offshore, indicating these mantas
were foraging in their respective locations rather than moving between
nearshore and offshore environments (Stewart et al. 2016a).
Additionally, the genetic analysis provided evidence of population
structure between the coastal Mexico and offshore Mexico populations
(Stewart et al. 2016a). While the authors note that the species may be
capable of occasional long-distance movements, the results from their
study indicate that, for some populations, these types of long-distance
movements may be rare and may not contribute to substantial gene flow
or inter-population mixing of individuals (Stewart et al. 2016a).
Overall, the available data indicate that giant manta rays undergo
both short- and long-distance migrations; however, the space or any
specific migratory corridor used by the species during these migrations
remains unknown. In addition, we have no information on any potential
migratory corridors that may exist within waters under U.S.
jurisdiction for the giant manta ray. As mentioned previously, we are
currently supporting and conducting tracking studies of giant manta
rays within U.S. waters to better understand the fine-scale movements
of the species off the coast of Florida and within the FGBNMS. Data
from these or similar studies may reveal potential migratory corridors
preferred by giant manta rays. Similarly, survey efforts by the Georgia
Aquarium off the coast of St. Augustine, Florida, may also help
elucidate some of these questions in the future.
As noted previously in this determination, giant manta rays appear
to have a seasonal pattern of occurrence along the east coast of
Florida, showing up with greater frequencies (and in greater numbers)
in the spring and summer months. In fact, sightings of manta rays in
the region signal to fishermen the start of cobia fishing as fishermen
have found that cobia tend to closely associate with the manta rays as
they migrate along the east coast of Florida. Based on information from
recreational cobia fishing articles, manta rays tend to appear off
Florida's coast when water temperatures climb above 20 [deg]C to 21
[deg]C; however, Levesque (2019) notes that it is ``impossible to
predict when they will show up from one year to the next.'' Killer
(2010) states that in Florida's Treasure Coast waters, mantas may not
show up every year, and it is unclear where they come from or where
they go after they leave the area. Quoting two charter vessel captains,
Killer (2010) reports that the mantas have been observed along the
coast moving from south to north as waters warm, but have also been
observed doing the opposite migration,
[[Page 66662]]
with some potentially moving from offshore to inshore waters as well
during this time. McNally (2012) believes that the spring migration of
rays off northeast Florida is occurring much farther offshore than in
the past, noting that the mantas used to be observed just off the beach
breakers but are now more than 10 miles offshore. We also note that
during the migratory season, manta rays tend to be found in both
shallow and deep waters (Killer 2010; Levesque 2019), with no
information to suggest they are restricted to a certain area off the
coast of Florida.
While the available information confirms the migratory behavior of
the species in U.S. waters, the data do not indicate that there are any
specific routes or corridors that are consistently used by the species
during their migration. In fact, as noted previously, McNally (2012)
suggests that a dedicated corridor may not exist, or that some other
unknown feature may be influencing their spatial patterns during these
migrations. Additionally, Roberts (2016) notes that ``no studies have
shown a correlation of bottom structure (reef lines, continental shelf,
etc.) and the ray's migration pattern,'' nor have we come across any
studies since that article was published. Therefore, at this time, and
based on the foregoing information, we cannot identify any specific
essential features that define migratory habitat for giant manta rays.
The Physical and Biological Features of Breeding Habitat That Are
Essential to the Conservation of the Species
Little information exists on the reproductive ecology of the giant
manta ray as mating behavior of M. birostris is rarely observed in the
wild. However, based primarily on observations of M. alfredi mating
behavior, Stevens et al. (2018b) identified seven stages of courtship
for manta rays: (1) Initiation, (2) endurance, (3) evasion, (4) pre-
copulation positioning, (5) copulation, (6) post-copulation holding,
(7) separation. The initiation stage involves males shadowing females
at normal cruising speeds. During this stage, males will often attempt
to facilitate female receptiveness by using the cephalic fins to gently
stroke the females' dorsal surface. During the endurance stage,
swimming speeds increase and from 1 to 8 males follow closely behind a
single female. The evasion stage is characterized by continued close
following at increased speeds with the female incorporating rapid
maneuvers, somersaults, and flips, with males attempting to stay right
behind her. Pre-copulation positioning involves the male using his
cephalic fins to guide himself down the females' back along the leading
edge of her pectoral fin. Once at the fin's tip, the male grasps it
firmly with his mouth then rotates his body so that he is underneath
the female and the two are abdomen to abdomen. Copulation then occurs,
usually initiating near the surface, with the male continuing to move
his fins to maintain position while the female ceases movement. The
clasper is inserted in the cloaca and copulation lasts between 30 and
90 seconds, while the pair slowly sinks (Stevens et al. 2018b).
Only a few instances of courtship involving giant manta rays have
actually been observed, with only a single instance resulting in
copulation. On two separate occasions, in early August 1996 at the
Ogasawara Islands, Japan, Yano et al. (1999) witnessed a male M.
birostris chasing closely behind a female at relatively high speeds
(~10 km/hr). In both instances, the behavior was observed for
approximately 40 minutes but did not result in copulation. Stevens et
al. (2018b) also witnessed two occurrences of this ``endurance'' stage
in M. birostris, one involving a single female followed by a single
male, and the other involving a single female followed by eight males.
Both of these observations were made off of the remote island of
Fuvahmulah in the Maldives, lasted approximately one minute, and
neither resulted in observed copulation. The only observation of
successful copulation was reported by Yano et al. (1999) who witnessed
two males chasing a single female in a zigzag pattern off the Ogasawara
Islands in early July 1997. Speeds were similar to those witnessed
during other observations; however, these chases progressed all the way
through the rest of the stages of copulatory behavior (Yano et al.
1999). The chases occurred approximately 30 minutes apart, with both
males observed inserting their claspers into the same female (Yano et
al. 1999).
In terms of habitat characteristics, the mating behavior in the
Maldives location occurred at a known aggregation site for the species
(Stevens et al. 2018b). Females were chased along the reef crest of the
atolls in the area (Stevens et al. 2018b). However, while the authors
noted that most of the mating behavior for M. alfredi happened at
cleaning stations, for M. birostris, the mating occurred at locations
where giant manta rays tend to just pass through (Stevens et al.
2018b). In other words, the area where the mating behavior was observed
did not appear to have any other significance for the species. Off the
Ogasawara Islands, Japan, Yano et al. (1999) described the site of the
mating behavior as 100-200 m offshore of the east coast of Chichijima
(one of the Ogasawara Islands), within an area comprised of rocky reefs
in 10-20 m depth. The authors noted that each copulation event happened
within one meter of the surface (Yano et al. 1999).
Giant manta ray breeding sites are also thought to occur off
Ecuador and the Galapagos Islands based on the presence of pregnant
females and recent mating scars. In fact, some of the first pregnant
females ever seen in the wild have been sighted in the productive
coastal waters off Isla de la Plata in the Machalilla National Park,
Ecuador. According to Guerrero and Hearn (2017), between 2009 and 2015,
8 pregnant giant mantas were observed off Isla de La Plata, with 7 of
these reported in 2011. Additionally, photographic records from 2012 to
2015 showing fresh scars on the pectoral fins of mature female giant
manta rays around Isla de la Plata and Bajo Cop[eacute] indicate the
likely use of these Ecuadorian aggregation sites as mating areas
(Guerrero and Hearn 2017). In terms of habitat characteristics of these
areas, the authors note that the majority of giant manta rays seen in
Isla de la Plata are off the northwest area of the island, in Punta El
Faro, Roca Honda, and La Pared (Guerrero and Hearn 2017). These
particular areas are close to deep waters, with a bottom characterized
by coarse sand and scattered rocks. Calcareous coral formations can be
found between 0 and 14 m depths and soft corals (gorgonians) can be
found in deeper depths (Guerrero and Hearn 2017). La Pared, in
particular, contains pinnacles and rocks that extend to the northwest
and create an edge with a steep drop to 52 m depths (Guerrero and Hearn
2017). The authors state that giant manta rays do not remain in the
area for very long (usually around a few days to a week), but may
return in multiple years and hypothesize that their purpose for
visiting the island could be primarily for cleaning purposes, mating,
and/or feeding as all three behaviors are observed at this site
(Guerrero and Hearn 2017).
Within U.S. waters, there are very few observations of mating
behavior. In our collection of manta ray sightings and videos, there
are only 4 records of ``chasing'' or ``courtship'' behavior of M.
birostris. Three of the records are from diver observations off the
west coast of Hawaii (Manta Pacific Research Foundation 2019), and the
fourth is from an instagram video off Avon Fishing Pier, North
Carolina, taken in July 2019 (G. Stevens, Manta Trust,
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pers. comm. to C. Horn, NMFS SERO, 2019); however, there is no
corresponding information regarding habitat features related to these
records (just individual sightings data). Given that the areas where
giant manta ray mating occurs remain largely unknown, with only a few,
opportunistic observations of courtship behavior or evidence of
breeding (i.e., mating scars, pregnant females) in a couple of
locations, there has not been any systematic evaluation of the
particular physical or biological features that facilitate or are
necessary for mating to occur. The general habitat characteristics
mentioned above in relation to the observations of mating behavior,
including presence of rocky and coral reefs, shallow depths, coarse
sand, and reef crests adjacent to deep water, are found throughout the
species' range and are commonly associated with giant manta ray
sightings (Yano et al. 1999; Childs 2001; Kashiwagi et al. 2011;
Marshall et al. 2011; Stevens et al. 2018b; Stewart et al. 2018b).
However, not all areas with the above features provide meaningful
mating habitat as, for example, many of the observations from the
studies previously discussed (for foraging, pupping, and migratory
habitat) also noted the presence of these habitat features but did not
observe mating behavior in M. birostris. As such, at this time, the
available information does not allow us to identify any physical or
biological features within these areas where mating has been observed
that are essential to support this behavior.
Unoccupied Areas
Section 3(5)(A)(ii) of the ESA defines critical habitat to include
specific areas outside the geographical area occupied by a threatened
or endangered species at the time it is listed if the areas are
determined by the Secretary to be essential for the conservation of the
species. Regulations at 50 CFR 424.12(b)(2) address designation of
unoccupied area as critical habitat and the regulations at 50 CFR
424.12(g) state that critical habitat shall not be designated within
foreign countries or in other areas outside of United States
jurisdiction.
As discussed previously, the waters off the U.S. west coast are not
considered part of the geographical area occupied by giant manta ray at
the time of listing. We also conclude that it is not an unoccupied area
essential to the species' conservation given the rare, errant use of
the area by a vagrant giant manta ray in the past, and no information
to suggest the area is essential to the conservation of the species.
The other geographical areas under U.S. jurisdiction that were not
included in the discussion of occupied areas by the giant manta ray
(i.e., U.S. waters north of Long Island, New York) are considered to be
out of the species' livable range and, thus, would not be essential to
the conservation of the species. As such, we find that there are no
specific areas outside the geographical areas occupied by M. birostris
that would meet the definition of critical habitat for the giant manta
ray.
Critical Habitat Determination
Given the best available information and the above analysis of this
information, we find that there are no identifiable occupied areas
under the jurisdiction of the United States with physical or biological
features that are essential to the conservation of the species or
unoccupied areas that are essential to the conservation of the species.
Therefore, we conclude that there are no specific areas within the
giant manta ray range and under U.S. jurisdiction that meet the
definition of critical habitat. Per 50 CFR 424.12(a)(1)(iv), if no
areas meet the definition of ``critical habitat,'' then we can conclude
that a designation of critical habitat is not prudent.
Although we have made this ``not prudent'' determination, the areas
occupied by giant manta rays under U.S. jurisdiction will continue to
be subject to conservation actions implemented under section 7(a)(1) of
the ESA, as well as consultation pursuant to section 7(a)(2) of the ESA
for Federal activities that may affect the giant manta ray, as
determined on the basis of the best available information at the time
of the action. Through the consultation process, we will continue to
assess effects of Federal actions on the species and its habitat.
Additionally, we remain committed to promoting the recovery of the
giant manta ray through both domestic and international efforts. As
noted in the proposed and final rules (82 FR 3694, January 12, 2017; 83
FR 2916, January 22, 2018, respectively), the most significant threat
to the giant manta ray is overutilization by commercial and artisanal
fisheries operating within the Indo-Pacific and eastern Pacific
portions of its range, primarily in areas outside of U.S. jurisdiction.
Giant manta rays are both targeted and caught as bycatch in a number of
fisheries throughout their range, and while the majority of these
fisheries target manta rays for their meat, there has been an
increasing demand for manta ray gill plates for use in Asian medicine,
primarily in the Indo-West Pacific. Efforts to address overutilization
of the species through regulatory measures appear inadequate, with
evidence of targeted fishing of the species despite prohibitions in a
number of countries, and only one regional fisheries management
organization measure to address bycatch issues (Miller and Klimovich
2017). Thus, recovery of the giant manta ray is highly dependent upon
international conservation efforts. To address this, we have developed
a recovery plan outline that provides our preliminary strategy for the
conservation of the giant manta ray. This outline can be found on our
website at: https://www.fisheries.noaa.gov/species/giant-manta-ray#
resources and provides an interim recovery action plan as well as
preliminary steps we will take towards the development of a full
recovery plan.
Currently, we are actively engaged in manta ray research to gain a
better understanding of the biology, behavior, and ecology of this
threatened species. We are presently working on collecting and
assimilating anecdotal and survey-related manta sightings and effort
data to support the development of an ensemble species distribution
model for the southeastern United States. We are also collaborating
with partners to examine giant manta ray movements in U.S. waters off
Florida and within the FGBNMS. This data will provide a better
understanding of giant manta ray movements and habitat use, including
environmental drivers of movement. We are also supporting research
projects assessing the survivorship of giant manta rays caught in
Peruvian and Indonesian artisanal gillnet fisheries.
We have developed safe handling and release guidelines for
fishermen (available at: https://www.fisheries .noaa.gov/webdam/
download/91927887). In an effort to address species identification
issues during aerial surveys, we have also developed an aerial survey
mobulid species identification key that will facilitate accurate
species identification in the future. We added the giant manta ray to
our Northeast and Southeast Observer Program capture reports, logbooks,
and manuals/reports, and provided a guide to the identification of
mobulid rays to observers to gain more accurate information regarding
the species' distribution and prevalence in U.S. fisheries. In
addition, we have set up a dedicated email (i.e., [email protected])
for the public to report giant manta ray encounters to help us learn
more about M. birostris movement patterns, habitat use, and human
interactions in our waters. We will continue to work towards the
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conservation and recovery of giant manta rays, both on a domestic and
global level, including with our international partners and within
regional fisheries management organizations and other international
bodies to promote the adoption of conservation and management measures
for the threatened giant manta ray.
References
A complete list of all references cited herein is available upon
request (see FOR FURTHER INFORMATION CONTACT).
Authority
The authority for this action is the Endangered Species Act of
1973, as amended (16 U.S.C. 1531 et seq.).
Dated: December 2, 2019.
Samuel D. Rauch III,
Deputy Assistant Administrator for Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2019-26265 Filed 12-4-19; 8:45 am]
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