Endangered and Threatened Wildlife and Plants; Final Endangered Listing of Five Species of Sawfish Under the Endangered Species Act, 73977-74005 [2014-29201]
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National Oceanic and Atmospheric Administration
50 CFR Part 224
Endangered and Threatened Wildlife and Plants; Final Endangered Listing
of Five Species of Sawfish Under the Endangered Species Act; Final Rule
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Federal Register / Vol. 79, No. 239 / Friday, December 12, 2014 / Rules and Regulations
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
50 CFR Part 224
[Docket No 101004485–4999–03]
RIN 0648–XZ50
Endangered and Threatened Wildlife
and Plants; Final Endangered Listing
of Five Species of Sawfish Under the
Endangered Species Act
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
ACTION: Final rule.
AGENCY:
We, NMFS, issue this final
rule implementing our determination
that the narrow sawfish (Anoxypristis
cuspidata), dwarf sawfish (Pristis
clavata), largetooth sawfish (collectively
Pristis pristis; formerly Pristis pristis,
Pristis microdon, and Pristis perotteti),
green sawfish (Pristis zijsron), and the
non-U.S. distinct population segment
(DPS) of smalltooth sawfish (Pristis
pectinata) are endangered species under
the Endangered Species Act (ESA) of
1973, as amended. We also include a
change in the scientific name for
largetooth sawfish in this final rule to
codify the taxonomic reclassification of
P. perotteti to P. pristis. We are not
designating critical habitat because the
geographical areas occupied by the
species are entirely outside U.S.
jurisdiction and we have not identified
any unoccupied areas within U.S.
jurisdiction that are essential to the
conservation of any of the five species.
We have reviewed the status of the five
species of sawfish, considered public
and peer review comments, and
conservation efforts being made to
protect all five species, and we have
made our determination based on the
best available scientific and commercial
data that all five species of sawfish—the
narrow sawfish (Anoxypristis
cuspidata), dwarf sawfish (Pristis
clavata), largetooth sawfish (collectively
Pristis pristis; formerly Pristis pristis,
Pristis microdon, and Pristis perotteti),
green sawfish (Pristis zijsron), and the
non-U.S. DPS of smalltooth sawfish
(Pristis pectinata)—are at risk of
extinction throughout all of their ranges
and should be listed as endangered
species.
DATES: This final rule is effective
January 12, 2015.
ADDRESSES: Information regarding this
final rule may be obtained by contacting
NMFS, Protected Resources Division,
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SUMMARY:
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263 13th Avenue South, St. Petersburg,
Florida, 33701. The final rule and
citation list are located on our Web site
at https://sero.nmfs.noaa.gov/protected_
resources/sawfish/.
FOR FURTHER INFORMATION CONTACT:
Shelley Norton, NMFS, Southeast
Regional Office (727) 824–5312 or Dr.
Dwayne Meadows, NMFS, Office of
Protected Resources (301) 427–8403.
SUPPLEMENTARY INFORMATION:
Background
On September 10, 2010, we received
a petition from the WildEarth Guardians
(WEG) requesting we list six sawfish
species—knifetooth, narrow, or pointed
sawfish (A. cuspidata), hereinafter the
narrow sawfish; dwarf or Queensland
sawfish (P. clavata), hereinafter the
dwarf sawfish; largetooth sawfish (P.
pristis and P. microdon); green sawfish
(P. zijsron); and the non-listed
population(s) of smalltooth sawfish (P.
pectinata)—as endangered or threatened
under the ESA; or alternatively, list any
distinct population segments (DPS) that
exist under the ESA. On March 7, 2011,
we published a 90-day finding (76 FR
12308) stating the petitioned action may
be warranted for five of the six species.
The five species were A. cuspidata, P.
clavata, P. microdon, P. zijsron, and the
non-listed population(s) of P. pectinata.
Information in our records at the time
indicated that P. pristis, as described in
the petition, was not a valid species.
Our 90-day finding requested
information to inform our decision, and
announced the initiation of status
reviews for the five species. On June 4,
2013, we published a proposed rule (78
FR 33300) to list A. cuspidata, P.
clavata, P. pristis (formerly P. pristis, P.
microdon, and P. perotteti), P. zijsron,
and the non-U.S. DPS of P. pectinata as
endangered. We also included a change
in the scientific name for largetooth
sawfish in the proposed rule to codify
the taxonomic reclassification of P.
perotteti to P. pristis. The largetooth
sawfish (P. perotteti) was already listed
as endangered on July 12, 2011 (76 FR
40822), but this listing decision
concerns the entire largetooth sawfish
(P. pristis) species as it is currently
classified, which also includes the
species formerly classified as P.
perotteti and P. microdon. We did not
propose to designate critical habitat
because the geographical areas occupied
by the species are entirely outside U.S.
jurisdiction and we did not identify any
unoccupied areas that are currently
essential to the conservation of any of
these species. We solicited public and
peer reviewer comments on the
proposed rule and also coordinated
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outreach on the proposed rule with the
Department of State to give notice to
foreign nations where the species are
believed to occur.
We are responsible for determining
whether species are threatened or
endangered under the ESA (16 U.S.C.
1531 et seq.). To make this
determination, we first consider
whether a group of organisms
constitutes a ‘‘species’’ under the ESA,
then whether the status of the species
qualifies it for listing as either
threatened or endangered. Section 3 of
the ESA defines a ‘‘species’’ as ‘‘any
subspecies of fish or wildlife or plants,
and any distinct population segment of
any species of vertebrate fish or wildlife
which interbreeds when mature.’’ On
February 7, 1996 (61 FR 4722), NMFS
and the U.S. Fish and Wildlife Service
(USFWS; collectively, the Services)
adopted a policy identifying two
elements that must be considered when
identifying a DPS: (1) The discreetness
of the population segment in relation to
the remainder of the species (or
subspecies) to which it belongs; and (2)
the significance of the population
segment to the remainder of the species
(or subspecies) to which it belongs. As
stated in the DPS policy, Congress
expressed its expectation that the
Services would exercise their authority
with regard to the use of DPSs sparingly
and only when the biological evidence
indicates such action is warranted.
Section 3 of the ESA defines an
endangered species as ‘‘any species
which is in danger of extinction
throughout all or a significant portion of
its range’’ and a threatened species as
one ‘‘which is likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range.’’ Thus
we interpret an ‘‘endangered species’’ to
be one that is presently in danger of
extinction. A ‘‘threatened species,’’ is
not presently in danger of extinction,
but is likely to become so in the
foreseeable future (that is, at a later
time). In other words, the primary
statutory difference between a
threatened and endangered species is
the timing of when a species may be in
danger of extinction— either presently
(endangered) or in the foreseeable future
(threatened).
Section 4(a)(1) of the ESA requires us
to determine whether any species is
endangered or threatened due to any
one or a combination of the following
five factors: (A) The present or
threatened destruction, modification, or
curtailment of its habitat or range; (B)
overutilization for commercial,
recreational, scientific, or educational
purposes; (C) disease or predation; (D)
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the inadequacy of existing regulatory
mechanisms; or (E) other natural or
manmade factors affecting its continued
existence. We are required to make
listing determinations based solely on
the best scientific and commercial data
available after conducting a review of
the status of the species and after taking
into account efforts being made by any
state or foreign nation to protect the
species.
Accordingly, we have followed a
stepwise approach in making our listing
determinations for A. cuspidata, P.
clavata, P. pristis (formerly P. pristis, P.
microdon, and P. perotteti), P. zijsron,
and the non-U.S.DPS of P. pectinata.
For the non-U.S. DPS of P. pectinata
that may qualify as a DPS, we
considered biological evidence, such as
genetic information to determine if the
population met the DPS policy criteria.
Using the best available information
gathered during the status reviews, we
completed an extinction risk assessment
using the general procedure of
Wainwright and Kope (1999). We then
assessed the threats affecting the status
of each species using the five factors
identified in section 4(a)(1) of the ESA,
and then assessed public and peer
reviewer comments.
Once we determined the threats, we
assessed the efforts being made to
protect each species to determine if
these conservation efforts were adequate
to mitigate the existing threats and alter
extinction risk. We evaluated
conservation efforts using the criteria
outlined in the joint NMFS and U.S.
Fish and Wildlife Service (USFWS)
Policy for Evaluating Conservation
Efforts (PECE; 68 FR 15100; March 28,
2003) to determine the certainty of
implementation and effectiveness for
future conservation efforts not yet fully
implemented or effective. Finally, we
re-assessed the extinction risk of each
species after considering the existing
conservation efforts.
In order to conduct a comprehensive
review, NMFS Southeast Region
Protected Resources Division and NMFS
Southeast Fisheries Science Center staff
members collaborated to identify the
best available information. Unlike some
of our previous 12-month findings, we
did not develop a separate status review
report. Instead, we presented all
information available for these species
in the proposed rule, and we present
that information again, as modified by
public comment on the proposed rule,
in this final rule. We first discuss
background information relative to all
five species, and then we include
descriptions of the natural history
specific to each species.
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Sawfish General Species Description
Sawfishes are a group of shark-like
rays. Taxonomically, they are classified
in the Family Pristidae (sawfishes),
Order Rajiformes (skates, rays, and
sawfishes), subclass (Elasmobrancii),
and Class Chondrichthyes (cartilaginous
fish). The overall body form of
sawfishes is similar to sharks, but they
are flattened dorso-ventrally. Sawfishes
are covered with dermal denticles
(teeth-like scales) and possess enlarged
pectoral fins.
The most distinct characteristic of
sawfishes is their large, flat, toothed
rostrum or ‘saw’ with large teeth on
each side. The rostral teeth are made
from calcified tissue that is neither
dentin nor enamel, though it is more
similar to the latter (Bradford, 1957).
Rostral teeth develop inside sockets on
the rostrum and are held in place by
strong fibers. Unlike sharks, sawfish
rostral teeth are not replaced, although
partially broken teeth may continue to
grow (Miller, 1974). For some species of
sawfish, the number of rostral teeth can
vary by geographic region.
Sawfishes use their rostrum to locate,
stun, and kill prey, generally small
schooling fishes such as mullet, herring,
shad, and sardines (Bigelow and
Schroeder, 1953). Breder (1952), in
summarizing the literature on
observations of sawfish feeding
behavior, noted that they attack fish by
slashing sideways through schools of
fish, and then impale the fish on their
rostral teeth. Prey are subsequently
scraped off their rostral teeth by rubbing
the rostrum on the bottom and then
ingesting the whole fish. Bigelow and
Schroeder (1953) also report that
sawfish feed on crustaceans and other
benthic species. Recent studies indicate
that sawfishes may use their toothed
rostrum to sense their prey’s electric
fields (Wueringer et al., 2011; 2012).
Sawfish species are distributed
primarily in circumtropical shallow
coastal waters that generally vary in
salinity. While sawfishes are commonly
found in shallow water, adults are
known to also inhabit deeper waters
(greater than 130 ft, 39.6 m). Some
sawfishes are found in freshwater, with
established populations in major rivers
and lakes of South America, Africa,
Australia, and Southeast Asia. The
physical characteristics of habitat, such
as salinity and temperature, likely
influence a sawfish’s movement
patterns. Tides limit the physical habitat
area available, which may explain
movement into shallow water areas
during specific tidal cycles (Blaber et
al., 1989).
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Life history data on sawfishes are
limited. Fertilization is internal by
means of male claspers and
reproduction is ovoviviparous; females
carry eggs with a yolk sac that nourishes
developing young until they hatch
within the body. Sawfishes are born
with a gelatinous substance around their
rostral teeth to protect the mother
during birth (Last and Stevens, 1994;
Rainboth, 1996; Compagno and Last,
1999; Raje and Joshi, 2003; Field et al.,
2009). It is thought that most sawfishes
breed every two years and have a
gestation period of about four to five
months (Bigelow and Schroeder, 1953;
Thorson, 1976a). The number of young
in a litter varies by species, as does the
age at sexual maturity.
Like most chondrichthyes, sawfishes
occupy the mid- to upper-level of their
food web. Smaller sawfishes, including
juveniles, may be preyed upon by larger
sharks like the bull shark (Carcharhinus
leucas), estuarine crocodiles
(Crocodylus porosus), or alligators
(Alligator mississippiensis). Sawfishes
may use their saw as a weapon for
defense against these predators (Brewer
et al., 1997; Wueringer et al., 2009).
Previously, seven valid species of
sawfish were recognized worldwide
(Compagno, 1999). Compagno and Cook
(1995) and Compagno (1999) identified
these seven species of sawfish as A.
cuspidata Latham 1794, P. microdon
Latham 1794, P. perotteti Muller and
Henle 1841, P. pristis Linnaeus 1758, P.
clavata Garman 1906, P. pectinata
Latham 1794, and P. zijsron Bleeker
1851. Since then, the taxonomy,
delineation, and identification of these
species have proven problematic (Oijen
et al., 2007; Wiley et al., 2008;
Wueringer et al., 2009). Most recently,
Faria et al. (2013) hypothesized that the
taxonomic uncertainty occurred due to
several factors: many original species
descriptions were abbreviated, few
holotypes are available for examination,
reference material is not available for
comparison in museum collections, and
it is difficult to obtain fresh specimens
because of the infrequent captures of all
sawfishes. The majority of the confusion
regarding taxonomic classification of
Pristidae was related to the species P.
pristis. To resolve questions regarding
the taxonomy of pristids, Faria et al.
(2013) used historical taxonomy,
external morphology, and mitochondrial
DNA (mtDNA) sequences (NADH–2
loci) to conclude that sawfishes have
five species in two genera: P. pristis, P.
clavata, P. pectinata, P. zijsron, and A.
cuspidata. We accept this proposed
taxonomy as the best available science.
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Natural History of the Narrow Sawfish
(Anoxypristis cuspidata)
Taxonomy and Morphology
The narrow sawfish was first
described by Latham in 1794 as P.
cuspidatus. It was later reclassified as
Anoxypristis due to morphological
differences from Pristis that include its
narrow rostral saw, which lacks teeth on
the first quarter of the saw closest to the
head in adults, as well as the distinct
shape of the lower lobe of the caudal fin
(Compagno et al., 2006a). In juveniles,
the portion of the rostrum without teeth
is only about one-sixth of the saw length
(Wueringer et al., 2009).
In addition, the narrow sawfish is
characterized by dagger-shaped rostral
teeth (Fowler, 1941; Blegvad and
Loppenthin, 1944; Compagno and Last,
1999; Faria et al., 2013). The narrow
sawfish also has a second pair of hollow
cartilaginous tubes in its rostrum that
are not present in other sawfishes.
These canals contain an additional
connection to the ampullae of Lorenzini
(special sensory receptors) located on
the underside of the rostrum (Wueringer
et al., 2009).
Rostral tooth count varies for this
species between 18 and 22 (Last and
Stevens, 1994), 24 and 28 (Hussakof,
1912), and 27–32 (Miller, 1974). The
total number of teeth has been found to
vary by individual, region, and sex.
Some studies report males having fewer
rostral teeth than females, while others
report the opposite (Last and Stevens,
1994; Compagno and Last, 1999). While
total rostral tooth count is often
inconsistent among individuals or
studies, the number of teeth an
individual has is fixed during
development (Wueringer et al., 2009).
The pectoral fins of the narrow
sawfish are narrow, short, and sharklike in shape. The first dorsal fin is
located posterior to the insertion of the
pelvic fins (Compagno and Last, 1999).
Within the jaw, there are 94 teeth on the
upper jaw and 102 on the lower jaw
(Taniuchi et al., 1991a). The eyes are
large and very close to the spiracles.
Coloration is dark grey dorsally and
whitish ventrally (Fowler, 1941;
Compagno and Last, 1999).
Narrow sawfish are the only sawfish
having tricuspid (three-pointed)
denticles (White and Moy-Thomas,
1941). These denticles first appear on
sawfish at 25.6 to 28 in (65 to 71 cm)
total length (TL), after they are born. In
general, the narrow sawfish is
considered ‘‘naked’’ because denticle
coverage in adults is often sporadic and
widely spaced, usually only covering
the rostrum and anterior fin margins,
making the skin appear smooth (Fowler,
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1941; Gloerfelt-Tarp and Kailola, 1984;
Last and Stevens, 1994; Wueringer et al.,
2009). Narrow sawfish also have
buccopharyngeal denticles (tooth-like
structures) present in their mouth. This
species does not have tubercles or
thorns on their skin (Deynat, 2005).
Habitat Use and Migration
The narrow sawfish is largely
euryhaline and moves between
estuarine and marine environments
(Gloerfelt-Tarp and Kailola, 1984; Last,
2002; Compagno, 2002b; Compagno et
al., 2006a; Peverell, 2008). It is generally
found in inshore waters in depths of
less than 130 ft (39.6 m) with salinities
between 25 and 35 parts per thousand
(ppt), spending most of its time near the
substrate or in the water column over
coastal flats (Compagno and Last, 1999;
Last, 2002; Peverell, 2005; Peverell,
2008; Wueringer et al., 2009). While
Smith (1936) described it as a possible
freshwater species, there are only a few
reports from freshwater (Taniuchi and
Shimizu, 1991; Last and Compagno,
2002; Bonfil and Abdallah, 2004;
Wueringer et al., 2009). We are not
aware of any fresh or salt water
tolerance studies on the species
(Compagno, 2002a; Compagno, 2002b)
and conclude its habitat is euryhaline.
In studies conducted by Peverell
(2008), the narrow sawfish in the Gulf
of Carpentaria, Australia, undergo an
ontogenetic shift in habitat. Larger
individuals were commonly
encountered offshore, while smaller
individuals were mostly found in
inshore waters. Peverell (2008) also
found females were more likely to be
offshore compared to males, at least
during the months of the study
(February to May). This suggests that
smaller narrow sawfish use the
protection and prey abundance found in
shallow, coastal waters (Dan et al., 1994;
Peverell, 2005; Peverell, 2008).
Age and Growth
Two studies have been conducted on
age and growth of narrow sawfish. Field
et al. (2009) compared previously-aged
vertebrae with aged rostral teeth and
found a direct correlation up to age 6.
After age 6, an individual’s age was
often underestimated using tooth
growth bands as the teeth become worn
over time (Field et al., 2009). Peverell
(2008) then used aged vertebrae to
develop more accurate growth curves
for both sexes. While the maximum
observed age of narrow sawfish from
vertebrae was 9 years, the theoretical
longevity was calculated at 27 years
(Peverell, 2008). A 1-year-old animal
has a saw length of approximately 4.5 in
(11.5 cm). Female narrow sawfish begin
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to mature at 8 ft 1 in (246 cm) TL and
all are mature at 15 ft 5 in (470 cm) TL;
males are mature at 8 ft (245 cm) TL
(Pogonoski et al., 2002; Bonfil and
Abdallah 2004; Peverell, 2005; 2008).
The maximum recorded length of a
narrow sawfish is 15 ft 5 in (4.7 m) TL,
with unconfirmed records of 20 ft (6.1
m) TL (Last and Stevens, 1994;
Compagno and Last, 1999; Pogonoski et
al., 2002; Bonfil and Abdallah, 2004;
Faria et al., 2013).
Reproduction
The narrow sawfish gives birth to a
maximum of 23 pups in the spring. The
total length (TL) of pups at birth is
between 17–24 in (43–61 cm)
(Compagno and Last, 1999; Peverell,
2005; 2008). The reproductive cycle is
assumed to be annual, with an average
of 12 pups per litter (Peverell, 2005;
D’Anastasi, 2010). The number of pups
is related to female body size, as smaller
females produce fewer offspring than
larger females (Compagno and Last,
1999). Preliminary genetic research
suggests that the narrow sawfish may
not have multiple fathers per litter
(D’Anastasi, 2010).
Mating season may vary by
geographic region. Female narrow
sawfish captured in August (dry season)
in the Gulf of Carpentaria, Australia, all
contained large eggs indicating they
were mature (Peverell, 2005). Mature
males were also captured in similar
locations during the same time of year
(McDavitt, 2006). Although animals are
sexually mature in the dry season,
mating may not occur until the rainy
season in March-May in the Indo-West
Pacific (Raje and Joshi, 2003).
Age at maturity for narrow sawfish is
2 years for males and 3 years for females
(Peverell, 2008). The intrinsic rate of
population increase (rate of growth of
the population) based on life history
data from the exploited population in
the Gulf of Carpentaria, Australia, has
been estimated at 0.27 per year (Moreno
Iturria, 2012), with a potential
population doubling time of 2.6 years.
Diet and Feeding
Narrow sawfish feed on small fish and
cuttlefish (Compagno and Last, 1999;
Field et al., 2009) and likely on
crustaceans, polychaetes, and
amphipods (Raje and Joshi, 2003).
Population Structure
Genetic and morphological data
support the division of the global
species of narrow sawfish into
populations. Based on gene sequence
data, there is a very low level of gene
flow between the northern Indian Ocean
(n = 2) and west Pacific (n = 11)
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populations. Four haplotypes
(combinations of deoxyribonucleic acid
sequences or DNA) were identified:
northern Indian Ocean; Indonesian;
New Guinean–Australian; and one
specimen that lacked locality
information, but had a northern Indian
Ocean haplotype. Specimens collected
from the Indian Ocean had a higher
number of rostral teeth per side than
those collected from the western Pacific
(Faria et al., 2013).
Field et al. (2009) examined the
primary chemical elements of rostral
teeth (i.e., oxygen, calcium, and
phosphorous) from narrow sawfish
captured throughout Australia in an
attempt to separate subpopulations
based on the isotopes of these
chemicals. They found distinctions
between regions indicating two separate
subpopulations within the Gulf of
Carpentaria Australia: one in the west
(Northern Territory) and one in the east
(Queensland). Using isotopes to separate
elasmobranch subpopulations is in its
infancy, however, and, coupled with the
limited number of samples, it is not
clear whether these results agree with
the above genetic studies of population
structure. Isotopic signatures indicate
the location where an animal spends
most of its time and identifies its major
prey resources and do not necessarily
provide information on reproductive
connectivity between regions.
Therefore, we conclude that the best
available information on isotopic
signatures does not support separating
narrow sawfish into subpopulations.
Distribution and Abundance
The narrow sawfish is found
throughout the eastern and western
portions of the Indian Ocean as well as
much of the western Pacific Ocean. The
range once extended from as far west as
the Red Sea in Egypt and Somalia (M.
McDavitt, National Legal Research
Group, Inc. pers. comm. to IUCN,
London, 2012) to as far north as
Honshu, Japan, including India, Sri
Lanka, and China (Blaber et al., 1994;
Last and Stevens, 1994; Compagno and
Last, 1999; Compagno et al., 2006a; Van
Oijen et al., 2007). The species has also
been recorded in rivers in India, Burma,
Malaysia, and Thailand (Compagno,
2002b).
While uncertain, the current status of
narrow sawfish populations across its
range has declined substantially from
historic levels. The species was
previously commonly reported
throughout its range, but it is now
becoming rare in catches by both
commercial and recreational fishers
(Brewer et al., 2006; Compagno et al.,
2006a). To evaluate the current and
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historic distribution and abundance of
the narrow sawfish, we conducted an
extensive search of peer-reviewed
publications and technical reports,
newspaper, and magazine articles. We
also reviewed records from the Global
Biodiversity Information Facility (GBIF)
database (www.gbif.org). The results of
that search are summarized by major
geographic region.
Indian Ocean
The earliest reports of narrow sawfish
in the Indian Ocean were from 1937 and
1938. Two sawfish were captured from
the northern Indian Ocean (no specific
location was reported). A third
specimen was later caught in the same
area (Blegvad and Loppenthin, 1944).
From areas in the western Indian
Ocean around the Arabian Sea, three
rostra were collected in 1938: Two near
Bushire, Iran, presumably from the Gulf
of Oman, and a third in Jask, Iran, also
adjacent to the Gulf of Oman (Blegvad
and Loppenthin, 1944). The most
extensive report was 13 rostra from the
Persian Gulf (one of those was from
Iran) but it did not include date
information. Four juveniles were
recorded in Pakistan waters in 1975:
Two females and two males (Faria et al.,
2013). The last published record of
narrow sawfish from the western edge of
the range, in the Straits of Hormuz, was
in 1997 (A. Moore, RSK Environment
Ltd., pers. comm. to IUCN, 2012).
Most records of narrow sawfish in the
Indian Ocean are from the Bay of
Bengal. In 1960 and 1961, 118 sawfish,
mostly narrow sawfish, were captured
during fishery surveys using gillnets
and long lines (James, 1973). There are
several additional records of rostra from
Bangladesh in the 1960s (Faria et al.,
2013). One record from the California
Academy of Sciences is from a fish
market in Bangkok, Thailand in 1961. A
narrow sawfish was used for a 1969
parasitological study in Bangladesh, but
no further information was recorded
(Moravec et al., 2006). Faria et al. (2013)
also reported one specimen from 1976,
as well as 11 more records off India, but
no dates were recorded. Narrow sawfish
were recorded from the Kirachi West
Wharf Fish Market in Pakistan in 1978
(GBIF Database). From 1982 to 1994,
one juvenile female, one juvenile male,
and three rostra were recorded in
Pondicherry, India (Deynat, 2005). Two
female neonate specimens were
recorded in Sri Lanka, and three
juveniles (two males and one female)
from Malabar in Southwest India were
also reported from 1982–1994 (Deynat,
2005). Between 1981 and 2000, in the
Bay of Bengal, total elasmobranch
landings records are dominated by rays
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and include narrow sawfish (Raje and
Joshi, 2003). Landings of narrow sawfish
are currently reported from the Indian
Ocean off India although they are
infrequent (K.K. Bineesh, Marine
Fisheries Research Institute, Department
of Pelagic Fisheries, India, pers. comm.
to IUCN, 2012).
Indo-Pacific Ocean (excluding
Australia)
There are several accounts of narrow
sawfish over time from various
unspecified locations throughout the
Indo-Pacific. One narrow sawfish
specimen was recorded from Mabe,
India in 1835, making it the oldest
museum record from the region (GBIF
Database). The first records of narrow
sawfish were for juvenile males in 1852
and 1854 (Faria et al., 2013). A female
and male were recorded in 1867, but no
exact location was specified (Faria et al.,
2013). In 1879, one male and one female
were also recorded from Indonesia and
four rostra were reported from China in
1898 (Faria et al., 2013).
The next reports of narrow sawfish
from the Indo-Pacific occurred in the
1930s. A female was reported in 1931 in
Indonesia (no specific location), and a
male was reported in Singapore in 1937
(Blegvad and Loppenthin, 1944). A
narrow sawfish was caught in the Gulf
of Thailand in March 1937 (Blegvad and
Loppenthin, 1944). A single report from
Papua New Guinea was recorded in
1938 (Faria et al., 2013). In 1945, narrow
sawfish were reported in the Chao
Phraya River, Thailand and its
tributaries (Smith, 1945). In 1952, two
females were captured from Batavia,
Semarang, Indonesia along with a third
female without a rostrum (Van Oijen et
al., 2007).
Records of narrow sawfish throughout
the Indo-Pacific were scattered and
infrequent throughout the 1950s. Faria
et al. (2013) recorded rostra from Papua
New Guinea; two from 1955 and one
each from 1966, 1980, and 2000. A male
was caught in 1989 from the Oriomo
River, Papua New Guinea (Taniuchi et
al., 1991b; Taniuchi and Shimizu, 1991;
Taniuchi, 2002). There are other reports
of narrow sawfish from Papua New
Guinea around the Gulf of Papua and in
Bootless Bay from the 1970s, but there
are no recent records (Taniuchi et al.,
1991b). In a comprehensive literature
search for the period 1923 to 1996 on
the biodiversity of elasmobranchs in the
South China Sea, Compagno (2002a)
found no records of sawfishes. Yet, fresh
dorsal and caudal fins of narrow sawfish
were found during a survey of fish
markets from 1996 to 1997 in Thailand
(Manjaji, 2002b).
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There are even fewer records of
narrow sawfish from the Indo-Pacific
over the last few decades. The only
known specimen in the twenty-first
century is a single report from New
Guinea in 2001 (L. Harrison, IUCN, pers.
comm. to John Carlson, NMFS, 2012).
Australia
Australia may have larger populations
of narrow sawfish than any other area
within the species’ range (Peverell,
2005). According to the GBIF Database
for Australia flora and fauna, the first
museum record of the narrow sawfish in
Australia is from the Australia Museum
in Townsville, Queensland in 1963.
This database also lists observations of
narrow sawfish throughout the 1980s,
mostly recorded by the Commonwealth
Scientific and Industrial Research
Organization (CSIRO) Marine and
Atmospheric Research group. One
individual was observed in Western
Australia in 1982 and in 1983. In 1984,
CSIRO observed one narrow sawfish just
west of Darwin, Northern Territory, and
five in the Gulf of Carpentaria (three in
the east and two in the northwest). Five
additional records in 1984 were from
the northwest tip of the western Gulf of
Carpentaria, one from outside the Daly
River, and three outside of Kakadu
National Park. In 1985, two narrow
sawfish were observed near Marchinbar
Island, Northern Territory. In the
eastern Gulf of Carpentaria, four narrow
sawfish were observed in 1986, with
single observations in 1987 and 1988. In
1988, a narrow sawfish was observed in
Western Australia. Two narrow sawfish
were reported from the Gulf of
Carpentaria in 1990 (Blaber et al., 1994).
Single specimens were captured in 1991
from the west coast of Australia
(Alexander, 1991), the Gulf of
Carpentaria in 1995 (Brewer et al.,
1997), and the Arafura Sea in 1999
(Beveridge et al., 2005). Faria et al.
(2013) reported three rostra records from
private collections in Australia from
1998–1999, but no other information on
the collection location was reported.
Narrow sawfish have been reported in
multiple studies between 2000 and
2011, mostly from northern Australia. In
a bycatch reduction device study
conducted in 2001 in the Gulf of
Carpentaria, 25 narrow sawfish were
captured in trawling gear (Brewer et al.,
2006). Later in 2001, a bycatch
reduction device study conducted in the
Queensland shallow-water eastern king
prawn (Penaeus plebejus) trawl fishery
did not capture a single specimen
(Courtney et al., 2006). The European
Molecular Biology Lab recorded narrow
sawfish in 2003 in the Northern
Territory (GBIF database). A review of
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fisheries data and records from 2000 to
2002, identified 74 offshore and 37
inshore records of narrow sawfish in the
Gulf of Carpentaria (Peverell, 2005).
Between April 2004 and April 2005, 16
narrow sawfish were caught in the Gulf
of Carpentaria during a trawl bycatch
study; the mean catch rate was 0.16
sawfish per hour (Dell et al., 2009).
Observers on commercial fishing boats
recorded nine captures of narrow
sawfish in 2007 within the Great Barrier
Reef World Heritage Area, Queensland,
which accounted for 0.86 percent of the
shark and ray catch in the commercial
fisheries (Williams, 2007). Observers in
the Northern Territory’s Offshore Net
and Line Fishery encountered several
narrow sawfish from 2007 to 2010
(Davies, 2010). Data from the Kimberley
(R. McAuley, Department of Fisheries,
Western Australia, pers. comm. to Colin
Simpfendorfer, 2012), the Northern
Territory (Field et al., 2009), the Gulf of
Carpentaria (Peverell, 2005), and parts
of the Queensland east coast (Harry et
al., 2011) suggest viable subpopulations
may remain locally, but at significantly
lower levels compared to historic levels.
In summary, it appears the current
range of narrow sawfish is restricted
largely to Australia. Narrow sawfish are
considered very rare in many places
where evidence is available, including
parts of India (Roy, 2010), Bangladesh
(Roy, 2010), Burma (FIRMS, 2007–
2012), Malaysia (including Borneo;
Almada-Villela, 2002; Manjaji, 2002),
Indonesia (White and Kyne, 2010),
Thailand (CITES, 2007; Compagno,
2002a; Vidthayanon, 2002), and
Singapore (CITES, 2007). In Australia,
narrow sawfish are primarily located in
the north. The most recent museum
record for narrow sawfish in southern
Australia was from New South Wales in
the 1970s (Pogonoski et al., 2002). Data
from the Queensland Shark Control
Program, conducted along the east coast
of Queensland, from 1969 to 2003 show
a clear decline in sawfish catch
(although not species-specific) with the
complete disappearance of sawfish in
southern regions of Queensland by 1993
(Stevens et al., 2005). Although we
cannot rule out underreporting of
narrow sawfish, especially in remote
areas of its historic range, we conclude
from the consistent lack of records that
narrow sawfish have been severely
depleted in numbers and their range has
contracted.
Natural History of Dwarf Sawfish
(Pristis clavata)
Taxonomy and Morphology
Due to its size and the geographic
location where it was described, P.
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clavata is referred to as the dwarf or the
Queensland sawfish. The species was
first described by Garman in 1906;
however, it has often been confused
with largetooth sawfish (Last and
Stevens, 1994; Cook et al., 2006; Morgan
et al., 2010a). This species can be
distinguished from largetooth sawfish
based on rostral tooth morphology
(Thorburn et al., 2007).
The dwarf sawfish is olive brown in
color dorsally with a white underside.
The rostrum of this species is quite
short, with 19 to 23 rostral teeth that are
moderately flattened, elongated, and
peg-like. Studies indicate that this
species does not display significant
differences in the number of rostral
teeth between males (19 to 23 teeth) and
females (20 to 23 teeth) (Ishihara et al.,
1991a; Thorburn et al., 2008; Morgan et
al., 2010a; Morgan et al., 2011). The
rostrum makes up 21 to 26 percent of
the total length of the dwarf sawfish
(Blaber et al., 1989; Grant, 1991; Last
and Stevens, 1994; Compagno and Last,
1999; Larson et al., 2006; Wueringer et
al., 2009; Morgan et al., 2011).
Morphologically, the origin of the first
dorsal fin is slightly posterior to the
insertion of the pelvic fins, and the
second dorsal fin is smaller than the
first. The pectoral fins are small
compared to other sawfish species, and
are ‘‘poorly developed’’ (Ishihara et al.,
1991a). There is no lower lobe on the
caudal fin. Lateral and low keels are
present along the base of the tail
(Compagno and Last, 1999; Wueringer et
al., 2009; Morgan et al., 2010a; Morgan
et al., 2011). Within the mouth are 82–
84 tooth rows on the upper jaw. The
total vertebrae number is 225–231. The
dwarf sawfish has regularly overlapping
monocuspidate denticles on its skin. As
a result, there are no keels or furrows
formed on the skin (Fowler, 1941; Last
and Stevens, 1994; Deynat, 2005).
Habitat Use and Migration
The dwarf sawfish has been found
along tropical coasts in marine and
estuarine waters, mostly from northern
Australia; it may inhabit similar habitats
in other areas. Dwarf sawfish are
reported on mudflats in water 6 ft 7 in
to 9 ft 10 in (2 to 3 m) deep that is often
turbid and influenced heavily by tides.
Thorburn et al. (2008) reported dwarf
sawfish occur in waters 2 to 22 ft (0.7
to 7 m) deep, while Stevens et al. (2008)
recorded a maximum depth of 65 ft (20
m). This species has also been reported
in rivers (Last and Stevens, 1994;
Wueringer et al., 2009; Morgan et al.,
2010a) and as commonly occurring in
both brackish and freshwater, and in
both marine and estuarine habitats
(Rainboth, 1996; Thorburn et al., 2008).
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For example, two dwarf sawfish were
found 31 miles (50 km) upstream from
the mouth of the south Alligator River,
Kakadu National Park, Northern
Territory, Australia in 2013 at salinities
of 0.12 and 7.64 ppt (P. Kyne, Charles
Darwin University, pers. comm. to S.
Norton, NMFS, June 2013).
Juvenile dwarf sawfish may use the
estuaries associated with the Fitzroy
River, Australia as nursery habitat for
up to three years (Thorburn et al., 2008).
Dwarf sawfish are also known to use the
Gulf of Carpentaria, Australia as nursery
area in a variety of habitats (Gorham,
2006). However, physical characteristics
such as salinity, temperature, and
turbidity may limit seasonal movements
(Blaber et al., 1989).
Age and Growth
Dwarf sawfish are considered to be
small compared to other sawfishes.
Their maximum size has been reported
as 4 ft 11 in (1.5 m) total length (TL)
(Grant, 1991) and 4 ft 7 in (140 cm) TL
(Last and Stevens, 1994; Rainboth, 1996;
Compagno and Last, 1999). But more
recently, much larger sizes have been
reported, as high as 19.7 ft (6000 cm) TL
(Peverell, 2005). Specimens from
Western Australia in 2008 indicate that
females reach at least 10 ft 2 in (310 cm)
TL (Morgan et al., 2010a; Morgan et al.,
2011).
Thorburn et al. (2008) and Peverell
(2008) estimated age and growth for this
species based on the number of
vertebral rings and total length. The
average growth estimates for dwarf
sawfish are 16.1 in (41cm) TL in the first
year, slowing to 9.4 in (24 cm) in the
second year (Peverell 2008). Thorburn et
al. (2008) determined that animals close
to 3 ft (90 cm) TL were age 1, those
between 3.5 and 4 ft (110 cm and 120
cm) TL were age 2, and those around 5
ft (160 cm) TL were age 6. Peverell
(2008) reported dwarf sawfish between
2 ft 11 in and 3 ft 3 in (90 and 98 cm)
TL were age 0, those between 3 ft 7 in
and 5 ft 9 in (110 to 175 cm) TL were
considered 1 to 3 years old, and those
between 6 ft 7 in and 8 ft (201 to 244
cm) TL were considered 4 to 6 years old
(Peverell, 2008). Any dwarf sawfish over
9 ft 10 in (300 cm) TL is considered to
be at least 9 years old (Morgan et al.,
2010a). The theoretical maximum age
calculated from von Bertalanffy
parameters for dwarf sawfish is 94 years
(Peverell, 2008).
Reproduction
There is little information available
regarding the time or location of dwarf
sawfish mating. It is hypothesized that
dwarf sawfish move into estuarine or
fresh waters to breed during the wet
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season (Larson et al., 2006), although no
information on pupping habitat,
gestation period, or litter size has been
recorded (Morgan et al., 2010a).
Dwarf sawfish are born between 2 ft
2 in and 2 ft 8 in (65 cm and 81 cm)
TL (Morgan et al., 2010a; Morgan et al.,
2011). Males become sexually mature
between 9 ft 8 in and 10 ft (295 and 306
cm) TL with fully calcified claspers,
though they may mature at smaller
sizes, around 8 ft 5 in (255–260 cm) TL
(Peverell, 2005; Thorburn et al., 2008;
Last and Stevens, 2009; Morgan et al.,
2011). All males captured by Thorburn
et al. (2008) less than 7 ft 5 in (226 cm)
TL were immature; two females, both
smaller than 3 ft 11 in (120 cm) TL,
were also immature. There is little
specific information about sexual
maturation of females; females are
considered immature at 6 ft 11 in (210
cm) TL (Peverell, 2005; Peverell, 2008;
Morgan et al., 2010a). Wueringer et al.
(2009) indicates that neither males nor
females are mature before 7 ft 8 in (233
cm) TL.
Intrinsic rates of population increase,
based on life history data from Peverell
(2008), has been estimated to be about
0.10 per year (Moreno Iturria, 2012),
with a potential population doubling
time of 7.2 years.
Diet and Feeding
Dwarf sawfish, like other sawfishes,
use their saw to stun small schooling
fishes. They may also use the saw for
rooting in the mud and sand for
crustaceans and mollusks (Breder Jr.,
1952; Raje and Joshi, 2003; Larson et al.,
2006; Last and Stevens, 2009). In
Western Australia, the dwarf sawfish
eats shrimp (Natantia spp.), mullet
(Mugilidae), herring (Clupeidae), and
croaker (Sciaenidae) (Thorburn et al.,
2008; Morgan et al., 2010a).
Population Structure
Phillips et al. (2011) conducted a
genetic study looking at mtDNA of
dwarf sawfish and found no distinct
difference in dwarf sawfish from
Western Australia and those from the
Gulf of Carpentaria in northern
Australia. The genetic diversity of this
species was moderate overall; however,
dwarf sawfish from the Gulf of
Carpentaria may have a lower genetic
diversity than those of the west coast,
possibly due to either a small sample
size or a reduction in abundance
(Phillips et al., 2008). Further declines
in abundance as well as genetic drift
may result in reduced genetic diversity
(Morgan et al., 2010a; 2011).
Phillips et al. (2011) determined the
populations of the dwarf sawfish are
organized matrilineally (from mother to
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daughter), indicating the possibility that
females are philopatric (return to their
birth place). While the genetic diversity
of this species is considered low to
moderate across Australia, haplotype
diversity in the Gulf of Carpentaria was
very low, but was greater in the west
compared to the east. Low diversity
among and within groups of dwarf
sawfish may be detrimental (Phillips et
al., 2011).
Distribution and Abundance
Dwarf sawfish are thought to
historically occur in the Indo-Pacific,
western Pacific, and eastern Indian
Oceans, with the population largely
occurring in northern Australia (Last
and Stevens, 1994; Last and Compagno,
2002; Compagno, 2002a; Compagno,
2002b; Thorburn et al., 2008; Wueringer
et al., 2009; Morgan et al., 2010a; Kyne
et al., 2013). While dwarf sawfish may
have been historically more widespread
throughout the Indo-West Pacific
(Compagno and Last, 1999; Last and
Stevens, 2009), there are questions
regarding records outside of Australian
waters (DSEWPaC 2011; Kyne et al.,
2013; GBIF database).
In an effort to gather more information
on the species’ historic and current
range and abundance, we conducted an
extensive search of peer-reviewed
publications and technical reports,
newspaper, and magazine articles. We
also reviewed records from the Global
Biodiversity Information Facility (GBIF)
Database (www.gbif.com). A summary of
those findings is presented by major
geographic region.
Indian Ocean
Dwarf sawfish are considered
extremely rare in the Indian Ocean and
there are few records indicating its
current presence (Last, 2002). Faria et al.
´
(2013) report a female from the Reunion
Islands, a female from an unidentified
location in the Indian Ocean, and a
museum record of a male from Bay of
Bengal, India. A sawfish was landed at
a port in Arabian Peninsula (presumably
caught in the Gulf of Oman or the
Arabian Gulf) in January of 2006. It may
have been a dwarf sawfish, but
identification could not be confirmed
(Kyne et al., 2013). There are no reports
of dwarf sawfish from Sri Lanka in more
than a decade, although they have been
assumed to occur there (Last, 2002).
Indo-Pacific (excluding Australia)
Dwarf sawfish are considered very
rare in Indonesia, with only a few
records (Last, 2002). Faria et al. (2013)
compiled most reports of dwarf sawfish
in Indonesia; since the first record in
1894 from Borneo, there have been two
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rostral saws in 1910 and five other
rostra without date or length
information. There is also one museum
record of a dwarf sawfish from Papua
New Guinea in 1828 (Kyne et al., 2013).
Although reported historically, dwarf
sawfish have not been found in any
other areas in the Indo-Pacific in over a
decade. Rainboth’s (1996) guide to
fishes of the Mekong reported a dwarf
sawfish from the Mekong River Basin,
Laos, in the early 1900s but no
specimen exists to confirm this report.
No sawfish of any species, including the
dwarf sawfish, were reported from the
South China Sea from 1923–1996
(Compagno, 2002a). Faria et al. (2013)
reported on two specimens from the
Pacific Ocean, but no specifics were
provided.
Australia
The northern coast of Australia
represents the geographic center of
dwarf sawfish range that extends from
Cape York, Queensland west to the
Pilbara area in Western Australia
(Compagno and Last, 1999; Last and
Stevens, 2009; Kyne et al., 2013). Dwarf
sawfish may have occurred as far south
as Cairns, but reports are lacking. Most
records for dwarf sawfish are from the
north and northwest areas of Australia.
The earliest record of dwarf sawfish
in Australia is from 1877, but no
specific location was recorded (Faria et
al., 2013). A single rostrum from a dwarf
sawfish was found in 1916, but no other
information was recorded. In 1945, a
single specimen was reported from the
Northern Territory, Australia (Stevens et
al., 2005). There is a single record of a
dwarf sawfish from the Victoria River in
1964 that is currently housed at the
Museum Victoria (GBIF Database).
Five female and five male dwarf
sawfish (32 to 55 in; 82 to 140 cm TL)
were captured in 1990 in the Pentecost
River using gillnets (Taniuchi and
Shimizu, 1991; Taniuchi, 2002). CSIRO
recorded five dwarf sawfish in Western
Australia in 1990 (GBIF Database).
CSIRO also found one dwarf sawfish in
Walker Creek (a tributary of the Gulf of
Carpentaria) in 1991 (GBIF Database). In
1992, one specimen was found near
Darwin, Northern Territory, Australia
(GBIF Database). Between 1994 and
2010, almost 75 tissue samples were
taken from live dwarf sawfish or dried
rostra from the Gulf of Carpentaria and
the northwest coast of Australia
(Phillips et al., 2011). In 1997, two
specimens were collected near the
mouth of Buffalo Creek in Darwin,
Northern Territory (Chisholm and
Whittington, 2000). In 2005, Naylor et
al. (2005) collected one dwarf sawfish
from Darwin, Australia. One dwarf
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sawfish was captured in 1998 in the
upper reaches of the Keep River Estuary
(Larson, 1999; Gunn et al., 2010). CSIRO
reported one dwarf sawfish in Western
Australia (GBIF Database). In 2006, the
European Molecular Biology Lab
reported the occurrence of three dwarf
sawfish in Western Australia (GBIF
Database). One interaction was reported
between 2007 and 2010 by observers in
the Northern Territory Offshore Net and
Line Fishery (Davies, 2010). A single
specimen from Queensland
(northeastern Australia) is preserved at
the Harvard Museum of Comparative
Zoology (Fowler, 1941).
In a comprehensive survey of the Gulf
of Carpentaria from 2001 to 2002
(Peverell, 2005; 2008), indicated dwarf
sawfish were concentrated in the west
where 12 males and 10 females were
captured. Most individuals caught in
the inshore fishery were immature
except for two mature males: 10 ft and
9 ft 8 in (306 cm and 296 cm) TL
(Peverell, 2005; 2008).
Within specific riverine basins in
northwestern Australia, dwarf sawfish
have been reported in various surveys.
Forty-four dwarf sawfish were captured
between October 2002 and July 2004, in
the King Sound and the Robison, May,
and Fitzroy Rivers (Thorburn et al.,
2008). Between 2001 and 2002, one
dwarf sawfish was caught at the mouth
of the Fitzroy River in Western Australia
(Morgan et al., 2004). Morgan et al.
(2011) acquired 109 rostra from dwarf
sawfish from the King Sound area that
were part of museum or personal
collections.
In summary, there is some uncertainty
in the species identification of historic
records of dwarf sawfish, however, it
appears the dwarf sawfish has become
extirpated from much of the IndoPacific region and from the eastern coast
of Australia. An October 2001 study on
the effectiveness of turtle-excluder
devices in the prawn trawl fishery in
Queensland, Australia, reported no
dwarf sawfish (Courtney et al., 2006).
Dwarf sawfish are now considered rare
in the Gulf of Carpentaria. It is likely the
Kimberley region and Pilbara region
(Western Australia) may be the last
remaining areas for dwarf sawfish (P.
Kyne, Charles Darwin University, pers.
comm. to IUCN, 2012).
Natural History of the Largetooth
Sawfish (Pristis pristis)
Taxonomy and Morphology
Many taxonomists have suggested
classification of largetooth sawfish into
a single circumtropical species given
common morphological features of
robust rostrum, origin of first dorsal fin
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anterior to origin of pelvic fins, and
presence of a caudal-fin lower lobe
¨
(Gunther, 1870; Garman, 1913; Fowler,
1936; Poll, 1951; Dingerkus, 1983;
´
Daget, 1984; Seret and McEachran,
1986; McEachran and Fechhelm, 1998;
Carvalho et al., 2007). The recent
analysis by Faria et al. (2013) used
mtDNA (mitochondrial
deoxyribonucleic acid) and
contemporary genetic analysis to argue
that the previously classified P. pristis,
P. microdon, and P. perotteti should
now be considered one species named
P. pristis. After reviewing Faria et al.
(2013) and consulting other sawfish
experts, we conclude, based on the best
available information, that P. pristis
applies to all the largetooth sawfishes
previously identified as P. pristis, P.
microdon, and P. perotteti.
The largetooth sawfish has a robust
rostrum, noticeably widening
posteriorly (width between the two
posterior-most rostral teeth is 1.7 to 2
times the width between the second
anterior-most rostral teeth). Rostral
tooth counts are between 14 and 23 per
side with grooves on the posterior
margin. The body is robust with the
origin of the first dorsal-fin anterior to
the origin of the pelvic fin; dorsal fins
are high and pointed with the height of
the second dorsal fin greater than the
first. The lower lobe of the caudal-fin is
small, but well-defined, with the lower
anterior margin about half as long as the
upper anterior margin (Wallace, 1967;
Taniuchi et al., 1991a; Last and Stevens,
1994; Compagno and Last, 1999; Deynat,
2005; Wueringer et al., 2009; Morgan et
al., 2010a; Morgan et al., 2010b; Morgan
et al., 2011). The largetooth sawfish has
buccopharyngeal denticles and regularly
overlapping monocuspidate dermal
denticles on its skin. The denticles are
present on both dorsal and ventral
portions of the body (Wallace, 1967;
Deynat, 2005). Within the mouth, there
are between 70 and 72 tooth rows on the
upper jaw, and 64 to 68 tooth rows on
the lower jaw. The number of vertebrae
is between 226 and 228 (Morgan et al.,
2010a). Coloration of the largetooth
sawfish is a reddish brown dorsally and
dull white ventrally (Fowler, 1941;
Wallace, 1967; Compagno et al., 1989;
Taniuchi et al., 1991a; Compagno and
Last, 1999; Chidlow, 2007).
Male and female largetooth sawfish
differ in the number of rostral teeth.
Using largetooth sawfish teeth collected
from Papua New Guinea and Australia,
Ishihara et al. (1991b) found males to
have an average of 21 rostral teeth on
the left and 22 on the right; females
averaged 19 rostral teeth on both the left
and the right side of the rostrum.
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Rostrum length can vary between males
and females (Wueringer et al., 2009).
Habitat Use and Migration
Largetooth sawfish are found in
coastal and inshore waters and are
considered euryhaline (Compagno et al.,
1989; Last and Stevens, 1994;
Compagno and Last, 1999; Chisholm
and Whittington, 2000; Last, 2002;
Compagno, 2002b; Peverell, 2005;
Peverell, 2008; Wueringer et al., 2009),
being found in salinities ranging from 0
to 40 ppt (Thorburn et al., 2007). The
species has been found far upriver, often
occupying freshwater lakes and pools;
they are associated with freshwater
more than any other sawfish species
(Last and Stevens, 1994; Rainboth, 1996;
Peter and Tan, 1997; Compagno and
Last, 1999; Larson, 1999). Largetooth
sawfish have even been observed in
isolated fresh water billabongs or pools
until floodwaters allow them to escape;
juveniles often use these areas for
multiple years as deepwater refuges
(Gorham, 2006; Thorburn et al., 2007;
Wueringer et al., 2009; Morgan et al.,
2010b). Similarly, largetooth sawfish
have been found in Lake Nicaragua in
depths up to 400 ft (122 m) and are
found in deeper holes, occupying
muddy or sandy bottoms (Thorson,
1982). Adults more often use marine
habitats than juveniles, and are typically
found in waters with salinity at 31 ppt
(Wueringer et al., 2009).
Despite the variety of habitats
occupied, females have been found to be
highly philopatric as indicated by
mtDNA studies, while males often
undergo long movements (Lack et al.,
2009; Phillips et al., 2009; Morgan et al.,
2010a; Morgan et al., 2010b; Morgan et
al., 2011). Largetooth sawfish occurred
from the Caribbean and Gulf of Mexico
south through Brazil, and in the United
States, largetooth sawfish were reported
in the Gulf of Mexico, mainly along the
Texas coast (NMFS, 2010a). Largetooth
sawfish were rarely reported in U.S.
waters and may have been long-distance
migrants from the Caribbean or Brazil
(Feldheim et al., 2011).
The physical characteristics of habitat
strongly influence the movements of,
and areas used by, largetooth sawfish.
Recruitment of neonate largetooth
sawfish was correlated with the rise in
water levels during the wet season in
Australia (Whitty et al., 2009). A study
of juvenile largetooth sawfish
movements in the Fitzroy River in
Australia found young-of-the-year using
extremely shallow areas (0 to 1 ft 7 in
or 0 to 0.49 m) up to 80 percent of the
time, mostly to avoid predators
(Thorburn et al., 2007). Juvenile and
adult largetooth sawfish also use rivers
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(Compagno, 2002b; Gorham, 2006) and
can be found in areas up to 248.5 miles
(400 km) upstream (Morgan et al., 2004;
Chidlow, 2007). The space used on a
day to day basis by largetooth sawfish
increases with body length (Whitty et
al., 2009).
Age and Growth
There are several age and growth
studies for the largetooth sawfish;
results vary due to differences in aging
techniques, data collection, or location.
In Australia, largetooth sawfish are
between 2 ft 6 in and 3 ft (76 and 91
cm) TL at birth, with females being
slightly smaller than males on average
(Chidlow, 2007; Morgan et al., 2011).
Thorson (1982) found pups at birth
average 2 ft 4.7 in to 2 ft 7.5 in (73–80
cm) TL, with a growth rate of 1 ft 2 in
to 1 ft 3 in (35–40) cm per year in Lake
Nicaragua (NMFS, 2010a; Kyne and
Feutry, 2013). Peverell (2008) found that
largetooth sawfish in the Indo-West
Pacific are born at 2 ft 4 in to 2 ft 11
in (72–90 cm) TL. Juveniles (age 1 to age
at maturity) range in size from 2 ft 6 in
to 9 ft (76 to 277 cm) TL (Morgan et al.,
2011).
Size at maturity in the Western
Atlantic is estimated to be around 9 ft
10 in (300 cm) TL for both sexes at
around age 8 (Lack et al., 2009; Morgan
et al., 2010a; Morgan et al., 2010b;
NMFS, 2010; Morgan et al., 2011; Kyne
and Feutry, 2013). Thorson (1982)
estimated age of maturity to be 10 years
at 9 ft 10 in (300 cm) TL in Lake
Nicaragua. Peverell (2008) estimated age
at maturity in the Gulf of Carpentaria to
be between 8 and 10 years. In the IndoPacific, males tend to mature earlier
than other regions (9 ft 2 in (280 cm))
TL (Kyne and Feutry, 2013). Generally,
males under 7 ft 7 in (230 cm) TL and
females under 8 ft 10 in (270 cm) TL are
considered immature (Whitty et al.,
2009; Wueringer et al., 2009).
The largest recorded length of a
largetooth sawfish is 22 ft 11 in (700 cm)
TL (Compagno et al., 1989. The largest
largetooth sawfish recorded in the
Kimberley, Queensland measured 21 ft
6 in (656 cm) TL (Compagno and Last,
1999). In other areas of Australia,
largetooth sawfish can reach up to 15 ft
(457 cm) and at least 11 ft 10 in (361 cm)
TL (Fowler, 1941; Chidlow, 2007; Gunn
et al., 2010). Thorson (1982) estimated
that largetooth sawfish in Lake
Nicaragua only reach a maximum size of
about 14 ft 1 in (430 cm) TL.
Age and growth for largetooth sawfish
has been estimated by Tanaka (1991)
who generated a von Bertalanffy growth
model for specimens collected from
Papua New Guinea and Australia. For
both sexes combined, the theoretical
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maximum size (L∞) from the von
Bertalanffy growth equation was
calculated at 11 ft 11 in (363 cm) TL
with a growth rate (K) of 0.066 per year.
Largetooth sawfish grow around 7 in (18
cm) in the first year and 4 in (10 cm)
by the tenth year (Tanaka, 1991).
Thorson (1982a) estimated an early
juvenile growth rate of 13–15 in (35 to
40 cm) per year and annual adult
growth rate of 1 in (4.4 cm) per year
based on largetooth from Lake
Nicaragua. Simpfendorfer (2000)
estimated the theoretical maximum size
of largetooth sawfish to be 14 ft 11 in
(456 cm) TL with a growth rate (Brody
growth coefficient K) of 0.089 per year
based on Thorson’s (1982) data from
Lake Nicaragua. Peverell (2008)
calculated that largetooth sawfish from
the Gulf of Carpentaria, Australia grow
1 ft 8.5 in (52 cm) in the first year and
7 in (17 cm) during the fifth year.
Maximum size was estimated at 20 ft 11
in (638 cm) TL with a growth rate
(Brody growth coefficient K) of 0.08 per
year from the von Bertalanffy equation
(Peverell, 2008). Kyne and Feutry (2013)
summarize maximum age estimates of
30 years in Lake Nicaragua and 35 years
in the Gulf of Carpentaria. Based on the
von Bertalanffy equation, growth slows
at about 35 years or 19 ft 10 in (606 cm)
TL (Kyne and Feutry, 2013).
Reproduction
Largetooth sawfish are thought to
reproduce in freshwater environments
(Compagno and Last, 1999; Last, 2002;
Compagno, 2002b; Martin, 2005;
Thorburn and Morgan, 2005; Compagno
et al., 2006b). Pupping seems to vary
across the range, occurring during the
wet season from May to July in the IndoPacific (Raje and Joshi, 2003), and from
October to December in the western
Atlantic and Lake Nicaragua (Thorson,
1976a; Kyne and Feutry, 2013).
The number of pups in a largetooth
sawfish litter varies by location,
possibly due to a number of factors. One
of the earliest reproductive studies on
largetooth sawfish by Thorson (1976a)
reported the litter sizes of 67 females
ranged between 1 to 13 pups and an
embryonic sex ratio for this species is
0.86 males for every 1 female. Average
number of pups is 7 (NMFS, 2010a;
Kyne and Feutry, 2013). Thorson
(1976a) also found that both ovaries
appeared to be functional, with the left
ovary producing more eggs. Estimates of
litter size from other studies in the IndoWest Pacific (e.g., Wilson, 1999; Moreno
Iturria, 2012; Peverell, 2005) cannot be
confirmed (Kyne and Feutry, 2013).
Length of gestation for largetooth
sawfish is approximately five months in
Lake Nicaragua, with a biennial
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reproduction cycle (Thorson 1976a;
NMFS 2010a; Kyne and Feutry, 2013).
In the Indo-West Pacific, largetooth
sawfish may reproduce every year
(Peverell, 2008).
Intrinsic rates of population growth
vary tremendously throughout the
species’ range. Simpfendorfer (2000)
estimated that the largetooth sawfish in
Lake Nicaragua had an intrinsic rate of
population growth of 0.05 to 0.07 per
year, with a potential population
doubling time of 10.3 to 13.6 years.
Using data from Australia, rates of
population increase for the Indo-Pacific
were estimated to be around 0.12 per
year (Moreno Iturria, 2012), with a
population doubling time of
approximately 5.8 years and a
generation time of 14.6 years. Data from
the western Atlantic Ocean indicate an
intrinsic rate of increase of 0.03 per
year, with a population doubling time of
23.3 years and a generation time of 17.2
years (Moreno Iturria, 2012). Annual
natural mortality for the western
Atlantic has been estimated at 0.07 to
0.16 (Simpfendorfer, 2000) and 0.14 to
0.15 per year (Moreno Iturria, 2012).
Diet and Feeding
Largetooth sawfish diet is
predominantly fish, but varies
depending on geographic area. Small
fishes including seer fish, mackerels,
ribbon fish, sciaenids, and pomfrets are
likely main diet items of largetooth
sawfish in the Indian Ocean (Devadoss,
1978; Rainboth, 1996; Raje and Joshi,
2003). Small sharks, mollusks, and
crustaceans are also potential prey items
(Devadoss, 1978; Rainboth, 1996; Raje
and Joshi, 2003). Taniuchi et al. (1991a)
found small fishes and shrimp in the
stomachs of juveniles in Lake Murray,
Papua New Guinea, while juveniles in
Western Australia had catfish, cherabin,
mollusks, and insect parts in their
stomachs (Thorburn et al., 2007; Whitty
et al., 2009; Morgan et al,. 2010a).
Largetooth sawfish have also been found
to feed on catfish, shrimp, croaker,
small crustaceans, croaker, and
mollusks (Chidlow, 2007; Thorburn et
al., 2007; Morgan et al., 2010a; Morgan
et al., 2010b). Largetooth sawfish
captured off South Africa had bony fish
and shellfish as common diet items
(Compagno et al., 1989; Compagno and
Last, 1999). In general, largetooth
sawfish subsist on the most abundant
small schooling fishes in the area
(NMFS, 2010a).
Population Structure
Genetic analyses based on specific
sequences of mitochondrial DNA
indicated largetooth sawfish can be
found in populations based on ocean
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basin: Atlantic, Indo-West Pacific, and
Eastern Pacific. There is also restricted
flow of genes in largetooth sawfish
between these geographic areas: Atlantic
and Indo-West Pacific; Atlantic and
eastern Pacific; and Indo-West Pacific
and eastern Pacific (Faria et al. 2013).
Genetic analyses based on a 480-base
pair sequencing of the mtDNA gene
NADH–2 sequence also revealed
information indicating largetooth
sawfish subpopulations. West and East
Atlantic subpopulations differed as did
samples from Australia and the wider
Indian Ocean. Collectively, a total of 19
haplotypes were identified across
largetooth sawfish: One east Pacific
haplotype, 12 western Atlantic
haplotypes, two eastern Atlantic
haplotypes, one Indian Ocean
haplotype, one Vietnamese-New
Guinean haplotype, and two Australian
haplotypes (Faria et al., 2013). This finescale structuring by haplotypes was
only partially corroborated by the
regional variation in the number of
rostral teeth. While the rostral tooth
count differed significantly in largetooth
sawfish collected from the western and
eastern Atlantic Ocean, it did not vary
significantly between specimens
collected from the Indian Ocean and
western Pacific (Faria et al., 2013).
Largetooth sawfish collected from the
western Atlantic specimens had a
higher rostral teeth count than those
collected from the eastern Atlantic. Data
from separate protein and genetics
studies indicates some evidence of
distinction among populations of
largetooth sawfish in the Indo-Pacific.
At a broad scale, Watabe (1991) found
that there was limited genetic variability
between samples taken from Australia
and Papua New Guinea based on lactate
dehydrogenase (LDH) isozyme patterns.
Largetooth sawfish might be genetically
subdivided within the Gulf of
Carpentaria, Australia, with both eastern
and western Gulf populations (Lack et
al., 2009).
Phillips et al. (2011) found that the
population of largetooth sawfish in the
Gulf of Carpentaria is different from
animals on the west coast of Australia
(Fitzroy River) based on mtDNA. Recent
data (Phillips, 2012) suggests that
matrilineal structuring is found at
relatively small spatial scales within the
Gulf of Carpentaria region (i.e., this
region contains more than one maternal
‘population’), although the precise
location and nature of population
boundaries are unknown. The difference
in the genetic structuring using markers
with different modes of inheritance
(maternal versus bi-parental) suggests
that largetooth sawfish may have malebiased dispersal and females remaining
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at, or returning to, their birth place to
mate (Phillips et al., 2009; Phillips,
2012). Phillips (2012) noted that the
presence of male gene flow between
populations in Australian waters
suggests that a decline of males in one
location could affect the abundance and
genetic diversity of assemblages in other
locations.
The genetic diversity for largetooth
sawfish throughout Australia seems to
be low to moderate. Genetic diversity
was greater in the Gulf of Carpentaria
than in Australian rivers, also
suggesting potential philopatry:
Animals return to or stay in their home
range (Lack et al., 2009). Yet, given
limited sampling, additional research is
needed to better understand potential
population structure of largetooth
sawfish in Australia (Lack et al., 2009;
Phillips et al., 2009; Morgan et al.,
2010a; Morgan et al., 2010b).
Distribution and Abundance
Largetooth sawfish have the largest
historical range of all sawfishes. The
species historically occurred throughout
the Indo-Pacific near Southeast Asia and
Australia and throughout the Indian
Ocean to east Africa. Older literature
notes the presence of this species in
Zanzibar, Madagascar, India, and the
southwest Pacific (Fowler, 1941;
Wallace, 1967; Taniuchi et al., 2003).
Largetooth sawfish have also been noted
in the Eastern Pacific Ocean from
Mexico to Ecuador (Cook et al., 2005) or
possibly Peru (Chirichigno and Cornejo,
2001). In the Atlantic Ocean, largetooth
sawfish inhabit warm temperate to
tropical marine waters from Brazil to the
Gulf of Mexico in the western Atlantic,
and Namibia to Mauritania in the
eastern Atlantic (Burgess et al., 2009).
Given the recent taxonomic changes
for largetooth sawfish, we examined all
current and historic records of P.
microdon, P. perotteti, and P. pristis for
a comprehensive overview on
distribution and abundance. We
conducted an extensive search of peerreviewed publications and technical
reports, newspaper, records from the
GBIF Database, and magazine articles.
The results of that search are
summarized below by major geographic
region.
Indian Ocean
Largetooth sawfish historically
occurred throughout the Indian Ocean;
however, current records are rare for
many areas. The earliest record of
largetooth sawfish was in 1936 from
Grand Lac near the Gulf of Aden, Indian
Ocean (Kottelat, 1985). A second record
in 1936 is from the Mangoky River,
Madagascar (Taniuchi et al., 2003).
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Records from the 1960s and 1970s are
largely from India and South Africa.
One largetooth sawfish was reported
from the confluence of the Lundi and
Sabi Rivers, South Africa in 1960, over
200 miles (mi) inland (Jubb, 1967).
Between 1964 and 1966, several
largetooth sawfish were caught in the
Zambesi River, South Africa during a
general survey of rays and skates;
largetooth sawfish have also been
recorded in the shark nets off Durban,
South Africa (Wallace, 1967). In 1966, a
male (10 ft; 305 cm TL) was captured in
a trawl net in the Gulf of Mannar, Sri
Lanka (Gunn et al., 2010). Largetooth
sawfish were commonly caught between
1973 and 1974 in the Bay of Bengal
during the wet season (July and
September) but rarely during other
times of the year. Largetooth sawfish
were also reported in three major rivers
that empty into the Bay of Bengal: The
Pennaiyar, Paravanar, and Gadilam
(Devadoss, 1978).
Current reports of largetooth sawfish
throughout the Indian Ocean are
isolated and rare. Largetooth sawfish
were recorded in South Africa 1992 and
1993 between Nelson Mandela Bay and
Cape Town. Eight additional
observations are reported in South
Africa but associated date information
was not included (GBIF database).
While the species could not be
confirmed, a survey of fishing landing
sites and interviews with 99 fishers in
Kenya by Nyingi found 71 reports of
sawfishes over the last 40 years
(unpublished report from Dorothy
Wanja Nyingi to J. Carlson, NMFS,
2007). The longest time series of
largetooth sawfish catches is from the
swimmer protection beach nets off
Natal, South Africa with a yearly
average capture rate of 0.2 sawfish per
0.6 mi (1 km) net per year from 1981 to
1990; since then only two specimens
have been caught (CITES, 2007).
Largetooth sawfish were reported in
Cochin, India by the Central Marine
Fisheries Research Institute in 1994, but
no information about location, size, or
number of animals is available (Dan et
al., 1994). Commercial landings of
elasmobranchs from 1981 to 2000 in the
Bay of Bengal were mostly rays with
some largetooth sawfish (Raje and Joshi,
2003). In the Betsiboka River,
Madagascar, four largetooth sawfish
were caught in 2001. The most recent
capture of a largetooth sawfish (18 ft;
550 cm TL) in India occurred on January
18, 2011, between Karnataka and Goa
(www.mangalorean.com).
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Indo-Pacific Ocean (Excluding
Australia)
Many islands within the Indo-Pacific
region contain suitable habitat for
largetooth sawfish, but few reports are
available, perhaps due to the lack of
surveys or data reporting. The earliest
records of largetooth sawfish from the
Indo-Pacific are from a compilation
study of elasmobranchs in the waters off
Thailand that reports a largetooth
sawfish in the Chao Phraya River and its
tributaries in 1945 (Vidthayanon, 2002).
In 1955, two largetooth sawfish were
captured from Lake Sentani (present day
Intan Jaya, Indonesia). Juvenile
largetooth sawfish have also been
reported around the same time in a
freshwater river close to Genjem,
Indonesia (Boeseman, 1956). In 1956,
largetooth sawfish were recorded in
Lake Sentani (present day Intan Jaya,
Indonesia), (Boeseman, 1956; Thorson et
al., 1966). In a study by Munro (1967)
in the Laloki River in the southeastern
portion of New Guinea, no sawfish were
captured. From 1967 to 1977, five
largetooth sawfish were captured from
the Indragiri River, Sumatra (Taniuchi,
2002). The presence of largetooth
sawfish in the Mahakam River, Borneo
was recorded in 1987 (Christensen,
1992). Three largetooth sawfish rostra
were acquired from local fish markets in
Sabah in 1996 (Manjaji, 2002a).
Additional surveys of local fish markets
indicate largetooth sawfish are still
present in these areas, although locals
have noticed a decline in their
abundance (Manjaji, 2002a). In 1996,
two specimens were found in Malaysia:
One in Palau Nangka and one in Palau
Besar (GBIF Database).
Multiple records of largetooth sawfish
have occurred in areas throughout
Papua New Guinea. From 1970 to 1971,
Berra et al. (1975) collected five
largetooth sawfish from the Laloki
River, Papua New Guinea. Four
largetooth sawfish were recorded in
1975 from the Fly River system, Papua
New Guinea and one in 1979 in the
northern part of Papua New Guinea near
new Tangu (GBIF Database). In a survey
of the Fly River system, Papua New
Guinea, 23 individuals were captured in
1978 (Roberts, 1978; Taniuchi and
Shimizu, 1991; Taniuchi et al., 1991b;
Taniuchi, 2002). There are two reports
of largetooth sawfish in the 1980s in
Papua New Guinea: One in 1987 and
one in 1988 (GBIF Database). More
recently, 36 largetooth sawfish were
captured in September 1989 in Papua
New Guinea (Taniuchi and Shimizu,
1991; Taniuchi, 2002).
The scarcity of records from IndoPacific led to an increased effort to
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document species presence. Anecdotal
evidence suggests that largetooth
sawfishes have not been recorded in
Indo-Pacific for more than 25 years
(White and Last, 2010). Largetooth
sawfish have not been recorded in the
Mekong River, Laos for decades
(Rainboth, 1996). In a comprehensive
study compiled by Compagno (2002a),
no sawfishes were found in the South
China Sea between the years of 1923
and 1996. Data from 200 survey days at
fish landing sites in eastern Indonesia
between 2001 and 2005 recorded over
40,000 elasmobranchs, but only 2
largetooth sawfish (White and
Dharmadi, 2007; Kyne and Feutry,
2013).
Australia
Australia may have a higher
abundance of largetooth sawfish than
other areas within the species’ current
range (Thorburn and Morgan, 2005;
Field et al., 2009). Despite their current
abundance levels, we only identified a
few historic records from Australia. The
first record of a largetooth sawfish was
in 1945 in the Northern Territory
(Stevens et al., 2005). There was a
subsequent record in 1947, and two
largetooth sawfish from the Gulf of
Carpentaria, Queensland were reported
in 1959 (GBIF Database). Faria et al.
(2013) obtained a rostrum that was
collected in Australia in 1960.
Since the 1980s, we found
significantly more records of largetooth
sawfish in Australia than other regions.
A largetooth sawfish was captured from
the Keep River, Australia in 1981
(Compagno and Last, 1999). Three
largetooth sawfish were recorded in
1984 near Marchinbar Island, Northern
Territory (GBIF Database). Blaber et al.
(1990) found that largetooth sawfish
were among the top twenty-five most
abundant species in the trawl fisheries
of Albatross Bay from 1986 to 1988.
Three largetooth sawfish were reported
from the Gulf of Carpentaria,
Queensland: One in 1987 in Walker
Creek, one in 1988 in the Gilbert River,
and one in 1991 in Marrakai Creek, a
tributary of the Adelaide River,
Northern Territory (GBIF Database).
Eight individuals were captured in the
Leichhardt River in 2008 (Morgan et al.,
2010b). In a preliminary survey of the
McArthur River, Northern Territory,
Gorham (2006) reported two largetooth
sawfish captured between 2002 and
2006. Surveys (Peverell, 2005; Gill et al.,
2006; Peverell, 2008) in the Gulf of
Carpentaria found largetooth sawfish
widely distributed throughout the
eastern portion of the Gulf with most
catches occurring near the mouth of
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many rivers (Mitchell, et al., 2005;
2008).
Juvenile largetooth sawfish in
Australia use the Fitzroy River and
other tributaries of King Sound (Morgan
et al., 2004) as nursery areas while
adults are found more often offshore
(Morgan et al., 2010a). In Western
Australia, besides the Fitzroy River and
King Sound, the only other areas where
juvenile sawfish have been recently
recorded are in Willie Creek and
Roebuck Bay (Gill et al., 2006; Morgan
et al., 2011). Nursery areas for largetooth
sawfish are also reported in northern
Australia in the Gulf of Carpentaria
(Gorham, 2006). Juvenile largetooth
sawfish have been captured within the
Adelaide River, Australia in 2013 (P.
Kyne, Charles Darwin University, pers.
comm., 2013). Abundance estimates for
the largetooth sawfish from areas that
support higher human populations may
be declining (Taniuchi and Shimizu,
1991; Taniuchi et al., 1991a; Morgan et
al., 2010a). Whitty et al. (2009) found
that the population of juvenile
largetooth sawfish in the Fitzroy River
had declined; catch per unit effort was
56.7 sawfish per 100 hours in 2003
compared to 12.4 in 2009. There were
no reported captures of largetooth
sawfish in 2008 from the Roper River
system, which drains into the western
Gulf of Carpentaria, Northern Territory
(Dally and Larson, 2008). No adult
sawfish were captured in any of the
prawn trawl fisheries in Queensland,
Australia during the month of October
2001 (Courtney et al., 2006).
Outside the northern and western
areas of Australia, largetooth sawfish do
occur but reports are less frequent. In
southwestern Australian waters, one
female sawfish was captured by a
commercial shark fisherman in February
2003 east of Cape Naturaliste (Chidlow,
2007). Data from the Queensland,
Australia Shark Control Program shows
a clear decline in sawfish catch over a
30 year period from the 1960s, and the
complete disappearance of sawfish in
southern regions by 1993 (Stevens et al.,
2005).
Eastern Pacific
In the eastern Pacific, the historic
range of largetooth sawfish was from
Mazatlan, Mexico to Guayaquil, Ecuador
(Cook et al., 2005) or possibly Peru
(Chirichigno and Cornejo, 2001). There
is very little information on the
population status in this region and few
reports of capture records. The species
has been reported in freshwater in the
Tuyra, Culebra, Tilapa, Chucunaque,
Bayeno, and Rio Sambu Rivers, and at
the Balboa and Miraflores locks in the
Panama Canal, Panama; in Rio San Juan,
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Colombia; and in the Rio Goascoran,
along the border of El Salvador and
Honduras (Fowler, 1936, 1941; Beebe
and Tee-Van, 1941; Bigelow and
Schroeder, 1953; Thorson et al., 1966a;
Dahl, 1971; Thorson, 1974, 1976, 1982a,
1982b, 1987; Compagno and Cook, 1995;
all as cited in Cook et al., 2005). There
are 4 records of largetooth sawfish south
of Purto Vallarta, Mexico in 1975, and
several reports from Panama with no
associated dates (GBIF Database). The
only recent reports of largetooth sawfish
in this area are anecdotal reports from
Colombia, Nicaragua, and Panama (R.
Graham, Wildlife Conservation Society,
pers. comm. to IUCN, 2012).
Western Atlantic Ocean
In the western Atlantic Ocean,
largetooth sawfish were widely
distributed throughout the marine and
estuarine waters in tropical and
subtropical climates and historically
found from Brazil through the
Caribbean, Central America, the Gulf of
Mexico, and seasonally into waters of
the United States (Burgess et al., 2009).
Largetooth sawfish also occurred in
freshwater habitats in Central and South
America. Throughout the Caribbean Sea,
the historical presence of the largetooth
sawfish is uncertain and early records
might have been misidentified
smalltooth sawfish (G. Burgess, Florida
Museum of Natural History, pers.
comm. to IUCN, 2012).
Historic records of largetooth sawfish
in the western north Atlantic have been
previously reported in NMFS (2010a).
Sawfish were documented in Central
America in Nicaragua as early as 1529
by a Spanish chronicler (Gill and
Bransford, 1877). This species was also
historically reported in Nicaragua by
Meek (1907), Regan (1908), Marden
(1944), Bigelow and Schroeder (1953)
and Hagberg (1968). Five largetooth
sawfish were reported from a survey of
Lake Izabal, Guatemala from 1946 to
1947, and sawfishes were reported to be
important to inland fisheries (Saunders
et al., 1950). There is a single largetooth
sawfish report from Honduras, but the
true origin of the rostrum and the date
of capture could not be confirmed
(NMFS, 2010a).
In Atlantic drainages, largetooth
sawfish has been found in freshwater at
least 833 miles (1,340 km) from the
ocean in the Amazon River system
(Manacapuru, Brazil), as well as in Lake
Nicaragua and the San Juan River; the
Rio Coco, on the border of Nicaragua
and Honduras; Rio Patuca, Honduras;
Lago de Izabal, Rio Motagua, and Rio
Dulce, Guatemala; and the Belize River,
Belize. Largetooth sawfish are found in
Mexican streams that flow into the Gulf
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of Mexico; Las Lagunas Del Tortuguero,
Rio Parismina, Rio Pacuare, and Rio
Matina, Costa Rica; and the Rio San
Juan and the Magdalena River,
Colombia (Thorson, 1974, 1982b;
Castro-Augiree, 1978 as cited in
Thorson, 1982b; Compagno and Cook,
1995; C. Scharpf and M. McDavitt,
National Legal Research Group, Inc., as
cited in Cook et al., 2005).
In the United States, largetooth
sawfish were reported in the Gulf of
Mexico mainly along the Texas coast
east into Florida waters, though nearly
all records of largetooth sawfish
encountered in U.S. waters were limited
to the Texas coast (NMFS, 2010a).
Though reported in the United States, it
appears that largetooth sawfish were
never abundant, with approximately 39
confirmed records (33 in Texas) from
1910 through 1961.
The Amazon River basin and adjacent
waters are traditionally the most
abundant known range of largetooth
sawfish in Brazil (Bates, 1964; Marlier,
1967; Furneau, 1969). Most of the
records for which location is known
originated in the state of Amazonas,
which encompasses the middle section
of the Amazon River basin along with
the confluence of the Rio Negro and Rio
Solimoes Rivers. The other known
locations are from the states of Rio
Grande do Norte, Sergipe, Bahia,
Espirito Santo, Rio de Janeiro, Sao
Paulo, Para, and Maranhao (NMFS,
2010a). Most records of largetooth
sawfish in the Amazon River
(Amazonia) predate 1974. The
Magdalena River estuary was the
primary source for largetooth sawfish
encounters in Colombia from the 1940’s
(Miles, 1945), while other records
originated from the Bahia de Cartagena
and Isla de Salamanca (both marine),
and Rio Sinu (freshwater) from the
1960’s through the 1980’s (Dahl, 1964;
1971; Frank and Rodriguez, 1976;
Alvarez and Blanco, 1985). In other
areas of South America, there are only
single records from Guyana, French
Guiana, and Trinidad from the late
1800’s and early 1900’s. Of the 5 records
from Suriname, the most recent was
1962. Though thought to have once been
abundant in some areas of Venezuela
(Cervignon, 1966a, 1966b), the most
recent confirmed records of largetooth
sawfish from that country was in 1962.
Many records in the 1970’s and 1980’s
are largely due to Thorson’s (1982a,
1982b) research on the Lake NicaraguaRio San Juan system in Nicaragua and
Costa Rica. Bussing (2002) indicated
that this species was known to inhabit
the Rio Tempisque and tributaries of the
San Juan basin in Costa Rica. Following
Thorson’s (1982a, 1982b) studies,
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records of largetooth sawfish in the
western North Atlantic decline
considerably. By 1981, Thorson (1982a)
was unable to locate a single live
specimen in the original areas he
surveyed. There are no known
Nicaraguan records of the largetooth
sawfish outside of the Lake NicaraguaRio San Juan-Rio Colorado system
(Burgess et al., 2009), although
largetooth sawfish are still captured
incidentally by fishers netting for other
species (McDavitt, 2002). Of the known
largetooth sawfish reported from
Mexico, most records are prior to 1978
(NMFS, 2010a). Caribbean records are
very sparse (NMFS, 2010a). The last
record of a largetooth sawfish in U.S.
waters was in 1961 (Burgess et al.,
2009).
Most recent records for largetooth
sawfish are in isolated areas. While
many reports of largetooth sawfish from
Brazil were from the 1980’s and 1990’s
(Lessa, 1986; Martins-Juras et al., 1987;
Stride and Batista, 1992; Menni and
Lessa, 1998; and Lessa et al., 1999),
recent records indicate largetooth
sawfish are primarily found in fish
markets near the Amazon-Orinoco
estuaries (Charvet-Almeida, 2002;
Burgess et al., 2009). A Lake Nicaragua
fisherman reports he encounters a few
sawfish annually (McDavitt, 2002).
Other records are rare for the area. Three
recent occurrences were found in
Internet searches, one being a 200 lb.
(90.7 kg) specimen caught recreationally
in Costa Rica (Burgess et al., 2009).
Though reported by Thorson et al.
(1966a, 1966b) to be common
throughout the area, there are no recent
reports of encounters with sawfishes in
Guatemala. Scientists in Colombia have
not reported any sawfish sightings
between 1999 and 2009 (Burgess et al.,
2009).
Eastern Atlantic Ocean
Historic records indicate that
largetooth sawfish were once relatively
common in the coastal estuaries along
the west coast of Africa. Verified records
exist from Senegal (1841–1902), Gambia
(1885–1909), Guinea-Bissau (1912),
Republic of Guinea (1965), Sierra Leone
ˆ
(date unknown), Liberia (1927), Cote
d’Ivoire (1881–1923), Congo (1951–
1958), Democratic Republic of the
Congo (1951–1959), and Angola (1951).
Most records, however, lacked species
identification and locality data and may
have been confused taxonomically with
other species. Unpublished notes from a
1950’s survey detail 12 largetooth
sawfish from Mauritania, Senegal,
ˆ
Guinea, Cote d’Ivoire, and Nigeria,
ranging in size from 35–275 in (89–700
cm) TL (Burgess et al., 2009).
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A more recent status review by
Ballouard et al. (2006) reported that
sawfishes, including the largetooth
sawfish, were once common from
Mauritania to the Republic of Guinea,
but are now rarely captured or
encountered. According to this report,
the range of sawfishes has decreased to
the Bissagos Archipelago (Guinea
Bissau). The most recent sawfish
encounters outside Guinea Bissau were
in the 1990’s in Mauritania, Senegal,
Gambia, and the Republic of Guinea.
The most recent documented largetooth
sawfish capture was from 2005 in Nord
de Caravela (Guinea Bissau), along with
anecdotal accounts from fishers of
captures off of two islands in the same
area in 2008 (Burgess et al., 2009).
In summary, on a global scale,
largetooth sawfish appear to have been
severely fragmented throughout their
historic range into isolated populations
of low abundance. Largetooth sawfish
are now considered very rare in many
places where evidence is available,
including parts of East Africa, India,
parts of the Indo-Pacific region, Central
and South America and West Africa.
Even within areas like Australia and
Brazil, the species is primarily located
in remote areas. Information from
genetic studies indicates that largetooth
sawfish display strong sex-biased
dispersal patterns; with females
exhibiting patterns of natal philopatry
while males move more broadly
between populations (Phillips et al.,
2011). Thus, the opportunity for reestablishment of these isolated
populations is limited because any
reduction in female abundance in one
region is not likely to be replenished by
movement from another region
(Phillips, 2012).
Natural History of Green Sawfish
(Pristis zijsron)
Taxonomy and Morphology
Pristis zijsron (Bleeker, 1851) is
frequently known as the narrowsnout
sawfish or the green sawfish.
Synonymous names include P. dubius
(Gloerfelt-Tarp and Kailola, 1984; Van
Oijen et al., 2007; Wueringer et al.,
2009). An alternative spelling for this
species’ scientific name (P. zysron) is
found in older literature, due to either
inconsistent writing or errors in
translation or transcription (Van Oijen
et al., 2007).
The green sawfish has a narrow saw
with 25–32 small, slender rostral teeth;
tooth count may vary geographically
(Marichamy, 1969; Last and Stevens,
1994; Morgan et al., 2010a). Specimens
collected along the west coast of
Australia have 24–30 left rostral teeth
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73989
and 23–30 right rostral teeth (Morgan et
al., 2010a), although other reports are
23–34 (Morgan et al., 2011). There have
been no studies to determine sexual
dimorphism from rostral tooth counts
for green sawfish. The rostral teeth are
generally denser near the base of the
saw than at the apical part of the saw
(Blegvad and Loppenthin, 1944). The
total rostrum length is between 20.6–
29.3 percent of the total length of the
animal and may vary based on the
number and size of individuals. In
general, green sawfish have a greater
rostrum length to total length ratio than
other sawfish species (Morgan et al.,
2010a, 2011).
In terms of body morphology, the
origin of the first dorsal fin on green
sawfish is slightly posterior to the origin
of pelvic fins. The lower caudal lobe is
not well defined and there is no
subterminal notch (Gloerfelt-Tarp and
Kailola, 1984; Compagno et al., 1989;
Last and Stevens, 1994; Compagno and
Last, 1999; Bonfil and Abdallah, 2004;
Wueringer et al., 2009; Morgan et al.,
2010a; Morgan et al., 2011). The green
sawfish has limited buccopharyngeal
denticles and regularly overlapping
monocuspidate dermal denticles on its
skin. As a result, there are no keels or
furrows formed on the skin (Deynat,
2005). The green sawfish is greenish
brown dorsally and white ventrally.
This species might be confused with the
dwarf or smalltooth sawfish due to its
similar size and range (Compagno et al.,
2006c).
Habitat Use and Migration
The green sawfish mostly uses
inshore, marine habitats, but it has been
found in freshwater environments
(Gloerfelt-Tarp and Kailola, 1984;
Compagno et al., 1989; Compagno,
2002b; Stevens et al., 2008; Wueringer
et al., 2009). In the Gilbert and Walsh
Rivers of Queensland, Australia,
specimens have been captured as far as
149 miles (240 km) upriver (Grant,
1991). However, Morgan et al. (2010a,
2011) report green sawfish do not move
into freshwater for any portion of their
lifecycle. Like most sawfishes, the green
sawfish prefers muddy bottoms in
estuarine environments (Last, 2002).
The maximum depth recorded for this
species is 131 ft (40 m) but it is often
found in much shallower waters,
around 16 ft (5 m; Compagno and Last,
1999; Wueringer et al., 2009). Adults
tend to spend more time in offshore
waters in Australia, as indicated by
interactions with the offshore Pilbara
Fish Trawl Fishery, while juveniles
prefer protected, inshore waters
(Morgan et al., 2010a; Morgan et al.,
2011).
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Age and Growth
At birth pups are between 2 ft and 2
ft 7 in (61 and 80 cm) TL. At age 1 green
sawfish are generally around 4 ft 3 in
(130 cm) TL (Morgan et al., 2010a).
Peverell (2008) found between ages 1
and 5, green sawfish measure between
4 ft 2 in and 8 ft 5 in (128 and 257 cm)
TL, based on the vertebral analysis of 6
individuals (Peverell, 2008; Morgan et
al., 2010a; Morgan et al., 2011). A 12 ft
6 in (380 cm) TL green sawfish was
found to be age 8, a 14 ft 4 in (438 cm)
TL individual was found to be age 10,
a 14 ft 9 in (449 cm) TL specimen was
found to be age 16, and a 15 ft (482 cm)
TL specimen was found to be age 18
(Peverell, 2008; Morgan et al., 2011).
Adult green sawfish often reach 16 ft
5 in (5 m) TL, but may grow as large as
23 ft (7 m) TL (Compagno et al., 1989;
Grant, 1991; Last and Stevens, 1994;
Compagno and Last, 1999; Bonfil and
Abdallah, 2004; Compagno et al., 2006c;
Morgan et al., 2010a). The largest green
sawfish collected in Australia was
estimated to be 19 ft 8 in (600 cm) TL
based on a rostrum length of 5 ft 5 in
(165.5 cm; Morgan et al., 2010a; Morgan
et al., 2011).
Peverell (2008) completed an age and
growth study for green sawfish using
vertebral growth bands. Von Bertalanffy
growth model parameters from both
sexes combined resulted in estimated
maximum theoretical size of 16 ft (482
cm) TL, relative growth rate of 0.12 per
year and theoretical time at zero length
of 1.12 yrs. The theoretical maximum
age for this species is calculated to be
53 years (Peverell, 2008; Morgan et al.,
2010a).
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Reproduction
Last and Stevens (2009) reported size
at maturity for green sawfish at 9 ft 10
in (300 cm) TL, corresponding to age 9.
In contrast, Peverell (2008) reported one
mature individual of 12 ft 4 in (380 cm)
TL and estimated its age as 9 yrs. Using
the growth function from Peverell
(2008) and assuming length of maturity
at 118 in (300 cm), Moreno Iturria
(2012) determined maturation is likely
to occur at age 5. Demographic models
based on life history data from the Gulf
of Carpentaria indicate the generation
time is 14.6 years, the intrinsic rate of
population increase is 0.02 per year, and
population doubling time is
approximately 28 years (Moreno Iturria,
2012).
Green sawfish give birth to as many
as 12 pups during the wet season
(January through July); Last and
Stevens, 1994; Peverell, 2008; Morgan et
al., 2010a, 2011). In Western Australia,
females are known to pup in areas
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between One Arm Point and Whim
Creek, with limited data for all other
areas (Morgan et al., 2010a; Morgan et
al., 2011). The Gulf of Carpentaria,
Australia is also a known nursery area
for green sawfish (Gorham, 2006). It is
not known where the green sawfish
breed or their length of gestation.
Diet and Feeding
Like other sawfish, green sawfish use
their rostra to stun small, schooling
fishes, such as mullet, or use it to dig
up benthic prey, including mollusks
and crustaceans (Breder Jr., 1952;
Rainboth, 1996; Raje and Joshi, 2003;
Compagno et al., 2006c; Last and
Stevens, 2009). One specimen captured
in 1967 in the Indian Ocean had jacks
and razor fish (Caranx and Centriscus)
species in its stomach (Marichamy,
1969). In Australia, the diet of this
species often includes shrimp, croaker,
salmon, glassfish, grunter, and ponyfish
(Morgan et al., 2010a).
Population Structure
Faria et al. (2013) found no global
population structure for green sawfish
in their genetic studies. However,
geographical variation was found in the
number of rostral teeth per side,
suggesting some population structure
may occur. Green sawfish from the
Indian Ocean have a higher number of
rostral teeth per side than those from
western Pacific specimens (Faria et al.,
2013).
In Australia, genetic analysis found
differences in green sawfish between the
west coast, the east coast, and the Gulf
of Carpentaria (Phillips et al., 2011).
Genetic data suggests these populations
are structured matrilineally (from the
mother to daughter) but there is no
information on male gene flow at this
time. These results may be indicative of
philopatry where adult females return to
or remain in the same area they were
born (Morgan et al., 2010a; Morgan et
al., 2011; Phillips et al., 2011). Phillips
et al. (2011) also found low levels of
genetic diversity for green sawfish in the
Gulf of Carpentaria, suggesting the
population may have undergone a
genetic bottleneck.
Distribution and Abundance
The green sawfish historically ranged
throughout the Indo-West Pacific from
South Africa northward along the east
coast of Africa, through the Red Sea,
Persian Gulf, Southern Asia, IndoAustralian archipelago, and east to Asia
as far north as Taiwan and Southern
China (Fowler, 1941; Blegvad and
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17:42 Dec 11, 2014
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sawfish in Australia: two from the
Northern Territory, and one from the
Gulf of Carpentaria (GBIF Database). A
green sawfish was observed in the Gulf
of Carpentaria in 1981 by CSIRO. Two
were observed in Western Australia, one
in 1982 and one in 1983 (GBIF
Database). Two green sawfish were
captured from Balgal, Queensland,
Australia in 1985 (Beveridge and
Campbell, 2005). In the Gulf of
Carpentaria, two green sawfish were
recorded in 1986, and one was recorded
in 1987 (GBIF Database).
One green sawfish was caught in the
southern portion of the Gulf of
Carpentaria in late 1990 during a fish
fauna survey (Blaber et al., 1994).
Alexander (1991) captured a female
green sawfish from the west coast of
Australia that was used for a
morphological study. Between 1994 and
2010, almost 50 tissue samples were
taken from live green sawfish or dried
rostra from multiple areas around
Australia, primarily the Gulf of
Carpentaria and northwest and
northeast coasts (Phillips et al., 2011). In
1997, one green sawfish was found at
the mouth of Buffalo Creek near Darwin,
Northern Territory (Chisholm and
Whittington, 2000). In a survey from
1999 through 2001 by White and Potter,
(2004), one green sawfish was captured
in Shark Bay, Queensland. In 1999, one
green sawfish was captured by CSIRO
from the Gulf of Carpentaria (GBIF
Database). Peverell (2005, 2008) noted
the green sawfish was one of the least
encountered species in a survey from
the Gulf of Carpentaria. In 2004, one
green sawfish was reported near
Darwin, Northern Territory by the
European Molecular Biology Lab (GBIF
Database). No green sawfish were
captured from the Roper River system in
2008, which drains into the western
Gulf of Carpentaria, Northern Territory
(Dally and Larson, 2008). Some records
have been reported for the east coast of
Australia; one female green sawfish was
acoustically tracked for 27 hours in May
2004 (Peverell and Pillans, 2004;
Porteous, 2004). Peverell (2005, 2008)
noted the green sawfish was one of the
least encountered species in a survey
from the Gulf of Carpentaria.
In summary, limited data makes it
difficult to determine the current range
and abundance of green sawfish.
Nonetheless, given the uniqueness (size
and physical characteristics) of the
sawfish, we believe the lack of records
in the areas where the species was
historically found indicates the species
is no longer present or has declined to
extremely low levels. Extensive surveys
at fish landing sites throughout
Indonesia since 2001 have failed to
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record the green sawfish (White pers.
comm. to IUCN, 2012). There is some
evidence from the Persian Gulf and Red
Sea (e.g., Sudan) of small but extant
populations (A. Moore, RSK
Environment Ltd., pers. comm. to IUCN,
2012). Green sawfish are currently
found primarily along the northern
coast of Australia, but all sawfish
species have undergone significant
declines in Australian waters. The
southern extent of the range of green
sawfishes in Australia has contracted
(Harry et al., 2011). Green sawfish have
been reported as far south as Sydney,
New South Wales, but are rarely found
as far south as Townsville, Queensland
(Porteous, 2004).
Natural History of the Non-Listed
Population(s) of Smalltooth Sawfish
(Pristis pectinata)
This section includes information
from the listed U.S. DPS of smalltooth
sawfish. The U.S. DPS of smalltooth
sawfish was listed as endangered on
April 1, 2003 (68 FR 15674). The basis
of the U.S. DPS smalltooth sawfish
listing was the significant differences in
management across international
borders. We discuss information from
the U.S. DPS of smalltooth sawfish here
because there is very little basic
biological information on smalltooth
sawfish found outside the U.S. We
believe the information from the U.S.
DPS is likely representative of the nonU.S. population of smalltooth sawfish
and is useful for understanding its
biology and extinction risk.
Taxonomy and Morphology
The smalltooth sawfish was first
described as Pristis pectinatus, Latham
1794. The name was changed to the
currently valid P. pectinata to match
gender of the genus and species as
required by the International Code of
Zoological Nomenclature.
The smalltooth sawfish has a thick
body with a moderately sized rostrum.
As with many other sawfishes, tooth
count varies by individual or region.
While there is no reported difference in
rostral tooth count between sexes, there
have been reports of sexual dimorphism
in tooth shape, with males having
broader teeth than females (Wueringer
et al., 2009). Rostral teeth are denser
near the apex of the saw than the base.
Most studies report a rostral tooth count
of 25 to 29 for smalltooth sawfish
(Wueringer et al., 2009). The saw may
constitute up to one-fourth of the total
body length (McEachran and De
Carvalho, 2002).
The pectoral fins are broad and long
with the origin of the first dorsal fin
over or anterior to the origin of the
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pelvic fins (Faria et al., 2013). The lower
caudal lobe is not well defined and
lacks a ventral lobe (Wallace, 1967;
Gloerfelt-Tarp and Kailola, 1984; Last
and Stevens, 1994; Compagno and Last,
1999; Bonfil and Abdallah, 2004;
Wueringer et al., 2009). This species has
between 228 and 232 vertebrae
(Wallace, 1967).
The smalltooth sawfish has
buccopharyngeal denticles and regularly
overlapping monocuspidate (singlepointed) dermal denticles on their skin.
As a result, there are no keels or furrows
formed on the skin (Last and Stevens,
1994; Deynat, 2005). The body is an
olive grey color dorsally, with a white
ventral surface (Compagno et al., 1989;
Last and Stevens, 1994; Compagno and
Last, 1999). This species may be
confused with the narrow or green
sawfish (Compagno, 2002b).
Habitat Use and Migration
All research on habitat use and
migration has been conducted on the
U.S. DPS of smalltooth sawfish. A
summary of recent information (NMFS,
2010b) indicates smalltooth sawfish are
generally found in shallow waters with
varying salinity level that are associated
with red mangroves (Rhizophora
mangle). Juvenile sawfish appear to
have small home ranges and limited
movements. Simpfendorfer et al. (2011)
reported smalltooth sawfish have an
affinity for salinities between 18 and at
least 24 ppt, suggesting movements are
likely made, in part, to remain within
this salinity range. Therefore, freshwater
flow may affect the location of
individuals within an estuary. Poulakis
et al. (2011) found juvenile smalltooth
sawfish had an affinity for water less
than 3 ft (1.0 m) deep, water
temperatures greater than 86 degrees
Fahrenheit (30 degrees Celsius),
dissolved oxygen greater than 6 mg per
liter, and salinity between 18 and 30
ppt. Greater catch rates for smalltooth
sawfish less than 1 year old were
associated with shoreline habitats with
overhanging vegetation such as
mangroves. Poulakis et al. (2012) further
determined daily activity space of
smalltooth sawfish is less than 1 mi (0.7
km) of river distance. Hollensead (2012)
reported smalltooth sawfish activity
areas ranged in size from 837 square
yards to 240,000 square yards to
approximately 3 million square yards
(0.0007 to 2.59 km2) with average range
of movements of 2.3 yards to 6.67 yards
(2.4 to 6.1 m) per minute. Hollensead
(2012) also found no difference in
activity area or range of movement
between ebb and flood, or high and low
tide. Smalltooth sawfish movements at
night suggest possible nocturnal
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foraging. Using a combination of data
from pop-off archival transmitting tags
across multiple institutional programs,
movements and habitat use of adult
smalltooth sawfish were determined in
southern Florida and the Bahamas
(Carlson et al., 2013). Smalltooth
sawfish generally remained in coastal
waters at shallow depths less than 32 ft;
(10 m) for more than 96 percent of the
time that they were monitored.
Smalltooth sawfish also remained in
warm water temperatures of 71.6 to 82.4
degrees Fahrenheit (22 to 28 degrees
Celsius) within the region where they
were initially tagged. Tagged smalltooth
sawfish traveled an average of 49 mi
(80.2 km) from deployment to pop-off
location during an average of 95 days.
No smalltooth sawfish tagged in U.S. or
Bahamian waters have been tracked to
countries outside where they were
tagged.
Age and Growth
There is no age and growth data for
smalltooth sawfish outside of the U.S.
DPS. A summary of age and growth data
on the U.S. DPS of smalltooth sawfish
(NMFS, 2010b) indicates rapid juvenile
growth for smalltooth sawfish for the
first two years after birth. Recently,
Scharer et al. (2012) counted bands on
sectioned vertebrae from naturally
deceased smalltooth sawfish and
estimated von Bertalanffy growth
parameters. Theoretical maximum size
was estimated at 14.7 ft (4.48 m),
relative growth was 0.219 per year, with
theoretical maximum size at 15.8 years.
Reproduction
In the eastern Atlantic Ocean,
smalltooth sawfish have been recorded
breeding in Richard’s Bay and St. Lucia,
South Africa (Wallace, 1967; Compagno
et al., 1989; Compagno and Last, 1999).
Pupping grounds are usually inshore, in
marine or fresh water. Pupping occurs
year-around in the tropics, but in only
spring and summer at higher latitudes
(Compagno and Last, 1999). Records of
captive breeding have been reported
from the Atlantis Paradise Island Resort
Aquarium in Nassau, Bahamas;
copulatory behavior was observed in
2003 and six months later the female
aborted the pups for unknown reasons
(McDavitt, 2006). In October 2012, a
female sawfish gave birth to five live
pups at the Atlantis Paradise Island
Resort Aquarium in Nassau, Bahamas (J.
Choromanski, Ripley’s Entertainment
pers. comm to NMFS, 2013).
Several studies have examined
demography of smalltooth sawfish in
U.S. waters. Moreno Iturria (2012)
calculated demographic parameters for
smalltooth sawfish in U.S. waters and
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estimated intrinsic rates of increase at
seven percent annually with a
population doubling time of 9.7 years.
However, preliminary results of a
different model by Carlson et al. (2012)
indicates population increase rates may
be greater, up to 17.6 percent annually,
for the U.S. population of smalltooth
sawfish. It is not clear which of these
models is more appropriate for the nonU.S. population of smalltooth sawfish.
Diet and Feeding
Smalltooth sawfish often use their
rostrum saw in a side-sweeping motion
to stun their prey, which may include
small fishes, or to dig up invertebrates
from the bottom (Breder Jr., 1952;
Compagno et al., 1989; Rainboth, 1996;
McEachran and De Carvalho, 2002; Raje
and Joshi, 2003; Last and Stevens, 2009;
Wueringer et al., 2009).
Population Structure
A qualitative examination of genetic
sequences revealed no geographical
structuring of smalltooth sawfish
haplotypes; however, variation in the
number of rostral teeth per side was
found in specimens from the western
and eastern Atlantic Ocean (Faria et al.,
2013).
Distribution and Abundance
Smalltooth sawfish were thought to be
historically found in South Africa,
Madagascar, the Red Sea, Arabia, India,
the Philippines, along the coast of West
Africa, portions of South America
including Brazil, Ecuador, the Caribbean
Sea, the Mexican Gulf of Mexico, as
well as Bermuda (Bigelow and
Scheroder, 1953; Wallace, 1967; Van der
Elst 1981; Compagno et al., 1989; Last
and Stevens, 1994; IUCN, 1996;
Compagno and Last, 1999; McEachran
and De Carvalho, 2002; Monte-Luna et
al., 2009; Wueringer et al., 2009). Yet,
reports of smalltooth sawfish from other
than the Atlantic Ocean are likely
misidentifications of other sawfish
(Faria et al., 2013). The lack of
confirmed reports of smalltooth sawfish
from areas other than the Atlantic Ocean
indicates that smalltooth sawfish are
only found in the Atlantic Ocean. In the
eastern Atlantic Ocean, smalltooth
sawfish were historically found along
the west coast of Africa from Angola to
Mauritania (Faria et al., 2013). Although
smalltooth sawfish were included in
historic faunal lists of species found in
the Mediterranean Sea (Serena, 2005), it
is still unclear if smalltooth sawfish
occurred as part of the Mediterranean
ichthyofauna or were only seasonal
migrants.
To evaluate the current and historic
distribution and abundance of the
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smalltooth sawfish outside the U.S.
DPS, we conducted an extensive search
of peer-reviewed publications and
technical reports, newspaper, records
from the GBIF Database, and magazine
articles. The results of that search are
summarized by major geographic region.
Africa, in 2003 (M. Diop, pers. comm. to
IUCN, 2012). Two other countries have
recently reported sawfish (Guinea
Bissau, Africa in 2011, and Mauritania
in 2010), but these reports did not
identify the species as smalltooth
sawfish.
Eastern Atlantic Ocean
Smalltooth sawfish were once
common in waters off the west coast of
Africa, but are now rarely reported or
documented in the area. The earliest
record of a smalltooth sawfish is a
specimen from Namibia in 1874 (GBIF
Database). Other records of smalltooth
sawfish in Africa occurred in 1907 from
Cameroon, five males and two females.
Female specimens were recorded in the
Republic of the Congo in 1911 and 1948.
Other reports from the Republic of
Congo include a male and two females,
but dates were not recorded. An
undated female specimen from
Mauritania was recorded (Faria et al.,
2013). A rostrum from Pointe Noire,
Molez, Republic of the Congo was found
in 1958 (Deynat, 2005; Faria et al.,
2013). There are records of smalltooth
sawfish from Senegal as early as 1956
and another rostral saw was recorded in
1959. Faria et al. (2013) also reports on
four other rostra from Senegal, but no
other information is available.
Many records of smalltooth sawfish
from the eastern Atlantic Ocean are
reported in the GBIF database during
the 1960s, particularly between 1963
and 1964. The majority of these records
are from Nigeria (118), but others are
from Gabon (77), Ghana (51), Cameroon
(43), and Liberia (39). Another online
database, Fishbase (www.fishbase.org),
has the same records. It is unclear if
these records are duplicative due to the
lack of specific information.
In the 1970s, records of smalltooth
sawfish became limited to more
northern areas of West Africa. One
rostral saw from Senegal was recorded
in 1975 (Alexander, 1991). Similarly,
one rostral saw was reported from
Gambia in 1977, but information about
exact location or sex of the animal was
absent (Faria et al., 2013). Faria et al.
(2013) report a record of smalltooth
sawfish in Guinea-Bissau in 1983 and a
record of a saw in 1987. For a
morphological study, Deynat (2005)
obtained a juvenile female from Cacheu,
Guinea-Bissau in 1983, and another
from Port-Etienne, Mauritania, in 1986.
Two rostra were reported from the
Republic of Guinea, one in 1980 and
one in 1988 (Faria et al., 2013).
In the last 10 years, there has been
only one confirmed record of a
smalltooth sawfish in the eastern
Atlantic Ocean in Sierra Leone, West
Western Atlantic Ocean (Outside U.S
Waters)
Overall, records of smalltooth sawfish
in the western Atlantic Ocean are scarce
and show a non-continuous range,
potentially due to misidentification
with largetooth sawfish. Faria et al.
(2013) summarized most records of
smalltooth sawfish in these areas. Faria
et al. (2013) report the earliest records
are a female smalltooth sawfish from
Haiti in 1831 and a female sawfish from
Trinidad and Tobago in 1876 (Faria et
al., 2013). One smalltooth sawfish was
´
recorded in Belem, Brazil in 1863 (GBIF
Database). Two smalltooth sawfish saws
were reported from Guyana in 1886, and
an additional saw was later recorded in
1900. In Brazil, there is a 1910 report of
a female smalltooth sawfish. In 1914,
there is a report of a smalltooth sawfish
in Laguna de Terminos, Mexico (GBIF
Database).
In the middle part of the twentieth
century, there are reports of two female
smalltooth sawfish from Mexico in
1926. Rostral saws were found in
Suriname in 1943, 1944, and 1963, but
no additional location or specimen
information is known. One rostrum was
reported from Costa Rica in 1960 and
one rostral saw from Trinidad and
Tobago in 1944 (Faria et al., 2013).
Several whole individuals and one
rostrum were recorded from Guyana in
1958 and 1960. There are also several
other undated specimens recorded from
Guyana from this period (Faria et al.,
2013). There are other records of
smalltooth sawfish’s presence in the
western Atlantic Ocean but specific
information is lacking. For example,
Faria et al. (2013) report that 4 rostral
saws came from Mexico and two from
Belize. One female was reported from
Venezuela and two rostra from Trinidad
and Tobago. Despite lacking date
information, the GBIF Database and
Fishbase have reports of smalltooth
sawfish throughout South and Central
´
America: French Guiana (48), Mexico
´
(9), Guyana (6), Venezuela (3), Haitı (2),
and individual records from Colombia,
Nicaragua, and Belize.
In summary, while records are sparse,
it is likely the distribution of smalltooth
sawfish in the Atlantic Ocean is patchy
and has been reduced in a pattern
similar to largetooth sawfish. Data
suggests only a few viable populations
might exist outside the United States.
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The Caribbean Sea may have greater
numbers of smalltooth sawfish than
other areas given high quality habitats
and reduced urbanization. For example,
smalltooth sawfish have been repeatedly
reported along the western coast of
Andros Island, Bahamas (R.D. Grubbs,
Florida State University pers. comm. to
J. Carlson, NMFS, 2014) and The Nature
Conservancy noted two smalltooth
sawfish at the northern and southern
end of the island in 2006. Fishing
guides commonly encounter smalltooth
sawfish around Andros Island while
fishing for bonefish and tarpon (R.D.
Grubbs pers. comm. to J. Carlson,
NMFS, 2014), and researchers tagged
two in 2010 (Carlson et al., 2013). In
Bimini, Bahamas, generally one
smalltooth sawfish has been caught
every two years as part of shark surveys
conducted by the Bimini Biological
Station (D. Chapman pers. comm.to
Carlson, NMFS). In West Africa, Guinea
Bissau represents the last areas where
sawfish can be found (M. Diop pers.
comm. to IUCN, 2012). Anecdotal
reports indicate smalltooth sawfish may
also be found in localized areas off
Honduras, Belize, and Cuba (R. Graham,
Wildlife Conservation Society, pers.
comm. to IUCN, 2012).
Peer Review and Public Comments
In December 2004, the Office of
Management and Budget (OMB) issued
a Final Information Quality Bulletin for
Peer Review pursuant to the Information
Quality Act (IQA). The Bulletin was
published in the Federal Register on
January 14, 2005 (70 FR 2664). The
Bulletin established minimum peer
review standards, a transparent process
for public disclosure of peer review
planning, and opportunities for public
participation with regard to certain
types of information disseminated by
the Federal Government. The peer
review requirements of the OMB
Bulletin apply to influential or highly
influential scientific information. The
proposed rule and included status
review were considered influential
scientific information under this policy
and subject to peer review. Similarly, a
joint NMFS/FWS policy (59 FR 34270;
July 1, 1994) requires us to solicit
independent expert review from at least
three qualified specialists, concurrent
with the public comment period, on the
science that is the basis for listing
decisions. To ensure this final rule was
based on the best scientific and
commercial data available, we solicited
peer review comments from three
scientists familiar with elasmobranchs.
On June 4, 2013, we published a
proposed rule to list as endangered five
species of sawfish: Narrow sawfish (A.
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cuspidata), dwarf sawfish (P. clavata),
largetooth sawfish (P. pristis), green
sawfish (P. zijsron), and the non-U.S.
DPS of smalltooth sawfish (P.
pectinata), that occurs outside U.S.
waters, and opened a 90-day public
comment period (78 FR 33300). In the
proposed rule, we stated that we were
not proposing to designate critical
habitat for any of the five species
because they occur outside U.S. waters.
During our comment period we received
a request to extend the public comment
period by 45 days. On August 7, 2013,
we published a notice extending the
public comment period by 45 days (78
FR 48134). We received a total of four
public comments.
In the following sections of the
document we summarize and respond
to the comments received from the
public and peer reviewers on the
proposed rule.
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Peer Review Comments
Comment 1: One commenter noted
that the section of the proposed rule
addressing protective efforts did not
include details on the Sawfish
Conservation Strategy developed by the
IUCN Shark Specialist Group. The
commenter stated that the strategy is a
protective effort and will improve the
conservation status of sawfishes
worldwide. The commenter predicted a
medium to high certainty that the
actions identified in the Conservation
Plan, when implemented, will be
effective.
Response: We have included the
IUCN Sawfish Conservation Strategy in
the Protective Efforts section of this
final rule. The Services established two
basic criteria in the PECE for evaluating
conservation efforts: (1) The certainty
that the conservation efforts will be
implemented, and (2) the certainty that
the efforts will be effective. We
evaluated the IUCN Sawfish
Conservation Strategy and determined it
does not meet either criterion identified
in the PECE. The strategy identifies
actions for countries to develop
regulations or adopt management
actions to implement the strategy.
However, the strategy does not legally
bind any country to enact laws or
regulations, fund conservation actions,
or otherwise implement the strategy. We
believe there is considerable uncertainty
that the actions identified in the strategy
will be adopted by the various countries
within the range of the five species of
sawfish, and that resources are limited
to support these actions. Therefore, we
cannot find that the strategy will
decrease extinction risk for any of the
species.
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Comment 2: One commenter stated
that the Protective Efforts section of the
proposed rule did not include national
protective efforts except for the
Convention on International Trade of
Endangered Species of Wild Fauna and
Flora (CITES). The commenter stated
that sawfish protections in Australia
were likely effective, but protections in
India were likely ineffective.
Response: We updated the Protective
Efforts section of the rule and included
the new information on sawfish
protections and conservation efforts in
Australia from the Australian
Government’s recently published 2014
Draft Recovery Plan for Sawfish and
River Sharks (Department of
Environment, 2014). We also included
updated information on existing laws in
Australia and India designed to protect
sawfishes into the Inadequacy of
Existing Regulatory Mechanisms section
of this final rule.
Comment 3: It was suggested we use
information in Kyne et al. (2013) to
update the occurrence information for
P. clavata.
Response: We appreciate the new
information and updated the occurrence
information in the preceding sections.
The information did not impact our
evaluation of the status of P. clavata.
Comment 4: We received a question
about the origin of the 1996 record of
dwarf sawfish from the Mekong River
Basin, Laos.
Response: We cite Rainboth (1996) for
this report from the early 1900s that
assumed the dwarf sawfish was from the
Mekong River Basin, Laos. We
acknowledge no specimen exists to
confirm this report.
Comment 5: The validity of narrow
sawfish reports from Tasmania by
Deynat (2005) was questioned in one
comment given the cold, temperate
waters that do not support sawfish. The
commenter suggested the record of the
sawfish specimen in the fish collection
of CSIRO in Hobart, Tasmania was
erroneous.
Response: We reviewed the literature
and agree with the commenter. We
removed the reference to reports of
narrow sawfish in Tasmania.
Public Comments
Comment 1: One commenter
requested we cite a more recent
reference for the information on the
supply and demand of sawfish than the
1996 reference in the proposed rule.
Specifically, the commenter questioned
the statement that ‘‘sawfishes are in
high demand throughout the world for
display’’ and suggested that sawfishes
are no longer in high demand for
display in aquaria.
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Response: We updated our
information on the aquaria trade of
sawfishes on current supply and
demand of sawfishes in the Scientific
and Educational Uses section and
removed the statement cited by the
commenter. Although we believe that
sawfish are still in high demand in the
aquaria trade, we recognize that the
recent inclusion of all sawfishes under
CITES Appendix I limits the use of
sawfish for display and requires
acquisition of animals for aquaria from
captivity or captive breeding.
Comment 2: Several commenters
stated that they were concerned about
the impacts of including ‘‘injuring or
killing a captive sawfish through
experimental or potentially injurious
veterinary care or conducting research
or breeding activities on captive
sawfish, outside the bounds of normal
animal husbandry practices’’ in the list
of activities that could result in a
violation of the ESA Section 9
prohibitions. The concerns relate to the
impacts on captive propagation and
rearing programs being conducted by
aquaria, and on the use of the latest
advanced technological techniques
available for captive held animals. The
commenters requested clarification that
fish care and husbandry techniques
could continue to be used by aquaria.
Response: As stated in the proposed
rule, sawfish held in captivity at the
time of listing are afforded all of the
ESA protections and may not be killed
or injured or otherwise harmed, and,
therefore, must receive proper care. We
realize that the care of captive animals
necessarily entails handling or other
manipulation and we do not consider
such activities to constitute injury or
harm to the animals so long as adequate
care, including veterinary care, is
provided. Such veterinary care includes
confining, tranquilizing, and
anesthetizing sawfishes when such
practices, procedures, or provisions are
necessary and not likely to result in
injury.
On the effective date of a final listing,
ESA Section 9 take prohibitions
automatically apply for species listed as
endangered and any ‘take’ of the species
is illegal unless that take is authorized
under a permit or through an incidental
take statement. Incidental take
statements result from ESA Section 7
consultations on the effects of federal
activities. ESA Section 10 permits can
authorize directed take (e.g., for
scientific research or enhancement of
the species) or incidental take during an
otherwise lawful activity that would not
be subject to ESA section 7 consultation.
ESA Section 10 permits are issued to
entities or persons subject to the
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jurisdiction of the United States. We
encourage institutions with captive
sawfish who are considering activities
outside the bounds of normal animal
husbandry (e.g., breeding or research) to
contact NMFS Office of Protected
Resources, Permits and Conservation
Division, to determine if an ESA Section
10 permit is required to authorize the
proposed activity. We do not have
information regarding emerging
advances in fish care and animal
husbandry for sawfish held in captivity
so we cannot determine at this time if
they are outside the bounds of normal
care for captive animals.
Comment 3: Several commenters
requested clarification of the meaning of
the terms ‘‘non-commercial’’ and ‘‘noncommercially’’ as those terms are used
in the section titled Identification of
those Activities that Would Constitute a
Violation of Section 9 of the ESA.
Response: Section 3 of the ESA
defines the term ‘‘commercial activity’’
to mean ‘‘all activities of industry and
trade, including but not limited to, the
buying and selling of commodities and
activities conducted for the purposes of
facilitating such buying and selling:
Provided, however, That it does not
include exhibitions of commodities by
museums or similar cultural or
historical organizations.’’ NMFS will
use the definition of ‘‘commercial
activity’’ to evaluate whether an activity
is ‘‘non-commercial’’ or a sawfish is
being held ‘‘non-commercially’’ in
captivity.
Our listing determinations and
summary of the data on which it is
based, with the incorporated changes,
are presented in the remainder of this
document.
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Species Determinations
We first consider whether the narrow
sawfish (A. cuspidata), dwarf sawfish
(P. clavata), largetooth sawfish (P.
pristis), green sawfish (P. zijsron), and of
the non-U.S. DPS of smalltooth sawfish
(P. pectinata) meet the definition of
‘‘species’’ pursuant to section 3 of the
ESA. Then we consider if any
populations meet the DPS criteria.
Consideration as a ‘‘Species’’ Under the
Endangered Species Act
Based on the best available scientific
and commercial information described
above in the natural history sections for
each species, we have determined that
the narrow sawfish (A. cuspidata),
dwarf sawfish (P. clavata), largetooth
sawfish (P. pristis), and green sawfish
(P. zijsron) are taxonomically-distinct
species and therefore eligible for listing
under the ESA. The largetooth sawfish
(P. pristis) now includes the formerly
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recognized species P. microdon and the
previously listed P. perotteti. The
decision to list P. pristis will replace our
2011 listing determination for P.
perotteti.
Distinct Population Segments
In order to determine if the petitioned
and currently non-listed population
segment of smalltooth sawfish (P.
pectinata) constitutes a ‘‘species’’
eligible for listing under the ESA, we
evaluated it under our joint NMFSUSFWS Policy regarding the recognition
of distinct population segments (DPS)
under the ESA (61 FR 4722; February 7,
1996). We examined the three criteria
that must be met for a DPS to be listed
under the ESA: (1) The discreteness of
the population segment in relation to
the remainder of the species to which it
belongs; (2) the significance of the
population segment to the remainder of
the species to which it belongs; and (3)
the population segment’s conservation
status in relation to the Act’s standards
for listing (i.e., Is the population
segment, when treated as if it were a
species, endangered or threatened?).
A population may be considered
discrete, if it satisfies one of the
following conditions: (1) It is markedly
separated from other populations of the
same taxon as a consequence of
physical, physiological, ecological, or
behavioral factors; or (2) it is delimited
by international governmental
boundaries within which differences of
control of exploitation, management of
habitat, conservation status, or
regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D)
of the ESA.
We previously determined that
smalltooth sawfish in the United States
merited protection as a DPS and listed
the U.S. DPS of smalltooth sawfish as
endangered (68 FR 15674; April 1,
2003). At that time, there was no
information available to indicate
smalltooth sawfish in U.S. waters
interact with those in international
waters or other countries, suggesting
that the U.S. population may be
effectively isolated from other
populations. However, there were few
scientific data on the biology of
smalltooth sawfish, and it was not
possible to conclusively subdivide this
species into discrete populations on the
basis of genetics, morphology, behavior,
or other biological characteristics.
Because there were no identified
mechanisms regulating the exploitation
of this species anywhere outside of the
United States, we considered that lack
of protection as directly relevant to the
inadequacy of existing regulatory
mechanisms and a basis for considering
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the U.S. population as discrete across
international boundaries.
We now evaluate the non-U.S.
population of smalltooth sawfish to
determine if it meets the discreteness
criteria of the joint DPS policy. First, we
determine whether the non-U.S.
population of smalltooth sawfish is
discrete from the U.S. population
because it is delimited by international
governmental boundaries within which
differences of control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the ESA. Because we have
designated critical habitat for the U.S.
DPS population of smalltooth sawfish,
there is a significant regulatory
mechanism for protecting smalltooth
sawfish and their habitats in the United
States that does not exist for the nonU.S. population of smalltooth sawfish.
Movement data from smalltooth sawfish
tagged in U.S. and Bahamian waters also
indicate no movement to countries
outside where they were tagged. This
information provides support that the
non-U.S. population is discrete from the
already-listed U.S. DPS on the basis of
being markedly separate as a
consequence of ecological factors, in
addition to our previous determination
that the U.S. DPS is discrete on the basis
of international boundaries and
significant differences in regulatory
mechanisms. For smalltooth sawfish
outside the U.S., we have no
information regarding genetic or other
biological differences that would
provide a strong basis for further
separating the non-U.S. smalltooth
sawfish population into smaller,
discrete units. We, therefore, conclude
that the non-U.S. population of
smalltooth sawfish meets the
discreteness criterion of the joint DPS
policy and we consider this population
as a single potential DPS.
We next must consider whether the
non-U.S. population of smalltooth
sawfish meets the significance criterion.
The joint DPS policy gives examples of
potential considerations indicating the
population’s significance to the larger
taxon. Among these considerations is
evidence that the discrete population
segment would result in a significant
gap in the range of the taxon. Smalltooth
sawfish are limited in their distribution
outside of the United States to West
Africa, the Caribbean, Mexico, and
Central and South America. Loss of this
group of smalltooth sawfish would
result in a significant gap in the range
of this species and restrict distribution
to U.S. waters. Because the loss of
smalltooth sawfish in areas outside the
United States would result in a
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significant gap in the range of the
species, we conclude the non-U.S.
population of smalltooth sawfish is
significant as defined by the DPS policy.
Based on the above analysis of
discreteness and significance, we
conclude that the non-U.S. population
of smalltooth sawfish (P. pectinata)
meets the definition of a DPS and is
eligible for listing under the ESA, and
hereafter refer to it as the non-U.S. DPS
of smalltooth sawfish.
Extinction Risk
Our updated extinction risk analysis
provides a more detailed discussion of
the extinction risk analysis process that
we used to determine the risk of
extinction for narrow sawfish, dwarf
sawfish, green sawfish, largetooth
sawfish, and the non-U.S. DPS of
smalltooth sawfish to determine
whether the species are threatened or
endangered per the ESA’s definitions.
We used an adaptation of the approach,
including the primary concepts,
developed by Wainwright and Kope
(1999) to organize and summarize our
findings. This approach was originally
developed for salmonids and has been
adapted and applied in the review of
many other species (Pacific salmonid,
Pacific hake, walleye pollock, Pacific
cod, Puget Sound rockfishes, Pacific
herring, and black abalone) to
summarize the status of the species
according to demographic risk criteria.
The approach is useful when there is
insufficient quantitative data to support
development of population viability
models to investigate extinction risk and
it allows the incorporation of sparse and
qualitative data. Wainwright and Kope
(1999) identified key demographic
parameters that have a strong bearing on
extinction risk, with a focus on risks to
small populations from genetic effects
and population dynamics. Using these
concepts, adapted to the biology of these
sawfishes and our available data, we
estimated the extinction risk, based on
demographic factors, for each of the five
species under both current threats and
threats expected in the foreseeable
future. We also performed a threats
assessment by identifying the severity of
threats that exist now and in the
foreseeable future.
We defined the ‘‘foreseeable future’’
as the timeframe over which threats, or
the species’ response to those threats,
can be reliably predicted to impact the
biological status of the species. We
determined that the foreseeable future is
approximately three generation times,
calculated for each of the species based
on the demographic calculations of
Moreno Iturria (2012): Narrow sawfish,
14 years; dwarf sawfish, 49 years;
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largetooth sawfish, 48 years; green
sawfish, 38 years; and the non-U.S. DPS
of smalltooth sawfish, 30 years. After
considering the life history of each
species, availability of data, and type of
threats, we concluded that three
generations was an appropriate measure
to evaluate threats in the foreseeable
future. As a late-maturing species, with
slow growth rate and low productivity,
it would take more than one generation
for any conservation management action
to be realized and reflected in
population abundance indices. The
timeframe of three generations is a
widely used scientific indicator of
biological status, and has been applied
to decision making models by many
other conservation management
organizations, including the American
Fisheries Society, the CITES, and the
IUCN.
We considered three demographic
categories in which to summarize
available data and assess extinction risk
of each sawfish species: (1) Abundance,
(2) population growth rate/productivity,
and (3) genetic integrity which include
the connectivity and genetic diversity of
the species. We determined the
extinction risk for each category, for
both now and in the foreseeable future,
using a five level qualitative scale to
describe our assessment of the risk of
extinction. At the lowest level, a factor,
either alone or in combination with
other factors, is considered ‘‘unlikely’’
to significantly contribute to risk of
extinction for a species. The next lowest
level is considered to be a ‘‘low’’ risk to
contribute to the extinction risk, but
could contribute in combination with
other factors. The next level is
considered a ‘‘moderate’’ risk of
extinction for the species, but in
combination with other factors
contributes significantly to the risk of
extinction. A ranking of ‘‘high’’ risk
means that factor by itself is likely to
contribute significantly to the risk of
extinction. Finally, a ranking of ‘‘very
high’’ risk means that factor is
considered ‘‘highly likely’’ to contribute
significantly to the risk of extinction.
We ranked abundance as high or very
high risk which is likely to contribute
significantly to the current and
foreseeable risk of extinction for all five
species. While it appears the northern
coast of Australia supports the largest
remaining groups of dwarf, largetooth,
green, and narrow sawfish in the Pacific
and Indian Ocean, data from the
Queensland, Australia Shark Control
Program show a clear decline in sawfish
catch (non-species-specific) over a 30year period from the 1960s. In addition,
it shows the complete disappearance of
sawfish in southern regions (Stevens et
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al., 2005). The available data on
abundance of sawfishes indicates there
are still some isolated groups of sawfish
in the western and central Indo-Pacific
region, but their abundance has likely
declined from historic levels.
Smalltooth sawfish are still being
reported outside of U.S. waters in the
Caribbean Sea, but records are few and
mostly insular (e.g., Andros Island)
where habitat is available and gillnet
fisheries are not a threat to the species
(see below). There are only four records
of largetooth sawfish in the eastern
Atlantic Ocean over the last decade. In
the western Atlantic, recent largetooth
sawfish records are from only the
Amazon River basin and the Rio
Colorado-Rio San Juan area in
Nicaragua.
Wainright and Kope (1999) stated
short- and long-term trends in
abundance are a primary indicator of
extinction risk. These trends may be
calculated from a variety of quantitative
data such as research surveys,
commercial logbook or observer data,
and landings information when
accompanied by effort, but there is an
absence of long-term monitoring data for
all five sawfishes. We looked at the
available data closely to see if we could
support inferences about extinction risk
based on the trends in past observations
using the presence of a particular
species at specified places and times
(e.g., Dulvy et al., 2003; Rivadeneira et
al., 2009). The available museum
records, negative scientific survey
results, and anecdotal reports do
indicate the abundance trend for all five
sawfishes is declining and population
sizes are small. Information available on
the species’ distribution indicates the
species’ ranges have also contracted. In
many areas where sawfish still occur,
they are subject to commercial and
artisanal fisheries and potential habitat
loss. We therefore ranked the risk of
extinction posed by the sawfishes’
abundances as high, now and into the
foreseeable future.
We next considered the species’
potential growth rates and productivity
as measures of their ability to recover
from depleted levels and provide
inherent protection against extinction
risk. Sawfish have historically been
classified as having both low
reproductive productivity and low
recovery potential. The demography of
smalltooth and largetooth sawfish from
the northwest Atlantic Ocean that was
originally investigated using an agestructured life table (Simpfendorfer,
2000). Using known estimates of
growth, mortality, and reproduction at
the time, Simpfendorfer (2000)
determined that intrinsic rates of
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population increase ranged from 8 to 13
percent per year, and population
doubling times were approximately 5 to
8.5 years for both species. These
estimates included assumptions that
there was no fishing mortality, no
habitat limitations, no population
fragmentation, or other effects of small
population sizes. Simpfendorfer (2006)
further modeled the demography of
smalltooth sawfish using a method for
estimating the rebound potential of a
population by assuming that maximum
sustainable yield was achieved when
the total mortality was twice that of
natural mortality. This demographic
model produced intrinsic rates of
population increase that were from two
to seven percent per year for both
smalltooth and largetooth sawfish.
These values are similar to those
calculated by Smith et al. (2008) using
the same methodology corresponding to
elasmobranch species with the lowest
productivity. Musick et al. (2000) noted
that species with intrinsic rates of
increase of less than 10 percent were
particularly vulnerable to rapid
population declines and a higher risk of
extinction.
Some recent studies on the life history
of sawfish, however, indicate they are
potentially more productive than
originally proposed. Growth rates (von
Bertalanffy ‘‘K’’) for some species, like
narrow sawfish, approach 0.34 per year
(Peverell, 2008). Data from tag-recapture
studies and analysis of vertebral growth
bands from smalltooth sawfish indicate
that the first few years after birth
represent the time when growth is most
rapid (e.g., Simpfendorfer et al., 2008;
Scharer et al., 2012). Using updated life
history information, Moreno Iturria
(2012) calculated intrinsic rates of
increase for these five species of sawfish
and determined values ranging from a
low of 0.02 per year for green sawfish
to a high of 0.27 per year for narrow
sawfish with dwarf sawfish being
second highest at 0.10 per year.
Considering this information, and the
inferred declining trend in abundance,
we conclude productivity is a moderate
risk for the narrow sawfish but a high
risk for the other four species. We also
determined that productivity would
remain a moderate risk for the narrow
sawfish and is a high risk for the other
four species, in the foreseeable future.
We also assessed the species’
extinction risk, based on genetic
diversity, spatial structure and
connectivity. Population structure and
levels of genetic diversity have recently
been assessed for the green sawfish,
dwarf sawfish, and largetooth sawfish
across northern Australia using a
portion of the mtDNA control region.
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Phillips et al. (2011) found statistically
significant genetic structure within
species and moderate genetic diversity
among these species. These results
suggest that sawfish may be more
vulnerable to local extirpation along
certain parts of their range, especially in
areas where the population has been
fragmented and movement between
these areas is limited. However, these
results do not necessarily suggest a
higher risk of extinction throughout the
entire range of the species. Chapman et
al. (2011) investigated the genetic
diversity of the U.S. DPS of smalltooth
sawfish that has declined to between
one percent to five percent of its
abundance at the turn of the twentieth
century, while its core distribution has
contracted to less than 10 percent of its
former range (NMFS, 2009).
Surprisingly, given the magnitude of
this population decline and range
contraction, the U.S DPS of smalltooth
sawfish does not exhibit any sign of
genetic bottlenecks, and it has genetic
diversity that is similar to other, less
depleted elasmobranch populations
(Chapman et al., 2011). Given that all
five species of sawfish considered here
have suffered similar abundance
declines, we believe this conclusion
should serve as a surrogate for the other
sawfish species. Because the U.S. DPS
of smalltooth sawfish has not undergone
a genetic bottleneck, we ranked genetic
diversity as a moderate risk for all
sawfish species as it is likely, in
combination with other factors, to
contribute significantly to the risk of
extinction. However, we determined
that the risk of extinction due to the lack
of connectivity was high for all five
species, primarily because all
populations have undergone severe
fragmentation. While genetic results
provide optimism for the remaining
populations of sawfish, this does not
preclude the promotion of management
actions to enhance connectivity among
populations that have been historically
fragmented. We are also somewhat
optimistic that sawfish populations may
begin to rebuild in some areas and the
risk of connectivity was determined to
decrease for smalltooth and the narrow
sawfish in the foreseeable future,
although by only a small amount.
After reviewing the best available
scientific data and assessing the
extinction risk on the five species of
sawfishes based on their status and
demography, we conclude the risk of
extinction for all five species of sawfish
is high.
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Summary of Factors Affecting the Five
Species of Sawfishes
Next we consider whether any of the
five factors specified in section 4(a)(1) of
the ESA are contributing to the
extinction risk of these five sawfishes.
The Present or Threatened Destruction,
Modification, or Curtailment of Its
Habitat or Range
We identified destruction,
modification, or curtailment of habitat
or range as a potential threat to all five
species of sawfishes and determined
this factor is currently, and in the
foreseeable future, contributing
significantly to the risk of extinction of
these species.
Coastal and Riverine Habitats
Loss of habitat is one of the factors
determined to be associated with the
decline of smalltooth sawfish in the U.S.
(NMFS, 2009). As juveniles, sawfishes
rely on shallow nearshore
environments, primarily mangrovefringed estuaries as nurseries (e.g.,
Wiley and Simpfendorfer, 2010; Norton
et al., 2012). Coastal development and
urbanization have caused these habitats
to be reduced or removed from many
areas throughout the species’ historic
and current range. Habitat loss was
identified as one of the most serious
threats to the persistence of all species
of sawfish, posing high risks for
extinction. It is still unclear how
anthropogenic perturbations to habitats
affect the recruitment of juvenile
sawfish, and therefore adequate
protection of remaining natural areas is
essential. Given the threat from coastal
urbanization coupled with the predicted
reduction of mangroves globally
(Alongi, 2008), we believe the risk of
habitat loss would significantly
contribute to both the decline of sawfish
and their reduced viability.
We expect habitat modification
throughout the range of these sawfishes
to continue with human population
increases. As humans continue to
develop rural areas, habitat for other
species, like sawfish, becomes
compromised (Compagno, 2002b).
Habitat modification affects all five
species of sawfish, especially those
inshore, coastal habitats near estuaries
and marshes (Compagno and Last, 1999;
Cavanagh et al., 2003; Martin, 2005;
Chin et al., 2010; NMFS, 2010). Mining
and mangrove deforestation severely
alter the coast habitats of estuaries and
wetlands that support sawfish
(Vidthayanon, 2002; Polhemus et al.,
2004; Martin, 2005). In addition,
riverine systems throughout most of
these species’ historical range have been
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altered or dammed. For example, the
potential expansion of the McArthur
River Mine would permanently realign
channels that would in turn affect the
number of pools formed during the wet
and dry seasons, many of which are
used as refuge areas for dwarf, green, or
largetooth sawfish (Polhemus et al.,
2004; Gorham, 2006). In addition to the
potential expansion of the McArthur
River Mine, the Nicaragua government
is proposing to build a cross-country
canal through habitats currently used by
the remaining largetooth sawfish
population in Lake Nicaraugua (BBC
News, Latin America and Caribbean,
2013).
Although the status of habitats across
the global range of these sawfishes is not
well known, we expect the continued
development and human population
growth to have negative effects on
habitat, especially to nearshore nursery
habitats. For example, Ruiz-Luna et al.
(2008) acknowledge that deforestation of
mangrove forests in Mexico has
occurred from logging practices,
construction of harbors, tourism, and
aquaculture activities. Valiela et al.
(2001) reported on mangrove declines
worldwide. They showed that the area
of mangrove habitat in Brazil decreased
from 9652 to 5173 square miles (24,999
to 13,398 square kilometers) between
1983 and 1997, with similar trends in
Guinnea-Bissau 1837 to 959 square
miles (4758 to 2484 square kilometers)
from 1953 to 1995. The areas with the
most rapid mangrove declines in the
Americas included Venezuela, Mexico,
Panama, the U.S., and Brazil. Along the
western coast of Africa, the largest
declines have occurred in Senegal,
Gambia, Sierra Leone, and GuinneaBissau. World-wide mangrove habitat
loss was estimated at 35 percent from
1980 to 2000 (Valiela et al., 2001). These
areas where mangroves are known to
have decreased are within both the
historic and current ranges of these five
species.
Hydroelectric and Flood Control Dams
Hydroelectric and flood control dams
pose a major threat to freshwater inflow
into the euryhaline habitats of
sawfishes. Alterations of flow, physical
barriers, and increased water
temperature affect water quality and
quantity in the rivers, as well as
adjacent estuaries that are important
nursery areas for sawfish. Regulating
water flow affects the environmental
cues of monsoonal rains and increased
freshwater flow for pupping (Peverel,
2008; Morgan et al., 2011). Changes in
siltation due to regulated water flow
may also affect benthic habitat or prey
abundance for these sawfishes
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(Compagno, 2002; Polhemus et al.,
2004; Martin, 2005; Thorburn et al.,
2007; Chin et al., 2010; Morgan et al.,
2010a).
New dams being proposed to provide
additional irrigation to farmland
upstream may affect sawfish habitat. For
example, the Gilbert River, in
Queensland, Australia drains into the
Gulf of Carpentaria, which is the
nursery area for green, dwarf, and
largetooth sawfish. Further modification
of the McArthur and Gilbert Rivers,
along with increased commercial fishing
in coastal waters, will negatively affect
sawfishes by reducing available habitat
while increasing bycatch mortality
(Gorham, 2006).
Water Quality
Largetooth sawfish in particular, and
likely the other sawfishes, have
experienced a loss of habitat throughout
their range due to the decline in water
quality. Agriculture and logging
practices increase runoff, change
salinity, and reduce the flow of water
into freshwater rivers and streams that
affects the habitat of the largetooth
sawfish (Polhemus et al., 2004; IUCN
Red List, 2006); mining seems to be the
most detrimental activity to water
quality. Pollution from industrial waste,
urban and rural sewage, fertilizers and
pesticides, and tourist development all
end up in these freshwater systems and
eventually the oceans. Pollution from
these operations has caused a reduction
in the number of sawfish in these
freshwater systems (Vidthayanon, 2002;
Polhemus et al., 2004).
In summary, habitat alterations that
potentially affect sawfishes include
commercial and residential
development; agricultural, silvicultural,
and mining land uses; construction of
water control structures; and
modification to freshwater inflows. All
sawfishes are vulnerable to a host of
habitat impacts because they use rivers,
estuaries, bays, and the ocean at various
times of their life cycle. Based on our
review of current literature, scientific
surveys and anecdotal information on
the historic and current distribution, we
find that destruction, modification, and
curtailment of habitat or ranges are a
factor affecting the status of each
species. We conclude that this factor is
contributing, on its own or in
combination with other factors, to the
extinction risk of all five species of
sawfishes.
Overutilization for Commercial,
Recreational, Scientific, or Educational
Purposes
We identified overutilization for
commercial, recreational, scientific, or
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educational purposes as a potential
threat to all five species of sawfishes
and determined that it is currently and
in the foreseeable future contributing
significantly to their risk of extinction.
Commercial Fisheries
Commercial fisheries pose the biggest
threat to these sawfishes, as these
species are bycatch from many fisheries.
Their unusual morphology and
prominent saw makes sawfishes
particularly vulnerable to most types of
fishing gear, most notably any type of
net (Anak, 2002; Hart, 2002; Last, 2002;
Pogonoski et al., 2002; Cavanagh et al.,
2003; Porteous, 2004; Stevens et al.,
2005; Gorham 2006; IUCN Red List,
2006; Chidlow, 2007; Field, 2009; Chin
et al., 2010; NMFS, 2010; Morgan et al.,
2011). Trawling gear is of particular
concern as it is the most common gear
used within the range and habitat of
sawfishes (Compagno and Last, 1999;
Taniuchi, 2002; Walden and Nou, 2008).
In Thailand, all sawfish fins obtained
and sold to markets are a result of
bycatch by otter-board trawling and
gillnet fisheries as there are no directed
sawfish fisheries in the country (Pauly,
1988; Vidthayanon, 2002). The Lake
Nicaragua commercial fishery for
largetooth sawfish that collapsed prior
to the 1980’s was comprised mostly of
gillnet boats (Thorson, 1982a), and the
commercial small coastal shark fishery
in Brazil mainly uses gillnets and some
handlines (Charvet-Almeida, 2002).
Subadult and adult smalltooth sawfish
have been reported as bycatch in the
U.S. Gulf of Mexico and south Atlantic
shrimp trawl fishery (NMFS SEFSC,
2011); however, if proper techniques are
used, all sawfish species, particularly
adults, are fairly resilient and can be
released alive from most fishing gear
(Lack et al., 2009).
Live release of sawfishes from
commercial fishing gear does occur but
sawfishes are often retained. The meat
is generally consumed locally, but the
fins and rostra are of high value and
sold in markets where these products
are unregulated (CITES, 2007). In Brazil,
a captured sawfish is most likely
retained because of the value of their
products, as the rostra, rostral teeth, and
fins are valued at upwards of $1,000
U.S. in foreign markets (NMFS, 2010a).
The proportion of largetooth sawfish in
these markets is unknown, although as
many as 180 largetooth sawfish saws
were annually sold at a single market in
northern Brazil in the early 2000’s
(McDavitt and Charvet-Almeida, 2004).
The Trade Records Analysis of Flora
and Fauna in Commerce (TRAFFIC)
organization found that meat, liver oil,
fins, and skin are among the most
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preferred sawfish products in Asian
markets (Anak, 2002; Vidthayanon,
2002). In the Gulf of Thailand, over
5,291 US tons (4,800 tonnes) of rays
were caught annually from 1976 to
1989; at the same time over 1,102 US
tons (1,000 tonnes) of rays were caught
in the Andaman Sea (Vidthayanon,
2002). It is likely that most of these
products were sold in Asian markets
because of the high demand for sawfish
products. Reports of sawfish products in
various markets throughout Asia are
often inconsistent and inaccurate
despite international rules on trade and
possession of sawfish products (Fowler,
2002; Clarke et al., 2008; Kiessling et al.,
2009).
Recreational or commercial fishing
gear may be abandoned or lost at sea.
These ‘‘ghost nets’’ are an entanglement
hazard for sawfishes and have become
an increasing problem in the Gulf of
Carpentaria where over 5,500 ghost nets
were removed in 2009. Sawfish captures
are expected to occur in regions where
no quantitative information about ghost
nets exists (Gunn et al, 2010).
Misidentification, general speciescomposition grouping, and failure to
record information are all concerns for
reporting sawfish captures in direct or
indirect commercial fisheries (Stobutzki
et al., 2002b). With little enforcement of
regional and international laws, the
practice of landing sawfishes may
continue (NMFS, 2010a). All sawfish
populations have been declining
worldwide, partly due to the negative
effects of commercial fishing (Stevens et
al., 2000; Peverell, 2008).
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Recreational Fisheries
Sawfish are bycatch of many
recreational fisheries throughout their
range, even in areas where they are
protected, including many Australian
rivers (Walden and Nou, 2008; Field et
al., 2009). Peverell (2008) reports that
some sawfish are a target sport fish for
recreational fishermen in the Gulf of
Carpentaria, Queensland. Historical
information from the U.S. indicates that
recreational hook and line fishers in
Texas sometimes target large sharks as
trophy fish but may capture sawfish
(Burgess et al., 2009). Elsewhere in the
United States, the abundance of
sawfishes is low and likely never high
enough for recreational fishers to
encounter sawfish, much less target it
(NMFS, 2010a). With the increase in
human population along the coast,
recreational fishing has the potential to
put additional pressure on sawfish
species that use coastal habitats
(Walden and Nou, 2008).
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Indigenous Take
Due to the large populations of
various indigenous people throughout
the range of these five species, and the
lack of data on the animals they harvest,
the number of sawfish taken by local
peoples is unknown. Elasmobranchs are
caught for consumption throughout the
Indo-Pacific. In some areas, the meat
and fins of these animals are of high
market value, and therefore they are
sold rather than consumed locally. Due
to this unregulated consumption,
removal of elasmobranchs, which
includes sawfishes, is a threat to their
population(s) (Compagno and Last,
1999; Pogonoski et al., 2002;
Vidthayanon, 2002; Thorburn et al.,
2007; Peverell, 2008; Morgan et al.,
2010a).
Some studies have been conducted on
the use and value of elasmobranch parts
to various indigenous groups,
particularly those in eastern Sabah,
Malaysia. One study (Almada-Villela,
2002) found the majority of natives from
Pulau Tetabuan and Pulau Mabul only
take what is necessary for subsistence.
Sawfish rostra are also valued and kept
as decoration or given as gifts at the
expense of the animal (Almada-Villela,
2002; McDavitt et al., 1996;
Vidthayanon, 2002).
Protective Coastal Nets
Protective gillnets to prevent shark
attacks on humans is used in some areas
but can have a negative impact due to
bycatch. Sawfishes are highly
susceptible to capture in nets because
their saws are easily tangled in nets. The
Queensland Shark Control Program in
Australia places nets along beaches
during the summer months. From 1970
to 1990, sawfish bycatch in these nets
declined despite relatively constant
effort; likely due to an overall decline in
sawfish populations (Stevens et al.,
2005). In South Africa, the first
protective gillnets lined the southeast
tip of the continent’s coast as early as
1952. By 1990, over 27 mi (44 km) of
nets lined the area between Richards
Bay and Mzamba (Dudley and Cliff,
1993). About 350 sharks and rays were
captured in these nets between 1981
and 1990. A high percentage of
entangled sawfish are released alive
because of their ability to breathe while
motionless. Dudley and Cliff (1993)
reported that 100 percent of largetooth
sawfish and 67 percent of smalltooth
sawfish caught during that time were
released alive. Still, subsequent
mortality post-release due to stress or
injury from the process is unknown and
potentially detrimental given other
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fishing pressures (Dudley and Cliff,
1993).
Scientific and Educational Uses
Sawfishes are unique animals that are
currently on public display in many
large aquariums. Removal of sawfishes
from their natural habitats has caused
some concern for these sawfish species
and their ecosystems. No information is
available on the level of mortality that
occurs during the capture and
transporting of live sawfish to aquaria.
Removal of female sawfish from the
wild could have an effect on the future
reproductive capacity of that population
(Anak, 2002; Harsan and Petrescu-Mag,
2008). Limited information is available
regarding the number of sawfish that
have been removed from the wild for
display in aquaria. All sawfish removed
from Australian waters for aquaria
collections have been reported as
juveniles (S. Olson, Association of Zoos
and Aquariums (AZA), 2013 pers.
comm). The two most recent imports of
largetooth sawfish to an Association of
Zoos and Aquariums (AZA) accredited
facility were in 2007 and 2008 (S.
Olson, AZA, 2013 pers. comm).
In July 2011, the Australian CITES
Scientific Authority for Marine Species
reviewed their 2007 non-detriment
finding for the export of P. microdon
and found that it was not possible to
conclude with a reasonable level of
certainty that any harvest for export
purposes would not be detrimental to
the survival or recovery of the species
(DSEWPaC, 2011). Since then,
international trade in freshwater sawfish
from Australia has ceased.
Worldwide, we are not aware of any
narrow sawfish in captivity (Peverell,
2005, 2008). We are aware of 2 dwarf
sawfish held in captivity in Japan
(McDavitt, 2006). Largetooth sawfish are
the most common sawfish species in
captivity (NMFS, 2010a). Juvenile
largetooth measuring less than 3.5 ft (1
m) TL on average are most often caught
for the aquaria trade as they are easier
to transport than adults (Peter and Tan,
1997).
Globally, scientists are collecting
information on sawfish biology.
Research efforts began in 2003 on the
U.S. DPS population of smalltooth
sawfish and no negative impacts have
been associated with this research to
date.
In summary, while no quantitative
data on fishery impacts are available, we
conclude that given the susceptibility of
sawfish to entanglement in gillnets and
trawl nets that are commonly used
throughout their range, sawfishes are
likely captured as incidental take. We
are not aware of any fisheries
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specifically targeting sawfishes. This
impact from fisheries is the most likely
single cause of the observed range
contractions and reduced abundance in
many areas of their former range. Trade
of sawfish parts occurs throughout the
world. Sawfish have been exploited for
their fins, rostra, and teeth. Sawfish fins
have been report in the shark fin trade
since the early 1900s (Mountnorris,
1809). Trade of sawfish parts occurs on
Internet sites such as eBay and
Craigslist. Trade of sawfish parts (e.g.,
fins, rostral teeth, and rostra) are also
ongoing threats to all five species
(Harrison et al., 2014). Therefore, we
conclude the overutilization for
commercial, recreational, scientific, or
educational purposes, alone or in
combination with other factors as
discussed herein, is contributing
significantly to the risk of extinction of
the narrow, dwarf, largetooth, green,
and the non-U.S. DPS of smalltooth
sawfish.
Disease and Predation
We have determined that disease and
predation are not potential threats to
any of the five species of sawfish and
that it is unlikely that these factors, on
their own or in combination with other
factors, are contributing significantly to
their risk of extinction of all five sawfish
species.
These species co-occur with other
sawfishes and large sharks, but we are
not aware of any studies or information
documenting interspecific competition
in terms of either habitat or prey (NMFS
2010a). Thorson (1971) speculated that
the Lake Nicaragua bull shark
population may compete with
largetooth sawfish, as both were
prevalent, but he offered no additional
data. Sawfish have been documented
within the stomach of a dolphin
(Tursiops truncatus) near Bermuda
(Bigelow and Schroeder, 1953; MonteLuna et al., 2009), in the stomach of a
bull shark (C. leucas) in Australia
(Thorburn et al., 2004), and evidence of
bite marks from what appeared to be a
bull shark (C. leucas) on a juvenile
smalltooth sawfish in the United States
have been reported (T. Wiley-Lescher,
Haven Worth Consulting, 2012 pers.
comm). Crocodiles also prey on
sawfishes (Cook and Compagno, 2005).
There is no evidence that unusual levels
of disease or predation affect any of the
five sawfish species. Based on the
information available on disease and
predation for all five species of sawfish,
we have determined that disease and
predation on their own, or in
combination with other factors, do not
pose an extinction risk to any of these
sawfishes.
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Inadequacy of Existing Regulatory
Mechanisms
We identified inadequacy of existing
regulatory mechanisms as a potential
threat to each of the five species of
sawfish. We determined that this factor
alone, or in combination with other
factors, is contributing significantly to
their risk of extinction.
First, we reviewed general or global
regulatory protections for sawfish. The
use of turtle exclusion devices (TEDs) in
the nets of trawl fisheries to conserve
sea turtles occurs throughout much of
the range of sawfishes, but TEDs are not
efficient in directing sawfish out of nets
because sawfish rostra get entangled
(Stobutzki et al., 2002a; Brewer et al.,
2006) prior to reaching the TED. TEDs
are often used when trawling occurs
along the sea bottom at depths of 49 ft
to 131 ft (15 to 40 m), areas where
sawfish are likely to be found (Stobutzki
et al., 2002a). Most sawfishes show no
difference in recovery after going
through a trawl net, regardless of the
presence or absence of a TED (Griffiths,
2006). Stobutzki et al. (2002a) found
that large females are more likely to
survive capture after passing through a
trawling net and TED compared to
smaller males. Only narrow sawfish
were found to benefit from the presence
of TEDs in nets as 73.3 percent escaped
(Brewer et al., 2006; Griffiths, 2006). In
general, TEDs tend to have negligible
impact on sawfish that get captured by
trawling nets (Stobutzki et al., 2002a;
Griffiths, 2006), but they do provide an
escape route if the animal does not get
entangled.
Data reporting agencies (i.e., customs
and national fisheries) are often
inconsistent in their reporting of
wildlife trade (Anak, 2002). Reports are
often vague and include general
descriptions like ‘‘shark fin’’ or ‘‘ray,’’
providing practically no information of
trading rates of specific products (Lack
and Sant, 2011). Many countries in the
Indo-Pacific do not report bycatch
statistics or elasmobranchs taken
illegally (Holmes et al., 2009). In order
for effective management plans to be
implemented in fin markets and for
sawfish product trade, data need to be
consistent.
Next, we reviewed regional or country
specific regulatory protections for
sawfish. Many countries in the IndoPacific and the Middle East do not have
formal legislation for management or
national protection of the sawfish that
may occur in their waters. Presently,
Thailand has regulated some fisheries,
but has no protective legislation for any
elasmobranch in the country except for
export of marine species for aquaria
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(Vidthayanon, 2002). Among Middle
Eastern countries that fish for sharks,
only Iran has implemented an
International Plan of Action for the
Conservation and Management of
Sharks (IPOA Shark Plan). Nine Arab
countries have recently signed a
Memorandum of Understanding on the
Conservation of Migratory Sharks to
improve shark conservation measures
under the United Nations Environment
Programme Convention on Migratory
Species. Countries in Africa face similar
circumstances as enforcement for
sawfish protection is unknown (NMFS,
2010a). Countries that do have
protective legislation are often unable to
effectively patrol their waters, and
fishing restrictions are routinely
violated by foreign vessels (Lack. and
Sant, 2008). In one study, genetic testing
(DNA barcoding) was used to identify
fins from green sawfish confiscated from
foreign boats illegally fishing in
northern Australian waters (Holmes,
2009).
The Australian government listed the
largetooth, green, and dwarf sawfishes
as vulnerable on their Environmental
Protection and Biodiversity
Conservation (EPBC) Act list. The EPBC
Act protects these sawfish and prohibits
killing, injuring, taking, trading,
keeping, or moving an individual
without a permit. Even with these
protections in place, the Draft Recovery
Plan for Sawfish and River Sharks
(Department of the Environment, 2014)
reports that these three sawfish species
have experienced substantial population
declines.
In summary, several organizations are
trying to regulate and manage sawfish
but often these regulations and
management initiatives are inadequate.
Illegal exploitation by foreign fishers
often occurs when regulations exist but
are not enforced (Kiessling et al., 2009).
Preventative measures on existing
fishing mechanisms to avoid sawfish
catch, international monitoring of trade
and bycatch, and governmental
influence on fisheries are not presently
sufficient to protect sawfishes. Specific
regulation and monitoring of sawfishes
by country would provide better
protection (Vidthayanon, 2002; Walden
and Nou, 2008). Therefore, we conclude
the inadequacy of existing regulatory
mechanisms has and continues to
significantly contribute to the risk of
extinction of the narrow, dwarf,
largetooth, green, and the non-U.S. DPS
of smalltooth sawfish.
Other Natural or Manmade Factors
Affecting Its Continued Existence
In the proposed rule, we determined
this was not a factor contributing
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significantly to the risk of extinction of
all five species of sawfish. We reevaluated the information for this factor
and changed our conclusion from the
proposed rule based on the fact that
sawfish life history traits, which
consists of slow growth rates, late
maturity, long life spans, and low
fecundity rates. These life history traits
do not enable them to respond rapidly
to additional sources of mortality, such
as overexploitation and habitat
degradation. Scientific information
available on all five species of sawfish
indicates that other natural or manmade
factors are potential threats to all of the
five species of sawfish. We conclude it
is likely that these factors, on their own
or in combination with other factors, are
contributing significantly to the risk of
extinction for all five sawfish species.
An increase in global sea-surface
temperature and sea level may already
be influencing sawfish populations
(Clark, 2006; Walden and Nou, 2008;
Chin et al., 2010). Fish assemblages are
likely to change their distribution and
could affect the prey base for sawfishes.
Estuaries, including sawfish pupping
grounds, may be affected as climate
change changes patterns in freshwater
flow due to rainfall and droughts.
Skewed salinities in these areas or
extreme tide levels might discourage
adults from making up-river migrations
(Clark, 2006). Saltwater marsh grass and
mangrove areas play important roles in
sawfish habitat as well (Simpfendorfer
et al., 2010); any disruption to these
areas may affect sawfish populations.
There is little agreement, however, on
the effects that climate change will have
on sawfish and their environments
specifically (Clark, 2006; Chin et al.,
2010).
Red tide is the common name for a
harmful algal bloom (HAB) of marine
algae (Karenia brevis) that can make the
ocean appear red or brown. Karenia
brevis is one of the first species ever
reported to have caused a HAB and is
principally distributed throughout the
Gulf of Mexico, with occasional red
tides in the mid- and south-Atlantic
United States. Karenia brevis naturally
produces a brevetoxin that is absorbed
directly across the gill membranes of
fish or through ingestion of algal cells.
While many HAB species are nontoxic
to humans or small mammals, they can
have significant effects on aquatic
organisms. Fish mortalities associated
with K. brevis events are very common
and widespread. The mortalities affect
hundreds of species during various
stages of development. Red tide toxins
can cause intoxication in fish, which
may include violent twisting and
corkscrew swimming, defecation and
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regurgitation, pectoral fin paralysis,
caudal fin curvature, loss of
equilibrium, quiescence, vasodilation,
and convulsions, culminating in death.
However, it is known that fish can die
at lower cell concentrations and can
also apparently survive in much higher
concentrations. In some instances,
mortality from red tide is not acute, but
may occur over a period of days or
weeks after exposure to subacute toxin
concentrations. There is no specific
information on red tide effects on
sawfish, but a single report exists of a
smalltooth sawfish that was found dead
along the west coast of Florida, during
a red tide event (International Sawfish
Encounter Database, 2009). Therefore,
we conclude that sawfishes occurring in
the U.S. Gulf of Mexico are vulnerable
to red tide, but there is little information
documenting direct mortality resulting
from exposure to red tide (NMFS,
2010a). Harmful algal blooms also exist
in waters outside of the U.S. Gulf of
Mexico therefore, it is probable that all
sawfishes are vulnerable to harmful
algal blooms wherever they occur.
Collectively, these other natural or
manmade factors may be affecting the
continued existence of the narrow,
dwarf, largetooth, green, and the nonU.S. DPS of smalltooth sawfish. Based
on the results from our extinction risk
analysis and information on other manmade factors affecting all five species of
sawfish, this factor is contributing to
their extinction risk.
Overall Risk Summary
After considering the extinction risks,
both threat-based and demographic, for
each of the five species of sawfish, we
have determined the narrow, dwarf,
largetooth, and green sawfish and the
non-U.S. DPS of smalltooth sawfish are
in danger of extinction throughout all of
their ranges due to (1) present or
threatened destruction, modification or
curtailment of habitat, (2)
overutilization for commercial,
recreational, scientific, or educational
purposes, (3) inadequacy of existing
regulatory mechanisms, and (4) other
natural or manmade factors affecting
their continued existence, and low
abundance, lack of connectivity, and
genetic diversity.
Protective Efforts
Section 4(b)(1)(A) of the ESA requires
the Secretary, when making a listing
determination for a species, to take into
consideration those efforts, if any, being
made by any State or foreign nation to
protect the species. In judging the
effectiveness of efforts not yet
implemented, or those existing
protective efforts that are not yet fully
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effective, we rely on the Services’ joint
‘‘Policy for Evaluation of Conservation
Efforts When Making Listing Decisions’’
(‘‘PECE’’; 68 FR 15100; March 28, 2003).
The PECE policy is designed to ensure
consistent and adequate evaluation on
whether any conservation efforts that
have been recently adopted or
implemented, but not yet proven to be
successful, will result in recovering the
species to the point at which listing is
not warranted or contribute to forming
the basis for listing a species as
threatened rather than endangered. The
purpose of the PECE policy is to ensure
consistent and adequate evaluation of
future or recently implemented
conservation efforts identified in
conservation agreements, conservation
plans, management plans, and similar
documents when making listing
determinations. The PECE provides
direction for the consideration of
conservation efforts identified in these
documents that have not yet been
implemented, or have been
implemented but not yet demonstrated
effectiveness. The policy is expected to
facilitate the development of
conservation efforts by states and other
entities that sufficiently improve a
species’ status so as to make listing the
species as threatened or endangered
unnecessary.
Two basic criteria were established in
the PECE to use in evaluating efforts
identified in conservations plans,
conservation agreements, management
plans or similar documents: (1) The
certainty that the conservation efforts
will be implemented; and (2) the
certainty that the efforts will be
effective. When we evaluate the
certainty of whether or not the
formalized conservation effort will be
implemented, we may consider the
following: Do we have a high level of
certainty that that the resources
necessary to carry out the conservation
effort are available? Do the parties to the
conservation effort have the authority to
carry it out? Are regulatory or
procedural mechanisms in place to
carry out the efforts? If the conservation
effort relies on voluntary participation,
we will evaluate whether the incentives
that are included in the conservation
effort will ensure the level of
participation necessary to carry out the
conservation effort. In evaluating the
certainty that a conservation effort will
be effective, we may consider the
following: Does the effort describe the
nature and extent of the threats to the
species to be addressed and how these
threats are reduced by the conservation
effort? Does the effort establish specific
conservation objectives? Does the effort
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identify the appropriate steps to reduce
the threats to the species? And does the
effort include quantifiable performance
measures to monitor both compliance
and effectiveness? Overall, we need to
be certain that the formalized
conservation effort improves the status
of the species at the time we make a
listing determination. The PECE Policy
also states that last-minute agreements
(i.e., those that are developed just before
or after a species is proposed for listing)
often have little chance of affecting the
outcome of a listing decision. Lastminute efforts are also less likely to be
able to demonstrate that they will be
implemented and effective in reducing
or removing the threats to a species. In
addition, there are circumstances in
which the threats to a species are so
imminent and/or complex that is will be
almost impossible to develop an
agreement or plan that includes
conservation efforts that will result in
making the listing unnecessary. A
conservation effort that satisfies the
criteria for implementation and
effectiveness is considered when
making a listing determination, but may
not ultimately change the risk
assessment for the species. Using the
criteria identified in our PECE Policy we
evaluated conservation efforts to protect
and recover the five sawfish species that
are either underway but not yet fully
implemented, or are only planned.
CITES restricts the trade of live
animals to a vast array of wildlife
products derived from them, including
food products, musical instruments,
tourist curios and medicines. Many
wildlife species in trade are not
endangered, but the existence of an
agreement to ensure the sustainability of
the trade is important in order to
safeguard these resources for the future.
All sawfishes in the family Pristidae
were listed on Appendix I of CITES at
the 14th Conference of the Parties
meeting in 2007. An Appendix I listing
bans all commercial trade in parts (e.g.,
rostral teeth, rostra, liver, and fins) or
derivatives of sawfish with trade in
specimens of these species permitted
only in exceptional circumstances (e.g.,
for research purposes). At that time, an
annotation to the Appendix I listing
allowed the largetooth sawfish P.
microdon (herein P. pristis) to be treated
as Appendix II ‘‘for the exclusive
purpose of allowing international trade
in live animals to appropriate and
acceptable aquaria for primarily
conservation purposes.’’ The annotation
was accepted on the basis that
Australian populations of P. microdon
were robust relative to other
populations in the species’ range, and
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that the capture of individuals for
aquaria was not likely to be detrimental
to the population. Later, at the CITES
16th Annual Conference of the Parties
meeting in March of 2013, Australia
proposed the transfer of P. microdon
from Appendix II to Appendix I, and the
measure was adopted and became
effective on 12 June 2013. Therefore,
live trade of P. pristis (P. microdon) is
currently banned and all commercial
trade of all sawfishes is banned per
CITES Appendix I listing.
The recent banning of all trade of P.
pristis (P. microdon) for aquaria trade is
a good conservation measure for the
species and meets all of the criteria for
implementation and effectiveness. The
recently adopted CITES Appendix I
listing for largetooth sawfish only bans
the live trade of the fish from Australia
to approved foreign aquaria, all other
trade was banned with the 2007 listing.
Only 11 largetooth sawfish were
approved for aquaria trade when the
largetooth sawfish was listed under
CITES Appendix I with the annotation
for aquaria trade. The recent CITES
Appendix I listing for largetooth sawfish
is not likely to significantly affect the
species outside of the limited area
(Australia) where they were removed
from the wild for aquaria display. Given
live trade of P. pristis (P. microdon) for
aquaria use is not a threat leading to the
extinction risk of the species, we
conclude the full CITES Appendix I
listing may satisfy the PECE policy’s
standards for implementation and
effectiveness, but the impact of this
measure is considered insignificant.
Australia may be effective at enforcing
trade policies, but the recent Appendix
I listing of P. microdon (largetooth
sawfish) alone, is not sufficient to
protect the species throughout its range.
The IUCN Shark Specialist Group, in
collaboration with a large number of the
national and international stakeholders
in sawfish conservation, developed A
Global Strategy for Sawfish
Conservation (Harrison and Dulvy,
2014). The strategy identifies the actions
required to achieve recovery for all
sawfishes. The strategy outlines seven
objectives that are necessary to achieve
recovery of all sawfishes: Fisheries
management, species protection, habitat
conservation, trade limitation, strategic
research, education and
communication, and responsible
husbandry. We evaluated the certainty
of whether or not the strategy would be
implemented and determined that (1)
the strategy does not have a high level
of certainty that the resources necessary
to carry out the conservation effort are
available, (2) that the strategy team
members do not have the authority to
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carry out all of the objectives, (3)
regulatory or procedural mechanisms
are not in place to carry out the
objectives, (4) and the conservation
efforts rely on voluntary participation
that does not have incentives that are
included in the conservation effort that
will ensure the level of participation
necessary to effectively carry out the
conservation effort. Based on the lack of
certainty that the conservation efforts
will be implemented we determined the
strategy does not satisfy the PECE
policy’s standards for certainty of
implementation and effectiveness.
The Australian Government,
Department of the Environment,
published a Draft Recovery Plan for
Sawfish and River Sharks (Plan) in 2014
(Department of Environment, 2014). The
Draft Plan covers three sawfish species
(P. pristis, P. zijsron, and P. clavata).
The Plan identifies specific actions and
objectives necessary to stop local
decline of sawfish and river sharks and
promotes their recovery. The goal of the
Draft Plan is to assist with the recovery
of sawfish in Australian waters in two
ways: (1) Improving the population
status leading to the removal of the
sawfish from the protected species list
of EPBC; and (2) ensuring anthropogenic
actives do not hinder the recovery in the
near future, or impact the conservation
status of the species in the future. We
evaluated the certainty of whether or
not the Draft Plan would be
implemented. We determined that the
strategy has a high level of uncertainty
regarding implementation because: (1)
The Draft Plan does not have dedicated
funding so the resources necessary to
carry out the conservation efforts may
not be available, and (2) the Draft Plan
is dependent on the participation of
voluntary groups or organizations (e.g.,
indigenous community groups and nongovernmental organizations) to carry out
some of the actions. Based on the lack
of certainty that the Draft Plan will be
implemented, we determined the Draft
Plan does not satisfy the PECE policy’s
standards for certainty of
implementation and effectiveness.
Listing Determinations
Section 4(b)(1) of the ESA requires
that we make listing determinations
based solely on the best scientific and
commercial data available after
conducting a review of the status of the
species and taking into account those
efforts, if any, being made by any state
or foreign nation, or political
subdivisions thereof, to protect and
conserve the species. We have reviewed
the best available scientific and
commercial information including the
petition, and the information in the
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review of the status of the five species
of sawfishes, and we have consulted
with species experts.
We are responsible for determining
whether narrow sawfish (A. cuspidata),
dwarf sawfish (P. clavata), largetooth
sawfish (P. pristis), green sawfish (P.
zijsron), and the non-U.S. DPS of
smalltooth sawfish (P. pectinata) are
threatened or endangered under the
ESA (16 U.S.C. 1531 et seq.). We have
followed a stepwise approach as
outlined above in making this listing
determination for these five species of
sawfish. We have determined that
narrow sawfish (A. cuspidata); dwarf
sawfish (P. clavata); largetooth sawfish
(P. pristis); green sawfish (P. zijsron);
and the non-U.S. DPS of smalltooth
sawfish (P. pectinata) constitute species
as defined by the ESA. We have
conducted an extinction risk analysis
and concluded that the risk of
extinction for all five species of sawfish
is high, now and in the foreseeable
future. We have assessed the threats
affecting the status of each species using
the five factors identified in section
4(a)(1) of the ESA and concluded the
narrow, dwarf, largetooth, green, and
the non-U.S. DPS of smalltooth sawfish
face ongoing threats from habitat
alteration, overutilization for
commercial and recreational purposes,
inadequacy of existing regulatory
mechanisms, and other natural or
manmade factors affecting their
continued existence throughout their
ranges. Therefore, we find that all five
species of sawfishes are in danger of
extinction throughout all of their ranges.
After considering efforts being made to
protect these sawfishes, we could not
conclude the proposed conservation
efforts would alter the extinction risk for
any of these five sawfishes.
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Effects of Listing
Conservation measures provided for
species listed as endangered or
threatened under the ESA include
recovery actions (16 U.S.C. 1533(f));
Federal agency requirements to consult
with NMFS and to ensure its actions do
not jeopardize the species or result in
adverse modification or destruction of
critical habitat should it be designated
(16 U.S.C. 1536); designation of critical
habitat if prudent and determinable (16
U.S.C. 1533(a)(3)(A)); and prohibitions
on taking (16 U.S.C. 1538). An
additional benefit of listing beyond
these legal requirements is that the
recognition of the species’ plight
through listing promotes conservation
actions by Federal and state agencies,
foreign entities, private groups, and
individuals.
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Recovery Plans
NMFS may develop a recovery plan or
plans for these species after considering
the conservation benefit to the species
per ESA sections 4(f)(1) and 4(f)(1)(A).
Section 4 (f)(1) of the ESA directs NMFS
to develop and implement recovery
plans for the conservation and survival
of listed species, unless we find that
such a plan will not promote the
conservation of the species. Section
4(f)(1)(A) further directs us, to the
maximum extent practicable, to give
priority in developing plans to those
species that will most likely benefit
from such plans.
Identifying Section 7 Consultation
Requirements
Section 7(a)(2) (16 U.S.C. 1536(a)(2))
of the ESA and NMFS/USFWS
regulations require Federal agencies to
consult with us to ensure that activities
authorized, funded, or carried out are
not likely to jeopardize the continued
existence of listed species or destroy or
adversely modify critical habitat. The
requirement to consult applies to these
Federal agency actions in the United
States and on the high seas. The five
sawfishes all occur in the waters of
foreign nations, where there would be
no consultation requirement. It is
possible, but highly unlikely, that the
listing of the five species of sawfish
under the ESA may result in a minor
increase in the number of Section 7
consultations for high seas activities.
Critical Habitat
Critical habitat is defined in Section
3 of the ESA (16 U.S.C. 1532(5)) as: (1)
The specific areas within the
geographical area occupied by a species,
at the time it is listed in accordance
with the ESA, on which are found those
physical or biological features (a)
essential to the conservation of the
species and (b) that may require special
management considerations or
protection; and (2) specific areas outside
the geographical area occupied by a
species at the time it is listed upon a
determination that such areas are
essential for the conservation of the
species. Critical habitat shall not be
designated in foreign countries or other
areas outside U.S. jurisdiction (50 CFR
424.12 (h)).
The best available scientific and
commercial data show that the
geographical areas occupied by the
narrow sawfish (A. cuspidata), dwarf
sawfish (P. clavata), green sawfish (P.
zijsron), largetooth sawfish (P. pristis),
and the non-U.S. DPS of smalltooth
sawfish (P. pectinata) are entirely
outside U.S. jurisdiction, so we cannot
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designate critical habitat for these
species in their occupied range.
We can designate critical habitat in
unoccupied areas in U.S. jurisdiction, if
we determine the areas are essential for
the conservation of the species. Only the
largetooth sawfish (P. pristis, formerly P.
perotteti) has a range that once included
occasional use of U.S. waters, with
approximately 39 confirmed records (33
in Texas) from 1910 through 1961. All
records of P. pristis in U.S. waters were
adults, mostly during the summer
months. U.S. waters were a limited part
of the historic range, likely used for
periodic, seasonal foraging movements.
There is no evidence of U.S. waters
supporting any other biological
functions like breeding or nursery areas.
Therefore, we believe reestablishment
back into U.S. waters is not required for
the recovery of P. pristis. Based on the
best available information we have not
identified unoccupied areas in U.S.
jurisdiction that are essential to the
conservation of any of the five sawfish
species. Therefore, we do not intend to
designate critical habitat for the narrow,
dwarf, largetooth, green, or the non-U.S.
DPS of smalltooth sawfish.
Identification of Those Activities That
Would Constitute a Violation of Section
9 of the ESA
On July 1, 1994, NMFS and FWS
published a policy (59 FR 34272) that
requires us to identify, to the maximum
extent practicable at the time a species
is listed, those activities that would or
would not constitute a violation of
section 9 of the ESA. Because we are
listing all five sawfishes as endangered,
all of the prohibitions of section 9(a)(1)
of the ESA will apply to all five species.
These include prohibitions against the
import, export, use in foreign
commerce, and ‘‘take’’ of the species.
Take is defined as ‘‘to harass, harm,
pursue, hunt, shoot, wound, kill, trap,
capture, or collect, or to attempt to
engage in any such conduct.’’ These
prohibitions apply to all persons subject
to the jurisdiction of the United States,
including in the United States or on the
high seas. The intent of this policy is to
increase public awareness of the effects
of this listing on proposed and ongoing
activities within the species’ range.
Activities that we believe could result in
a violation of Section 9 prohibitions of
these five sawfishes include, but are not
limited to, the following:
(1) Take within the U.S. or its
territorial sea, or upon the high seas;
(2) Possessing, delivering,
transporting, or shipping any sawfish
part that was illegally taken;
(3) Delivering, receiving, carrying,
transporting, or shipping in interstate or
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foreign commerce any sawfish or
sawfish part, in the course of a
commercial activity, even if the original
taking of the sawfish was legal;
(4) Selling or offering for sale in
interstate commerce any sawfish part,
except antique articles at least 100 years
old;
(5) Importing or exporting sawfish or
any sawfish part to or from any country;
(6) Releasing captive sawfish into the
wild. Although sawfish held noncommercially in captivity at the time of
listing are exempt from certain
prohibitions, the individual animals are
considered listed and afforded most of
the protections of the ESA, including
most importantly the prohibitions
against injuring or killing. Release of a
captive animal has the potential to
injure or kill the animal. Of an even
greater conservation concern, the release
of a captive animal has the potential to
affect wild populations of sawfish
through introduction of diseases or
inappropriate genetic mixing.
Depending on the circumstances of the
case, NMFS may authorize the release of
a captive animal through a section
10(a)(1)(a) permit; and
(7) Engaging in experimental or
potentially injurious veterinary care or
conducting research or breeding
activities on captive sawfish, outside the
bounds of normal animal husbandry
practices. Normal care of captive
animals necessarily entails handling or
other manipulation of the animals, and
NMFS does not consider such activities
to constitute take or harassment of the
animals so long as adequate care,
including adequate veterinary care is
provided. Such veterinary care includes
confining, tranquilizing, or
anesthetizing sawfishes when such
practices, procedures, or provisions are
not likely to result in injury. Captive
breeding of sawfish is considered
experimental and potentially injurious.
Furthermore, the production of sawfish
progeny has conservation implications
(both positive and negative) for wild
populations. Experimental or
potentially injurious veterinary
procedures and research or breeding
activities of sawfish may, depending on
the circumstances, be authorized under
an ESA 10(a)(1)(a) permit for scientific
research or the enhancement of the
propagation or survival of the species.
We have identified, to the extent
known at this time, specific activities
that will not be considered likely to
result in a violation of Section 9.
Although not binding, we consider the
following actions, depending on the
circumstances, as not being prohibited
by ESA Section 9:
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(1) Take of a sawfish authorized by a
10(a)(1)(a) permit authorized by, and
carried out in accordance with the terms
and conditions of an ESA section
10(a)(1)(a) permit issued by NMFS for
purposes of scientific research or the
enhancement of the propagation or
survival of the species;
(2) Incidental take of a sawfish
resulting from Federally authorized,
funded, or conducted projects for which
consultation under section 7 of the ESA
has been completed, and when the
otherwise lawful activity is conducted
in accordance with any terms and
conditions granted by NMFS in an
incidental take statement in a biological
opinion pursuant to section 7 of the
ESA;
(3) Continued possession of sawfish
parts that were in possession at the time
of listing. Such parts may be noncommercially exported or imported;
however the importer or exporter must
be able to provide sufficient evidence to
show that the parts meet the criteria of
ESA section 9(b)(1) (i.e., held in a
controlled environment at the time of
listing, non-commercial activity);
(4) Continued possession of live
sawfish that were in captivity or in a
controlled environment (e.g., in aquaria)
at the time of this listing, so long as the
prohibitions under ESA section 9(a)(1)
are not violated. Again, facilities should
be able to provide evidence that the
sawfish were in captivity or in a
controlled environment prior to listing.
We suggest such facilities submit
information to us on the sawfish in their
possession (e.g., size, age, description of
animals, and the source and date of
acquisition) to establish their claim of
possession (see For Further Information
Contact);
(5) Provision of care for live sawfish
that were in captivity at the time of
listing. These individuals are still
protected under the ESA and may not be
killed or injured, or otherwise harmed,
and, therefore, must receive proper care.
Normal care of captive animals
necessarily entails handling or other
manipulation of the animals, and we do
not consider such activities to constitute
take or harassment of the animals so
long as adequate care, including
adequate veterinary care is provided.
Such veterinary care includes confining,
tranquilizing, or anesthetizing sawfish
when such practices, procedures, or
provisions are not likely to result in
injury; and
(6) Any importation or exportation of
live sawfish or sawfish parts with all
accompanying CITES import and export
permits and an ESA section 10(a)(1)(a)
permit for purposes of scientific
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research or the enhancement of the
propagation or survival of the species.
Section 11(f) of the ESA gives NMFS
authority to promulgate regulations that
may be appropriate to enforce the ESA.
Future regulations may be promulgated
to regulate trade or holding of sawfish,
if necessary. The public will be given
the opportunity to comment on future
proposed regulations.
Policies on Peer Review
In December 2004, the Office of
Management and Budget (OMB) issued
a Final Information Quality Bulletin for
Peer Review establishing a minimum
peer review standard. Similarly, a joint
NMFS/FWS policy (59 FR 34270; July 1,
1994) requires us to solicit independent
expert review from qualified specialists,
concurrent with the public comment
period. The intent of the joint peer
review policy is to ensure that listings
are based on the best scientific and
commercial data available. We formally
solicited expert opinion of three
appropriate and independent specialists
regarding the scientific and commercial
data or assumptions related to the
information considered for listing.
We considered peer reviewer
comments in making our determination.
We conclude that these experts’ reviews
satisfy the requirements for ‘‘adequate
[prior] peer review’’ contained in the
Information Quality Bulletin for Peer
Review and the joint NMFS/FWS policy
(59 FR 34270; July 1, 1994).
References
A complete list of the references used
in this final rule is available on the
Internet at https://sero.nmfs.noaa.gov/
protected_resources/sawfish/.
Classification
National Environmental Policy Act
The 1982 amendments to the ESA, in
section 4(b)(1)(A), restrict the
information that may be considered
when assessing species for listing. Based
on this limitation of criteria for a listing
decision and the opinion in Pacific
Legal Foundation v. Andrus, 675 F. 2d
825 (6th Cir. 1981), NMFS has
concluded that ESA listing actions are
not subject to the environmental
assessment requirements of the National
Environmental Policy Act (NEPA) (See
NOAA Administrative Order 216–6).
Executive Order 12866, Regulatory
Flexibility Act, and Paperwork
Reduction Act
As noted in the Conference Report on
the 1982 amendments to the ESA,
economic impacts cannot be considered
when assessing the status of a species.
Therefore, the economic analysis
E:\FR\FM\12DER3.SGM
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Federal Register / Vol. 79, No. 239 / Friday, December 12, 2014 / Rules and Regulations
requirements of the Regulatory
Flexibility Act are not applicable to the
listing process. In addition, this final
rule is exempt from review under
Executive Order 12866. This final rule
does not contain a collection-ofinformation requirement for the
purposes of the Paperwork Reduction
Act.
Executive Order 13132, Federalism
In accordance with E.O. 13132, we
determined that this final rule does not
have significant Federalism effects and
that a Federalism assessment is not
required.
List of Subjects in 50 CFR Part 224
Administrative practice and
procedure, Endangered and threatened
species, Exports, Imports, Reporting and
recordkeeping requirements, and
Transportation.
Dated: December 8, 2014.
Samuel D. Rauch, III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
For the reasons set out in the
preamble, 50 CFR part 224 is amended
as follows:
*
*
*
*
*
(h) The endangered species under the
jurisdiction of the Secretary of
Commerce are:
1. The authority citation for part 224
continues to read as follows:
■
Authority: 16 U.S.C. 1531–1543 and 16
U.S.C 1361 et seq.
Scientific name
*
§ 224.101 Enumeration of endangered
marine and anadromous species.
*
PART 224—ENDANGERED MARINE
AND ANADROMOUS SPECIES
Species1
Common name
2. In § 224.101, paragraph (h), amend
the table by:
■ A. Removing the ‘‘Sawfish,
largetooth’’ and the ‘‘Sawfish,
smalltooth (United States DPS)’’ entries.
■ B. Adding entries for five new sawfish
species in alphabetic order by Scientific
name under ‘‘Fishes’’:
■
Description of listed entity
*
*
Citation(s) for listing
determination(s)
*
Critical
habitat
*
ESA
rules
*
FISHES
*
Sawfish, dwarf ...................
*
*
Pristis clavata ....................
*
Entire species ...................
Sawfish, green ...................
Pristis zijsron .....................
Entire species ...................
Sawfish, largetooth ............
Pristis pristis (formerly
Pristis perotteti, Pristis
pristis, and Pristis
microdon).
Anoxypristis cuspidata ......
Entire species ...................
Sawfish, narrow .................
Sawfish, smalltooth (NonU.S. DPS).
*
Pristis pectinata ................
*
Entire species ...................
Smalltooth sawfish originating from non-U.S.
waters.
*
*
*
*
[Insert Federal Register
citation] 12/12/2014.
[Insert Federal Register
citation] 12/12/2014.
[Insert Federal Register
citation] 12/12/2014.
*
NA
NA
NA
NA
NA
NA
*
NA
NA
[Insert Federal Register
citation] 12/12/2014.
[Insert Federal Register
citation] 12/12/2014.
1 Species
*
NA
NA
*
includes taxonomic species, subspecies, distinct population segments (DPSs) (for a policy statement, see 61 FR 4722, February 7,
1996), and evolutionarily significant units (ESUs) (for a policy statement, see 56 FR 58612, November 20, 1991).
*
*
*
*
*
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Agencies
[Federal Register Volume 79, Number 239 (Friday, December 12, 2014)]
[Rules and Regulations]
[Pages 73977-74005]
From the Federal Register Online via the Government Printing Office [www.gpo.gov]
[FR Doc No: 2014-29201]
[[Page 73977]]
Vol. 79
Friday,
No. 239
December 12, 2014
Part III
Department of Commerce
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National Oceanic and Atmospheric Administration
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50 CFR Part 224
Endangered and Threatened Wildlife and Plants; Final Endangered
Listing of Five Species of Sawfish Under the Endangered Species Act;
Final Rule
Federal Register / Vol. 79 , No. 239 / Friday, December 12, 2014 /
Rules and Regulations
[[Page 73978]]
-----------------------------------------------------------------------
DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric Administration
50 CFR Part 224
[Docket No 101004485-4999-03]
RIN 0648-XZ50
Endangered and Threatened Wildlife and Plants; Final Endangered
Listing of Five Species of Sawfish Under the Endangered Species Act
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce.
ACTION: Final rule.
-----------------------------------------------------------------------
SUMMARY: We, NMFS, issue this final rule implementing our determination
that the narrow sawfish (Anoxypristis cuspidata), dwarf sawfish
(Pristis clavata), largetooth sawfish (collectively Pristis pristis;
formerly Pristis pristis, Pristis microdon, and Pristis perotteti),
green sawfish (Pristis zijsron), and the non-U.S. distinct population
segment (DPS) of smalltooth sawfish (Pristis pectinata) are endangered
species under the Endangered Species Act (ESA) of 1973, as amended. We
also include a change in the scientific name for largetooth sawfish in
this final rule to codify the taxonomic reclassification of P.
perotteti to P. pristis. We are not designating critical habitat
because the geographical areas occupied by the species are entirely
outside U.S. jurisdiction and we have not identified any unoccupied
areas within U.S. jurisdiction that are essential to the conservation
of any of the five species. We have reviewed the status of the five
species of sawfish, considered public and peer review comments, and
conservation efforts being made to protect all five species, and we
have made our determination based on the best available scientific and
commercial data that all five species of sawfish--the narrow sawfish
(Anoxypristis cuspidata), dwarf sawfish (Pristis clavata), largetooth
sawfish (collectively Pristis pristis; formerly Pristis pristis,
Pristis microdon, and Pristis perotteti), green sawfish (Pristis
zijsron), and the non-U.S. DPS of smalltooth sawfish (Pristis
pectinata)--are at risk of extinction throughout all of their ranges
and should be listed as endangered species.
DATES: This final rule is effective January 12, 2015.
ADDRESSES: Information regarding this final rule may be obtained by
contacting NMFS, Protected Resources Division, 263 13th Avenue South,
St. Petersburg, Florida, 33701. The final rule and citation list are
located on our Web site at https://sero.nmfs.noaa.gov/protected_resources/sawfish/.
FOR FURTHER INFORMATION CONTACT: Shelley Norton, NMFS, Southeast
Regional Office (727) 824-5312 or Dr. Dwayne Meadows, NMFS, Office of
Protected Resources (301) 427-8403.
SUPPLEMENTARY INFORMATION:
Background
On September 10, 2010, we received a petition from the WildEarth
Guardians (WEG) requesting we list six sawfish species--knifetooth,
narrow, or pointed sawfish (A. cuspidata), hereinafter the narrow
sawfish; dwarf or Queensland sawfish (P. clavata), hereinafter the
dwarf sawfish; largetooth sawfish (P. pristis and P. microdon); green
sawfish (P. zijsron); and the non-listed population(s) of smalltooth
sawfish (P. pectinata)--as endangered or threatened under the ESA; or
alternatively, list any distinct population segments (DPS) that exist
under the ESA. On March 7, 2011, we published a 90-day finding (76 FR
12308) stating the petitioned action may be warranted for five of the
six species. The five species were A. cuspidata, P. clavata, P.
microdon, P. zijsron, and the non-listed population(s) of P. pectinata.
Information in our records at the time indicated that P. pristis, as
described in the petition, was not a valid species. Our 90-day finding
requested information to inform our decision, and announced the
initiation of status reviews for the five species. On June 4, 2013, we
published a proposed rule (78 FR 33300) to list A. cuspidata, P.
clavata, P. pristis (formerly P. pristis, P. microdon, and P.
perotteti), P. zijsron, and the non-U.S. DPS of P. pectinata as
endangered. We also included a change in the scientific name for
largetooth sawfish in the proposed rule to codify the taxonomic
reclassification of P. perotteti to P. pristis. The largetooth sawfish
(P. perotteti) was already listed as endangered on July 12, 2011 (76 FR
40822), but this listing decision concerns the entire largetooth
sawfish (P. pristis) species as it is currently classified, which also
includes the species formerly classified as P. perotteti and P.
microdon. We did not propose to designate critical habitat because the
geographical areas occupied by the species are entirely outside U.S.
jurisdiction and we did not identify any unoccupied areas that are
currently essential to the conservation of any of these species. We
solicited public and peer reviewer comments on the proposed rule and
also coordinated outreach on the proposed rule with the Department of
State to give notice to foreign nations where the species are believed
to occur.
We are responsible for determining whether species are threatened
or endangered under the ESA (16 U.S.C. 1531 et seq.). To make this
determination, we first consider whether a group of organisms
constitutes a ``species'' under the ESA, then whether the status of the
species qualifies it for listing as either threatened or endangered.
Section 3 of the ESA defines a ``species'' as ``any subspecies of fish
or wildlife or plants, and any distinct population segment of any
species of vertebrate fish or wildlife which interbreeds when mature.''
On February 7, 1996 (61 FR 4722), NMFS and the U.S. Fish and Wildlife
Service (USFWS; collectively, the Services) adopted a policy
identifying two elements that must be considered when identifying a
DPS: (1) The discreetness of the population segment in relation to the
remainder of the species (or subspecies) to which it belongs; and (2)
the significance of the population segment to the remainder of the
species (or subspecies) to which it belongs. As stated in the DPS
policy, Congress expressed its expectation that the Services would
exercise their authority with regard to the use of DPSs sparingly and
only when the biological evidence indicates such action is warranted.
Section 3 of the ESA defines an endangered species as ``any species
which is in danger of extinction throughout all or a significant
portion of its range'' and a threatened species as one ``which is
likely to become an endangered species within the foreseeable future
throughout all or a significant portion of its range.'' Thus we
interpret an ``endangered species'' to be one that is presently in
danger of extinction. A ``threatened species,'' is not presently in
danger of extinction, but is likely to become so in the foreseeable
future (that is, at a later time). In other words, the primary
statutory difference between a threatened and endangered species is the
timing of when a species may be in danger of extinction-- either
presently (endangered) or in the foreseeable future (threatened).
Section 4(a)(1) of the ESA requires us to determine whether any
species is endangered or threatened due to any one or a combination of
the following five factors: (A) The present or threatened destruction,
modification, or curtailment of its habitat or range; (B)
overutilization for commercial, recreational, scientific, or
educational purposes; (C) disease or predation; (D)
[[Page 73979]]
the inadequacy of existing regulatory mechanisms; or (E) other natural
or manmade factors affecting its continued existence. We are required
to make listing determinations based solely on the best scientific and
commercial data available after conducting a review of the status of
the species and after taking into account efforts being made by any
state or foreign nation to protect the species.
Accordingly, we have followed a stepwise approach in making our
listing determinations for A. cuspidata, P. clavata, P. pristis
(formerly P. pristis, P. microdon, and P. perotteti), P. zijsron, and
the non-U.S.DPS of P. pectinata. For the non-U.S. DPS of P. pectinata
that may qualify as a DPS, we considered biological evidence, such as
genetic information to determine if the population met the DPS policy
criteria. Using the best available information gathered during the
status reviews, we completed an extinction risk assessment using the
general procedure of Wainwright and Kope (1999). We then assessed the
threats affecting the status of each species using the five factors
identified in section 4(a)(1) of the ESA, and then assessed public and
peer reviewer comments.
Once we determined the threats, we assessed the efforts being made
to protect each species to determine if these conservation efforts were
adequate to mitigate the existing threats and alter extinction risk. We
evaluated conservation efforts using the criteria outlined in the joint
NMFS and U.S. Fish and Wildlife Service (USFWS) Policy for Evaluating
Conservation Efforts (PECE; 68 FR 15100; March 28, 2003) to determine
the certainty of implementation and effectiveness for future
conservation efforts not yet fully implemented or effective. Finally,
we re-assessed the extinction risk of each species after considering
the existing conservation efforts.
In order to conduct a comprehensive review, NMFS Southeast Region
Protected Resources Division and NMFS Southeast Fisheries Science
Center staff members collaborated to identify the best available
information. Unlike some of our previous 12-month findings, we did not
develop a separate status review report. Instead, we presented all
information available for these species in the proposed rule, and we
present that information again, as modified by public comment on the
proposed rule, in this final rule. We first discuss background
information relative to all five species, and then we include
descriptions of the natural history specific to each species.
Sawfish General Species Description
Sawfishes are a group of shark-like rays. Taxonomically, they are
classified in the Family Pristidae (sawfishes), Order Rajiformes
(skates, rays, and sawfishes), subclass (Elasmobrancii), and Class
Chondrichthyes (cartilaginous fish). The overall body form of sawfishes
is similar to sharks, but they are flattened dorso-ventrally. Sawfishes
are covered with dermal denticles (teeth-like scales) and possess
enlarged pectoral fins.
The most distinct characteristic of sawfishes is their large, flat,
toothed rostrum or `saw' with large teeth on each side. The rostral
teeth are made from calcified tissue that is neither dentin nor enamel,
though it is more similar to the latter (Bradford, 1957). Rostral teeth
develop inside sockets on the rostrum and are held in place by strong
fibers. Unlike sharks, sawfish rostral teeth are not replaced, although
partially broken teeth may continue to grow (Miller, 1974). For some
species of sawfish, the number of rostral teeth can vary by geographic
region.
Sawfishes use their rostrum to locate, stun, and kill prey,
generally small schooling fishes such as mullet, herring, shad, and
sardines (Bigelow and Schroeder, 1953). Breder (1952), in summarizing
the literature on observations of sawfish feeding behavior, noted that
they attack fish by slashing sideways through schools of fish, and then
impale the fish on their rostral teeth. Prey are subsequently scraped
off their rostral teeth by rubbing the rostrum on the bottom and then
ingesting the whole fish. Bigelow and Schroeder (1953) also report that
sawfish feed on crustaceans and other benthic species. Recent studies
indicate that sawfishes may use their toothed rostrum to sense their
prey's electric fields (Wueringer et al., 2011; 2012).
Sawfish species are distributed primarily in circumtropical shallow
coastal waters that generally vary in salinity. While sawfishes are
commonly found in shallow water, adults are known to also inhabit
deeper waters (greater than 130 ft, 39.6 m). Some sawfishes are found
in freshwater, with established populations in major rivers and lakes
of South America, Africa, Australia, and Southeast Asia. The physical
characteristics of habitat, such as salinity and temperature, likely
influence a sawfish's movement patterns. Tides limit the physical
habitat area available, which may explain movement into shallow water
areas during specific tidal cycles (Blaber et al., 1989).
Life history data on sawfishes are limited. Fertilization is
internal by means of male claspers and reproduction is ovoviviparous;
females carry eggs with a yolk sac that nourishes developing young
until they hatch within the body. Sawfishes are born with a gelatinous
substance around their rostral teeth to protect the mother during birth
(Last and Stevens, 1994; Rainboth, 1996; Compagno and Last, 1999; Raje
and Joshi, 2003; Field et al., 2009). It is thought that most sawfishes
breed every two years and have a gestation period of about four to five
months (Bigelow and Schroeder, 1953; Thorson, 1976a). The number of
young in a litter varies by species, as does the age at sexual
maturity.
Like most chondrichthyes, sawfishes occupy the mid- to upper-level
of their food web. Smaller sawfishes, including juveniles, may be
preyed upon by larger sharks like the bull shark (Carcharhinus leucas),
estuarine crocodiles (Crocodylus porosus), or alligators (Alligator
mississippiensis). Sawfishes may use their saw as a weapon for defense
against these predators (Brewer et al., 1997; Wueringer et al., 2009).
Previously, seven valid species of sawfish were recognized
worldwide (Compagno, 1999). Compagno and Cook (1995) and Compagno
(1999) identified these seven species of sawfish as A. cuspidata Latham
1794, P. microdon Latham 1794, P. perotteti Muller and Henle 1841, P.
pristis Linnaeus 1758, P. clavata Garman 1906, P. pectinata Latham
1794, and P. zijsron Bleeker 1851. Since then, the taxonomy,
delineation, and identification of these species have proven
problematic (Oijen et al., 2007; Wiley et al., 2008; Wueringer et al.,
2009). Most recently, Faria et al. (2013) hypothesized that the
taxonomic uncertainty occurred due to several factors: many original
species descriptions were abbreviated, few holotypes are available for
examination, reference material is not available for comparison in
museum collections, and it is difficult to obtain fresh specimens
because of the infrequent captures of all sawfishes. The majority of
the confusion regarding taxonomic classification of Pristidae was
related to the species P. pristis. To resolve questions regarding the
taxonomy of pristids, Faria et al. (2013) used historical taxonomy,
external morphology, and mitochondrial DNA (mtDNA) sequences (NADH-2
loci) to conclude that sawfishes have five species in two genera: P.
pristis, P. clavata, P. pectinata, P. zijsron, and A. cuspidata. We
accept this proposed taxonomy as the best available science.
[[Page 73980]]
Natural History of the Narrow Sawfish (Anoxypristis cuspidata)
Taxonomy and Morphology
The narrow sawfish was first described by Latham in 1794 as P.
cuspidatus. It was later reclassified as Anoxypristis due to
morphological differences from Pristis that include its narrow rostral
saw, which lacks teeth on the first quarter of the saw closest to the
head in adults, as well as the distinct shape of the lower lobe of the
caudal fin (Compagno et al., 2006a). In juveniles, the portion of the
rostrum without teeth is only about one-sixth of the saw length
(Wueringer et al., 2009).
In addition, the narrow sawfish is characterized by dagger-shaped
rostral teeth (Fowler, 1941; Blegvad and Loppenthin, 1944; Compagno and
Last, 1999; Faria et al., 2013). The narrow sawfish also has a second
pair of hollow cartilaginous tubes in its rostrum that are not present
in other sawfishes. These canals contain an additional connection to
the ampullae of Lorenzini (special sensory receptors) located on the
underside of the rostrum (Wueringer et al., 2009).
Rostral tooth count varies for this species between 18 and 22 (Last
and Stevens, 1994), 24 and 28 (Hussakof, 1912), and 27-32 (Miller,
1974). The total number of teeth has been found to vary by individual,
region, and sex. Some studies report males having fewer rostral teeth
than females, while others report the opposite (Last and Stevens, 1994;
Compagno and Last, 1999). While total rostral tooth count is often
inconsistent among individuals or studies, the number of teeth an
individual has is fixed during development (Wueringer et al., 2009).
The pectoral fins of the narrow sawfish are narrow, short, and
shark-like in shape. The first dorsal fin is located posterior to the
insertion of the pelvic fins (Compagno and Last, 1999). Within the jaw,
there are 94 teeth on the upper jaw and 102 on the lower jaw (Taniuchi
et al., 1991a). The eyes are large and very close to the spiracles.
Coloration is dark grey dorsally and whitish ventrally (Fowler, 1941;
Compagno and Last, 1999).
Narrow sawfish are the only sawfish having tricuspid (three-
pointed) denticles (White and Moy-Thomas, 1941). These denticles first
appear on sawfish at 25.6 to 28 in (65 to 71 cm) total length (TL),
after they are born. In general, the narrow sawfish is considered
``naked'' because denticle coverage in adults is often sporadic and
widely spaced, usually only covering the rostrum and anterior fin
margins, making the skin appear smooth (Fowler, 1941; Gloerfelt-Tarp
and Kailola, 1984; Last and Stevens, 1994; Wueringer et al., 2009).
Narrow sawfish also have buccopharyngeal denticles (tooth-like
structures) present in their mouth. This species does not have
tubercles or thorns on their skin (Deynat, 2005).
Habitat Use and Migration
The narrow sawfish is largely euryhaline and moves between
estuarine and marine environments (Gloerfelt-Tarp and Kailola, 1984;
Last, 2002; Compagno, 2002b; Compagno et al., 2006a; Peverell, 2008).
It is generally found in inshore waters in depths of less than 130 ft
(39.6 m) with salinities between 25 and 35 parts per thousand (ppt),
spending most of its time near the substrate or in the water column
over coastal flats (Compagno and Last, 1999; Last, 2002; Peverell,
2005; Peverell, 2008; Wueringer et al., 2009). While Smith (1936)
described it as a possible freshwater species, there are only a few
reports from freshwater (Taniuchi and Shimizu, 1991; Last and Compagno,
2002; Bonfil and Abdallah, 2004; Wueringer et al., 2009). We are not
aware of any fresh or salt water tolerance studies on the species
(Compagno, 2002a; Compagno, 2002b) and conclude its habitat is
euryhaline.
In studies conducted by Peverell (2008), the narrow sawfish in the
Gulf of Carpentaria, Australia, undergo an ontogenetic shift in
habitat. Larger individuals were commonly encountered offshore, while
smaller individuals were mostly found in inshore waters. Peverell
(2008) also found females were more likely to be offshore compared to
males, at least during the months of the study (February to May). This
suggests that smaller narrow sawfish use the protection and prey
abundance found in shallow, coastal waters (Dan et al., 1994; Peverell,
2005; Peverell, 2008).
Age and Growth
Two studies have been conducted on age and growth of narrow
sawfish. Field et al. (2009) compared previously-aged vertebrae with
aged rostral teeth and found a direct correlation up to age 6. After
age 6, an individual's age was often underestimated using tooth growth
bands as the teeth become worn over time (Field et al., 2009). Peverell
(2008) then used aged vertebrae to develop more accurate growth curves
for both sexes. While the maximum observed age of narrow sawfish from
vertebrae was 9 years, the theoretical longevity was calculated at 27
years (Peverell, 2008). A 1-year-old animal has a saw length of
approximately 4.5 in (11.5 cm). Female narrow sawfish begin to mature
at 8 ft 1 in (246 cm) TL and all are mature at 15 ft 5 in (470 cm) TL;
males are mature at 8 ft (245 cm) TL (Pogonoski et al., 2002; Bonfil
and Abdallah 2004; Peverell, 2005; 2008). The maximum recorded length
of a narrow sawfish is 15 ft 5 in (4.7 m) TL, with unconfirmed records
of 20 ft (6.1 m) TL (Last and Stevens, 1994; Compagno and Last, 1999;
Pogonoski et al., 2002; Bonfil and Abdallah, 2004; Faria et al., 2013).
Reproduction
The narrow sawfish gives birth to a maximum of 23 pups in the
spring. The total length (TL) of pups at birth is between 17-24 in (43-
61 cm) (Compagno and Last, 1999; Peverell, 2005; 2008). The
reproductive cycle is assumed to be annual, with an average of 12 pups
per litter (Peverell, 2005; D'Anastasi, 2010). The number of pups is
related to female body size, as smaller females produce fewer offspring
than larger females (Compagno and Last, 1999). Preliminary genetic
research suggests that the narrow sawfish may not have multiple fathers
per litter (D'Anastasi, 2010).
Mating season may vary by geographic region. Female narrow sawfish
captured in August (dry season) in the Gulf of Carpentaria, Australia,
all contained large eggs indicating they were mature (Peverell, 2005).
Mature males were also captured in similar locations during the same
time of year (McDavitt, 2006). Although animals are sexually mature in
the dry season, mating may not occur until the rainy season in March-
May in the Indo-West Pacific (Raje and Joshi, 2003).
Age at maturity for narrow sawfish is 2 years for males and 3 years
for females (Peverell, 2008). The intrinsic rate of population increase
(rate of growth of the population) based on life history data from the
exploited population in the Gulf of Carpentaria, Australia, has been
estimated at 0.27 per year (Moreno Iturria, 2012), with a potential
population doubling time of 2.6 years.
Diet and Feeding
Narrow sawfish feed on small fish and cuttlefish (Compagno and
Last, 1999; Field et al., 2009) and likely on crustaceans, polychaetes,
and amphipods (Raje and Joshi, 2003).
Population Structure
Genetic and morphological data support the division of the global
species of narrow sawfish into populations. Based on gene sequence
data, there is a very low level of gene flow between the northern
Indian Ocean (n = 2) and west Pacific (n = 11)
[[Page 73981]]
populations. Four haplotypes (combinations of deoxyribonucleic acid
sequences or DNA) were identified: northern Indian Ocean; Indonesian;
New Guinean-Australian; and one specimen that lacked locality
information, but had a northern Indian Ocean haplotype. Specimens
collected from the Indian Ocean had a higher number of rostral teeth
per side than those collected from the western Pacific (Faria et al.,
2013).
Field et al. (2009) examined the primary chemical elements of
rostral teeth (i.e., oxygen, calcium, and phosphorous) from narrow
sawfish captured throughout Australia in an attempt to separate
subpopulations based on the isotopes of these chemicals. They found
distinctions between regions indicating two separate subpopulations
within the Gulf of Carpentaria Australia: one in the west (Northern
Territory) and one in the east (Queensland). Using isotopes to separate
elasmobranch subpopulations is in its infancy, however, and, coupled
with the limited number of samples, it is not clear whether these
results agree with the above genetic studies of population structure.
Isotopic signatures indicate the location where an animal spends most
of its time and identifies its major prey resources and do not
necessarily provide information on reproductive connectivity between
regions. Therefore, we conclude that the best available information on
isotopic signatures does not support separating narrow sawfish into
subpopulations.
Distribution and Abundance
The narrow sawfish is found throughout the eastern and western
portions of the Indian Ocean as well as much of the western Pacific
Ocean. The range once extended from as far west as the Red Sea in Egypt
and Somalia (M. McDavitt, National Legal Research Group, Inc. pers.
comm. to IUCN, London, 2012) to as far north as Honshu, Japan,
including India, Sri Lanka, and China (Blaber et al., 1994; Last and
Stevens, 1994; Compagno and Last, 1999; Compagno et al., 2006a; Van
Oijen et al., 2007). The species has also been recorded in rivers in
India, Burma, Malaysia, and Thailand (Compagno, 2002b).
While uncertain, the current status of narrow sawfish populations
across its range has declined substantially from historic levels. The
species was previously commonly reported throughout its range, but it
is now becoming rare in catches by both commercial and recreational
fishers (Brewer et al., 2006; Compagno et al., 2006a). To evaluate the
current and historic distribution and abundance of the narrow sawfish,
we conducted an extensive search of peer-reviewed publications and
technical reports, newspaper, and magazine articles. We also reviewed
records from the Global Biodiversity Information Facility (GBIF)
database (www.gbif.org). The results of that search are summarized by
major geographic region.
Indian Ocean
The earliest reports of narrow sawfish in the Indian Ocean were
from 1937 and 1938. Two sawfish were captured from the northern Indian
Ocean (no specific location was reported). A third specimen was later
caught in the same area (Blegvad and Loppenthin, 1944).
From areas in the western Indian Ocean around the Arabian Sea,
three rostra were collected in 1938: Two near Bushire, Iran, presumably
from the Gulf of Oman, and a third in Jask, Iran, also adjacent to the
Gulf of Oman (Blegvad and Loppenthin, 1944). The most extensive report
was 13 rostra from the Persian Gulf (one of those was from Iran) but it
did not include date information. Four juveniles were recorded in
Pakistan waters in 1975: Two females and two males (Faria et al.,
2013). The last published record of narrow sawfish from the western
edge of the range, in the Straits of Hormuz, was in 1997 (A. Moore, RSK
Environment Ltd., pers. comm. to IUCN, 2012).
Most records of narrow sawfish in the Indian Ocean are from the Bay
of Bengal. In 1960 and 1961, 118 sawfish, mostly narrow sawfish, were
captured during fishery surveys using gillnets and long lines (James,
1973). There are several additional records of rostra from Bangladesh
in the 1960s (Faria et al., 2013). One record from the California
Academy of Sciences is from a fish market in Bangkok, Thailand in 1961.
A narrow sawfish was used for a 1969 parasitological study in
Bangladesh, but no further information was recorded (Moravec et al.,
2006). Faria et al. (2013) also reported one specimen from 1976, as
well as 11 more records off India, but no dates were recorded. Narrow
sawfish were recorded from the Kirachi West Wharf Fish Market in
Pakistan in 1978 (GBIF Database). From 1982 to 1994, one juvenile
female, one juvenile male, and three rostra were recorded in
Pondicherry, India (Deynat, 2005). Two female neonate specimens were
recorded in Sri Lanka, and three juveniles (two males and one female)
from Malabar in Southwest India were also reported from 1982-1994
(Deynat, 2005). Between 1981 and 2000, in the Bay of Bengal, total
elasmobranch landings records are dominated by rays and include narrow
sawfish (Raje and Joshi, 2003). Landings of narrow sawfish are
currently reported from the Indian Ocean off India although they are
infrequent (K.K. Bineesh, Marine Fisheries Research Institute,
Department of Pelagic Fisheries, India, pers. comm. to IUCN, 2012).
Indo-Pacific Ocean (excluding Australia)
There are several accounts of narrow sawfish over time from various
unspecified locations throughout the Indo-Pacific. One narrow sawfish
specimen was recorded from Mabe, India in 1835, making it the oldest
museum record from the region (GBIF Database). The first records of
narrow sawfish were for juvenile males in 1852 and 1854 (Faria et al.,
2013). A female and male were recorded in 1867, but no exact location
was specified (Faria et al., 2013). In 1879, one male and one female
were also recorded from Indonesia and four rostra were reported from
China in 1898 (Faria et al., 2013).
The next reports of narrow sawfish from the Indo-Pacific occurred
in the 1930s. A female was reported in 1931 in Indonesia (no specific
location), and a male was reported in Singapore in 1937 (Blegvad and
Loppenthin, 1944). A narrow sawfish was caught in the Gulf of Thailand
in March 1937 (Blegvad and Loppenthin, 1944). A single report from
Papua New Guinea was recorded in 1938 (Faria et al., 2013). In 1945,
narrow sawfish were reported in the Chao Phraya River, Thailand and its
tributaries (Smith, 1945). In 1952, two females were captured from
Batavia, Semarang, Indonesia along with a third female without a
rostrum (Van Oijen et al., 2007).
Records of narrow sawfish throughout the Indo-Pacific were
scattered and infrequent throughout the 1950s. Faria et al. (2013)
recorded rostra from Papua New Guinea; two from 1955 and one each from
1966, 1980, and 2000. A male was caught in 1989 from the Oriomo River,
Papua New Guinea (Taniuchi et al., 1991b; Taniuchi and Shimizu, 1991;
Taniuchi, 2002). There are other reports of narrow sawfish from Papua
New Guinea around the Gulf of Papua and in Bootless Bay from the 1970s,
but there are no recent records (Taniuchi et al., 1991b). In a
comprehensive literature search for the period 1923 to 1996 on the
biodiversity of elasmobranchs in the South China Sea, Compagno (2002a)
found no records of sawfishes. Yet, fresh dorsal and caudal fins of
narrow sawfish were found during a survey of fish markets from 1996 to
1997 in Thailand (Manjaji, 2002b).
[[Page 73982]]
There are even fewer records of narrow sawfish from the Indo-
Pacific over the last few decades. The only known specimen in the
twenty-first century is a single report from New Guinea in 2001 (L.
Harrison, IUCN, pers. comm. to John Carlson, NMFS, 2012).
Australia
Australia may have larger populations of narrow sawfish than any
other area within the species' range (Peverell, 2005). According to the
GBIF Database for Australia flora and fauna, the first museum record of
the narrow sawfish in Australia is from the Australia Museum in
Townsville, Queensland in 1963. This database also lists observations
of narrow sawfish throughout the 1980s, mostly recorded by the
Commonwealth Scientific and Industrial Research Organization (CSIRO)
Marine and Atmospheric Research group. One individual was observed in
Western Australia in 1982 and in 1983. In 1984, CSIRO observed one
narrow sawfish just west of Darwin, Northern Territory, and five in the
Gulf of Carpentaria (three in the east and two in the northwest). Five
additional records in 1984 were from the northwest tip of the western
Gulf of Carpentaria, one from outside the Daly River, and three outside
of Kakadu National Park. In 1985, two narrow sawfish were observed near
Marchinbar Island, Northern Territory. In the eastern Gulf of
Carpentaria, four narrow sawfish were observed in 1986, with single
observations in 1987 and 1988. In 1988, a narrow sawfish was observed
in Western Australia. Two narrow sawfish were reported from the Gulf of
Carpentaria in 1990 (Blaber et al., 1994). Single specimens were
captured in 1991 from the west coast of Australia (Alexander, 1991),
the Gulf of Carpentaria in 1995 (Brewer et al., 1997), and the Arafura
Sea in 1999 (Beveridge et al., 2005). Faria et al. (2013) reported
three rostra records from private collections in Australia from 1998-
1999, but no other information on the collection location was reported.
Narrow sawfish have been reported in multiple studies between 2000
and 2011, mostly from northern Australia. In a bycatch reduction device
study conducted in 2001 in the Gulf of Carpentaria, 25 narrow sawfish
were captured in trawling gear (Brewer et al., 2006). Later in 2001, a
bycatch reduction device study conducted in the Queensland shallow-
water eastern king prawn (Penaeus plebejus) trawl fishery did not
capture a single specimen (Courtney et al., 2006). The European
Molecular Biology Lab recorded narrow sawfish in 2003 in the Northern
Territory (GBIF database). A review of fisheries data and records from
2000 to 2002, identified 74 offshore and 37 inshore records of narrow
sawfish in the Gulf of Carpentaria (Peverell, 2005). Between April 2004
and April 2005, 16 narrow sawfish were caught in the Gulf of
Carpentaria during a trawl bycatch study; the mean catch rate was 0.16
sawfish per hour (Dell et al., 2009). Observers on commercial fishing
boats recorded nine captures of narrow sawfish in 2007 within the Great
Barrier Reef World Heritage Area, Queensland, which accounted for 0.86
percent of the shark and ray catch in the commercial fisheries
(Williams, 2007). Observers in the Northern Territory's Offshore Net
and Line Fishery encountered several narrow sawfish from 2007 to 2010
(Davies, 2010). Data from the Kimberley (R. McAuley, Department of
Fisheries, Western Australia, pers. comm. to Colin Simpfendorfer,
2012), the Northern Territory (Field et al., 2009), the Gulf of
Carpentaria (Peverell, 2005), and parts of the Queensland east coast
(Harry et al., 2011) suggest viable subpopulations may remain locally,
but at significantly lower levels compared to historic levels.
In summary, it appears the current range of narrow sawfish is
restricted largely to Australia. Narrow sawfish are considered very
rare in many places where evidence is available, including parts of
India (Roy, 2010), Bangladesh (Roy, 2010), Burma (FIRMS, 2007-2012),
Malaysia (including Borneo; Almada-Villela, 2002; Manjaji, 2002),
Indonesia (White and Kyne, 2010), Thailand (CITES, 2007; Compagno,
2002a; Vidthayanon, 2002), and Singapore (CITES, 2007). In Australia,
narrow sawfish are primarily located in the north. The most recent
museum record for narrow sawfish in southern Australia was from New
South Wales in the 1970s (Pogonoski et al., 2002). Data from the
Queensland Shark Control Program, conducted along the east coast of
Queensland, from 1969 to 2003 show a clear decline in sawfish catch
(although not species-specific) with the complete disappearance of
sawfish in southern regions of Queensland by 1993 (Stevens et al.,
2005). Although we cannot rule out underreporting of narrow sawfish,
especially in remote areas of its historic range, we conclude from the
consistent lack of records that narrow sawfish have been severely
depleted in numbers and their range has contracted.
Natural History of Dwarf Sawfish (Pristis clavata)
Taxonomy and Morphology
Due to its size and the geographic location where it was described,
P. clavata is referred to as the dwarf or the Queensland sawfish. The
species was first described by Garman in 1906; however, it has often
been confused with largetooth sawfish (Last and Stevens, 1994; Cook et
al., 2006; Morgan et al., 2010a). This species can be distinguished
from largetooth sawfish based on rostral tooth morphology (Thorburn et
al., 2007).
The dwarf sawfish is olive brown in color dorsally with a white
underside. The rostrum of this species is quite short, with 19 to 23
rostral teeth that are moderately flattened, elongated, and peg-like.
Studies indicate that this species does not display significant
differences in the number of rostral teeth between males (19 to 23
teeth) and females (20 to 23 teeth) (Ishihara et al., 1991a; Thorburn
et al., 2008; Morgan et al., 2010a; Morgan et al., 2011). The rostrum
makes up 21 to 26 percent of the total length of the dwarf sawfish
(Blaber et al., 1989; Grant, 1991; Last and Stevens, 1994; Compagno and
Last, 1999; Larson et al., 2006; Wueringer et al., 2009; Morgan et al.,
2011).
Morphologically, the origin of the first dorsal fin is slightly
posterior to the insertion of the pelvic fins, and the second dorsal
fin is smaller than the first. The pectoral fins are small compared to
other sawfish species, and are ``poorly developed'' (Ishihara et al.,
1991a). There is no lower lobe on the caudal fin. Lateral and low keels
are present along the base of the tail (Compagno and Last, 1999;
Wueringer et al., 2009; Morgan et al., 2010a; Morgan et al., 2011).
Within the mouth are 82-84 tooth rows on the upper jaw. The total
vertebrae number is 225-231. The dwarf sawfish has regularly
overlapping monocuspidate denticles on its skin. As a result, there are
no keels or furrows formed on the skin (Fowler, 1941; Last and Stevens,
1994; Deynat, 2005).
Habitat Use and Migration
The dwarf sawfish has been found along tropical coasts in marine
and estuarine waters, mostly from northern Australia; it may inhabit
similar habitats in other areas. Dwarf sawfish are reported on mudflats
in water 6 ft 7 in to 9 ft 10 in (2 to 3 m) deep that is often turbid
and influenced heavily by tides. Thorburn et al. (2008) reported dwarf
sawfish occur in waters 2 to 22 ft (0.7 to 7 m) deep, while Stevens et
al. (2008) recorded a maximum depth of 65 ft (20 m). This species has
also been reported in rivers (Last and Stevens, 1994; Wueringer et al.,
2009; Morgan et al., 2010a) and as commonly occurring in both brackish
and freshwater, and in both marine and estuarine habitats (Rainboth,
1996; Thorburn et al., 2008).
[[Page 73983]]
For example, two dwarf sawfish were found 31 miles (50 km) upstream
from the mouth of the south Alligator River, Kakadu National Park,
Northern Territory, Australia in 2013 at salinities of 0.12 and 7.64
ppt (P. Kyne, Charles Darwin University, pers. comm. to S. Norton,
NMFS, June 2013).
Juvenile dwarf sawfish may use the estuaries associated with the
Fitzroy River, Australia as nursery habitat for up to three years
(Thorburn et al., 2008). Dwarf sawfish are also known to use the Gulf
of Carpentaria, Australia as nursery area in a variety of habitats
(Gorham, 2006). However, physical characteristics such as salinity,
temperature, and turbidity may limit seasonal movements (Blaber et al.,
1989).
Age and Growth
Dwarf sawfish are considered to be small compared to other
sawfishes. Their maximum size has been reported as 4 ft 11 in (1.5 m)
total length (TL) (Grant, 1991) and 4 ft 7 in (140 cm) TL (Last and
Stevens, 1994; Rainboth, 1996; Compagno and Last, 1999). But more
recently, much larger sizes have been reported, as high as 19.7 ft
(6000 cm) TL (Peverell, 2005). Specimens from Western Australia in 2008
indicate that females reach at least 10 ft 2 in (310 cm) TL (Morgan et
al., 2010a; Morgan et al., 2011).
Thorburn et al. (2008) and Peverell (2008) estimated age and growth
for this species based on the number of vertebral rings and total
length. The average growth estimates for dwarf sawfish are 16.1 in
(41cm) TL in the first year, slowing to 9.4 in (24 cm) in the second
year (Peverell 2008). Thorburn et al. (2008) determined that animals
close to 3 ft (90 cm) TL were age 1, those between 3.5 and 4 ft (110 cm
and 120 cm) TL were age 2, and those around 5 ft (160 cm) TL were age
6. Peverell (2008) reported dwarf sawfish between 2 ft 11 in and 3 ft 3
in (90 and 98 cm) TL were age 0, those between 3 ft 7 in and 5 ft 9 in
(110 to 175 cm) TL were considered 1 to 3 years old, and those between
6 ft 7 in and 8 ft (201 to 244 cm) TL were considered 4 to 6 years old
(Peverell, 2008). Any dwarf sawfish over 9 ft 10 in (300 cm) TL is
considered to be at least 9 years old (Morgan et al., 2010a). The
theoretical maximum age calculated from von Bertalanffy parameters for
dwarf sawfish is 94 years (Peverell, 2008).
Reproduction
There is little information available regarding the time or
location of dwarf sawfish mating. It is hypothesized that dwarf sawfish
move into estuarine or fresh waters to breed during the wet season
(Larson et al., 2006), although no information on pupping habitat,
gestation period, or litter size has been recorded (Morgan et al.,
2010a).
Dwarf sawfish are born between 2 ft 2 in and 2 ft 8 in (65 cm and
81 cm) TL (Morgan et al., 2010a; Morgan et al., 2011). Males become
sexually mature between 9 ft 8 in and 10 ft (295 and 306 cm) TL with
fully calcified claspers, though they may mature at smaller sizes,
around 8 ft 5 in (255-260 cm) TL (Peverell, 2005; Thorburn et al.,
2008; Last and Stevens, 2009; Morgan et al., 2011). All males captured
by Thorburn et al. (2008) less than 7 ft 5 in (226 cm) TL were
immature; two females, both smaller than 3 ft 11 in (120 cm) TL, were
also immature. There is little specific information about sexual
maturation of females; females are considered immature at 6 ft 11 in
(210 cm) TL (Peverell, 2005; Peverell, 2008; Morgan et al., 2010a).
Wueringer et al. (2009) indicates that neither males nor females are
mature before 7 ft 8 in (233 cm) TL.
Intrinsic rates of population increase, based on life history data
from Peverell (2008), has been estimated to be about 0.10 per year
(Moreno Iturria, 2012), with a potential population doubling time of
7.2 years.
Diet and Feeding
Dwarf sawfish, like other sawfishes, use their saw to stun small
schooling fishes. They may also use the saw for rooting in the mud and
sand for crustaceans and mollusks (Breder Jr., 1952; Raje and Joshi,
2003; Larson et al., 2006; Last and Stevens, 2009). In Western
Australia, the dwarf sawfish eats shrimp (Natantia spp.), mullet
(Mugilidae), herring (Clupeidae), and croaker (Sciaenidae) (Thorburn et
al., 2008; Morgan et al., 2010a).
Population Structure
Phillips et al. (2011) conducted a genetic study looking at mtDNA
of dwarf sawfish and found no distinct difference in dwarf sawfish from
Western Australia and those from the Gulf of Carpentaria in northern
Australia. The genetic diversity of this species was moderate overall;
however, dwarf sawfish from the Gulf of Carpentaria may have a lower
genetic diversity than those of the west coast, possibly due to either
a small sample size or a reduction in abundance (Phillips et al.,
2008). Further declines in abundance as well as genetic drift may
result in reduced genetic diversity (Morgan et al., 2010a; 2011).
Phillips et al. (2011) determined the populations of the dwarf
sawfish are organized matrilineally (from mother to daughter),
indicating the possibility that females are philopatric (return to
their birth place). While the genetic diversity of this species is
considered low to moderate across Australia, haplotype diversity in the
Gulf of Carpentaria was very low, but was greater in the west compared
to the east. Low diversity among and within groups of dwarf sawfish may
be detrimental (Phillips et al., 2011).
Distribution and Abundance
Dwarf sawfish are thought to historically occur in the Indo-
Pacific, western Pacific, and eastern Indian Oceans, with the
population largely occurring in northern Australia (Last and Stevens,
1994; Last and Compagno, 2002; Compagno, 2002a; Compagno, 2002b;
Thorburn et al., 2008; Wueringer et al., 2009; Morgan et al., 2010a;
Kyne et al., 2013). While dwarf sawfish may have been historically more
widespread throughout the Indo-West Pacific (Compagno and Last, 1999;
Last and Stevens, 2009), there are questions regarding records outside
of Australian waters (DSEWPaC 2011; Kyne et al., 2013; GBIF database).
In an effort to gather more information on the species' historic
and current range and abundance, we conducted an extensive search of
peer-reviewed publications and technical reports, newspaper, and
magazine articles. We also reviewed records from the Global
Biodiversity Information Facility (GBIF) Database (www.gbif.com). A
summary of those findings is presented by major geographic region.
Indian Ocean
Dwarf sawfish are considered extremely rare in the Indian Ocean and
there are few records indicating its current presence (Last, 2002).
Faria et al. (2013) report a female from the R[eacute]union Islands, a
female from an unidentified location in the Indian Ocean, and a museum
record of a male from Bay of Bengal, India. A sawfish was landed at a
port in Arabian Peninsula (presumably caught in the Gulf of Oman or the
Arabian Gulf) in January of 2006. It may have been a dwarf sawfish, but
identification could not be confirmed (Kyne et al., 2013). There are no
reports of dwarf sawfish from Sri Lanka in more than a decade, although
they have been assumed to occur there (Last, 2002).
Indo-Pacific (excluding Australia)
Dwarf sawfish are considered very rare in Indonesia, with only a
few records (Last, 2002). Faria et al. (2013) compiled most reports of
dwarf sawfish in Indonesia; since the first record in 1894 from Borneo,
there have been two
[[Page 73984]]
rostral saws in 1910 and five other rostra without date or length
information. There is also one museum record of a dwarf sawfish from
Papua New Guinea in 1828 (Kyne et al., 2013).
Although reported historically, dwarf sawfish have not been found
in any other areas in the Indo-Pacific in over a decade. Rainboth's
(1996) guide to fishes of the Mekong reported a dwarf sawfish from the
Mekong River Basin, Laos, in the early 1900s but no specimen exists to
confirm this report. No sawfish of any species, including the dwarf
sawfish, were reported from the South China Sea from 1923-1996
(Compagno, 2002a). Faria et al. (2013) reported on two specimens from
the Pacific Ocean, but no specifics were provided.
Australia
The northern coast of Australia represents the geographic center of
dwarf sawfish range that extends from Cape York, Queensland west to the
Pilbara area in Western Australia (Compagno and Last, 1999; Last and
Stevens, 2009; Kyne et al., 2013). Dwarf sawfish may have occurred as
far south as Cairns, but reports are lacking. Most records for dwarf
sawfish are from the north and northwest areas of Australia.
The earliest record of dwarf sawfish in Australia is from 1877, but
no specific location was recorded (Faria et al., 2013). A single
rostrum from a dwarf sawfish was found in 1916, but no other
information was recorded. In 1945, a single specimen was reported from
the Northern Territory, Australia (Stevens et al., 2005). There is a
single record of a dwarf sawfish from the Victoria River in 1964 that
is currently housed at the Museum Victoria (GBIF Database).
Five female and five male dwarf sawfish (32 to 55 in; 82 to 140 cm
TL) were captured in 1990 in the Pentecost River using gillnets
(Taniuchi and Shimizu, 1991; Taniuchi, 2002). CSIRO recorded five dwarf
sawfish in Western Australia in 1990 (GBIF Database). CSIRO also found
one dwarf sawfish in Walker Creek (a tributary of the Gulf of
Carpentaria) in 1991 (GBIF Database). In 1992, one specimen was found
near Darwin, Northern Territory, Australia (GBIF Database). Between
1994 and 2010, almost 75 tissue samples were taken from live dwarf
sawfish or dried rostra from the Gulf of Carpentaria and the northwest
coast of Australia (Phillips et al., 2011). In 1997, two specimens were
collected near the mouth of Buffalo Creek in Darwin, Northern Territory
(Chisholm and Whittington, 2000). In 2005, Naylor et al. (2005)
collected one dwarf sawfish from Darwin, Australia. One dwarf sawfish
was captured in 1998 in the upper reaches of the Keep River Estuary
(Larson, 1999; Gunn et al., 2010). CSIRO reported one dwarf sawfish in
Western Australia (GBIF Database). In 2006, the European Molecular
Biology Lab reported the occurrence of three dwarf sawfish in Western
Australia (GBIF Database). One interaction was reported between 2007
and 2010 by observers in the Northern Territory Offshore Net and Line
Fishery (Davies, 2010). A single specimen from Queensland (northeastern
Australia) is preserved at the Harvard Museum of Comparative Zoology
(Fowler, 1941).
In a comprehensive survey of the Gulf of Carpentaria from 2001 to
2002 (Peverell, 2005; 2008), indicated dwarf sawfish were concentrated
in the west where 12 males and 10 females were captured. Most
individuals caught in the inshore fishery were immature except for two
mature males: 10 ft and 9 ft 8 in (306 cm and 296 cm) TL (Peverell,
2005; 2008).
Within specific riverine basins in northwestern Australia, dwarf
sawfish have been reported in various surveys. Forty-four dwarf sawfish
were captured between October 2002 and July 2004, in the King Sound and
the Robison, May, and Fitzroy Rivers (Thorburn et al., 2008). Between
2001 and 2002, one dwarf sawfish was caught at the mouth of the Fitzroy
River in Western Australia (Morgan et al., 2004). Morgan et al. (2011)
acquired 109 rostra from dwarf sawfish from the King Sound area that
were part of museum or personal collections.
In summary, there is some uncertainty in the species identification
of historic records of dwarf sawfish, however, it appears the dwarf
sawfish has become extirpated from much of the Indo-Pacific region and
from the eastern coast of Australia. An October 2001 study on the
effectiveness of turtle-excluder devices in the prawn trawl fishery in
Queensland, Australia, reported no dwarf sawfish (Courtney et al.,
2006). Dwarf sawfish are now considered rare in the Gulf of
Carpentaria. It is likely the Kimberley region and Pilbara region
(Western Australia) may be the last remaining areas for dwarf sawfish
(P. Kyne, Charles Darwin University, pers. comm. to IUCN, 2012).
Natural History of the Largetooth Sawfish (Pristis pristis)
Taxonomy and Morphology
Many taxonomists have suggested classification of largetooth
sawfish into a single circumtropical species given common morphological
features of robust rostrum, origin of first dorsal fin anterior to
origin of pelvic fins, and presence of a caudal-fin lower lobe
(G[uuml]nther, 1870; Garman, 1913; Fowler, 1936; Poll, 1951; Dingerkus,
1983; Daget, 1984; S[eacute]ret and McEachran, 1986; McEachran and
Fechhelm, 1998; Carvalho et al., 2007). The recent analysis by Faria et
al. (2013) used mtDNA (mitochondrial deoxyribonucleic acid) and
contemporary genetic analysis to argue that the previously classified
P. pristis, P. microdon, and P. perotteti should now be considered one
species named P. pristis. After reviewing Faria et al. (2013) and
consulting other sawfish experts, we conclude, based on the best
available information, that P. pristis applies to all the largetooth
sawfishes previously identified as P. pristis, P. microdon, and P.
perotteti.
The largetooth sawfish has a robust rostrum, noticeably widening
posteriorly (width between the two posterior-most rostral teeth is 1.7
to 2 times the width between the second anterior-most rostral teeth).
Rostral tooth counts are between 14 and 23 per side with grooves on the
posterior margin. The body is robust with the origin of the first
dorsal-fin anterior to the origin of the pelvic fin; dorsal fins are
high and pointed with the height of the second dorsal fin greater than
the first. The lower lobe of the caudal-fin is small, but well-defined,
with the lower anterior margin about half as long as the upper anterior
margin (Wallace, 1967; Taniuchi et al., 1991a; Last and Stevens, 1994;
Compagno and Last, 1999; Deynat, 2005; Wueringer et al., 2009; Morgan
et al., 2010a; Morgan et al., 2010b; Morgan et al., 2011). The
largetooth sawfish has buccopharyngeal denticles and regularly
overlapping monocuspidate dermal denticles on its skin. The denticles
are present on both dorsal and ventral portions of the body (Wallace,
1967; Deynat, 2005). Within the mouth, there are between 70 and 72
tooth rows on the upper jaw, and 64 to 68 tooth rows on the lower jaw.
The number of vertebrae is between 226 and 228 (Morgan et al., 2010a).
Coloration of the largetooth sawfish is a reddish brown dorsally and
dull white ventrally (Fowler, 1941; Wallace, 1967; Compagno et al.,
1989; Taniuchi et al., 1991a; Compagno and Last, 1999; Chidlow, 2007).
Male and female largetooth sawfish differ in the number of rostral
teeth. Using largetooth sawfish teeth collected from Papua New Guinea
and Australia, Ishihara et al. (1991b) found males to have an average
of 21 rostral teeth on the left and 22 on the right; females averaged
19 rostral teeth on both the left and the right side of the rostrum.
[[Page 73985]]
Rostrum length can vary between males and females (Wueringer et al.,
2009).
Habitat Use and Migration
Largetooth sawfish are found in coastal and inshore waters and are
considered euryhaline (Compagno et al., 1989; Last and Stevens, 1994;
Compagno and Last, 1999; Chisholm and Whittington, 2000; Last, 2002;
Compagno, 2002b; Peverell, 2005; Peverell, 2008; Wueringer et al.,
2009), being found in salinities ranging from 0 to 40 ppt (Thorburn et
al., 2007). The species has been found far upriver, often occupying
freshwater lakes and pools; they are associated with freshwater more
than any other sawfish species (Last and Stevens, 1994; Rainboth, 1996;
Peter and Tan, 1997; Compagno and Last, 1999; Larson, 1999). Largetooth
sawfish have even been observed in isolated fresh water billabongs or
pools until floodwaters allow them to escape; juveniles often use these
areas for multiple years as deepwater refuges (Gorham, 2006; Thorburn
et al., 2007; Wueringer et al., 2009; Morgan et al., 2010b). Similarly,
largetooth sawfish have been found in Lake Nicaragua in depths up to
400 ft (122 m) and are found in deeper holes, occupying muddy or sandy
bottoms (Thorson, 1982). Adults more often use marine habitats than
juveniles, and are typically found in waters with salinity at 31 ppt
(Wueringer et al., 2009).
Despite the variety of habitats occupied, females have been found
to be highly philopatric as indicated by mtDNA studies, while males
often undergo long movements (Lack et al., 2009; Phillips et al., 2009;
Morgan et al., 2010a; Morgan et al., 2010b; Morgan et al., 2011).
Largetooth sawfish occurred from the Caribbean and Gulf of Mexico south
through Brazil, and in the United States, largetooth sawfish were
reported in the Gulf of Mexico, mainly along the Texas coast (NMFS,
2010a). Largetooth sawfish were rarely reported in U.S. waters and may
have been long-distance migrants from the Caribbean or Brazil (Feldheim
et al., 2011).
The physical characteristics of habitat strongly influence the
movements of, and areas used by, largetooth sawfish. Recruitment of
neonate largetooth sawfish was correlated with the rise in water levels
during the wet season in Australia (Whitty et al., 2009). A study of
juvenile largetooth sawfish movements in the Fitzroy River in Australia
found young-of-the-year using extremely shallow areas (0 to 1 ft 7 in
or 0 to 0.49 m) up to 80 percent of the time, mostly to avoid predators
(Thorburn et al., 2007). Juvenile and adult largetooth sawfish also use
rivers (Compagno, 2002b; Gorham, 2006) and can be found in areas up to
248.5 miles (400 km) upstream (Morgan et al., 2004; Chidlow, 2007). The
space used on a day to day basis by largetooth sawfish increases with
body length (Whitty et al., 2009).
Age and Growth
There are several age and growth studies for the largetooth
sawfish; results vary due to differences in aging techniques, data
collection, or location. In Australia, largetooth sawfish are between 2
ft 6 in and 3 ft (76 and 91 cm) TL at birth, with females being
slightly smaller than males on average (Chidlow, 2007; Morgan et al.,
2011). Thorson (1982) found pups at birth average 2 ft 4.7 in to 2 ft
7.5 in (73-80 cm) TL, with a growth rate of 1 ft 2 in to 1 ft 3 in (35-
40) cm per year in Lake Nicaragua (NMFS, 2010a; Kyne and Feutry, 2013).
Peverell (2008) found that largetooth sawfish in the Indo-West Pacific
are born at 2 ft 4 in to 2 ft 11 in (72-90 cm) TL. Juveniles (age 1 to
age at maturity) range in size from 2 ft 6 in to 9 ft (76 to 277 cm) TL
(Morgan et al., 2011).
Size at maturity in the Western Atlantic is estimated to be around
9 ft 10 in (300 cm) TL for both sexes at around age 8 (Lack et al.,
2009; Morgan et al., 2010a; Morgan et al., 2010b; NMFS, 2010; Morgan et
al., 2011; Kyne and Feutry, 2013). Thorson (1982) estimated age of
maturity to be 10 years at 9 ft 10 in (300 cm) TL in Lake Nicaragua.
Peverell (2008) estimated age at maturity in the Gulf of Carpentaria to
be between 8 and 10 years. In the Indo-Pacific, males tend to mature
earlier than other regions (9 ft 2 in (280 cm)) TL (Kyne and Feutry,
2013). Generally, males under 7 ft 7 in (230 cm) TL and females under 8
ft 10 in (270 cm) TL are considered immature (Whitty et al., 2009;
Wueringer et al., 2009).
The largest recorded length of a largetooth sawfish is 22 ft 11 in
(700 cm) TL (Compagno et al., 1989. The largest largetooth sawfish
recorded in the Kimberley, Queensland measured 21 ft 6 in (656 cm) TL
(Compagno and Last, 1999). In other areas of Australia, largetooth
sawfish can reach up to 15 ft (457 cm) and at least 11 ft 10 in (361
cm) TL (Fowler, 1941; Chidlow, 2007; Gunn et al., 2010). Thorson (1982)
estimated that largetooth sawfish in Lake Nicaragua only reach a
maximum size of about 14 ft 1 in (430 cm) TL.
Age and growth for largetooth sawfish has been estimated by Tanaka
(1991) who generated a von Bertalanffy growth model for specimens
collected from Papua New Guinea and Australia. For both sexes combined,
the theoretical maximum size (L[infin]) from the von Bertalanffy growth
equation was calculated at 11 ft 11 in (363 cm) TL with a growth rate
(K) of 0.066 per year. Largetooth sawfish grow around 7 in (18 cm) in
the first year and 4 in (10 cm) by the tenth year (Tanaka, 1991).
Thorson (1982a) estimated an early juvenile growth rate of 13-15 in (35
to 40 cm) per year and annual adult growth rate of 1 in (4.4 cm) per
year based on largetooth from Lake Nicaragua. Simpfendorfer (2000)
estimated the theoretical maximum size of largetooth sawfish to be 14
ft 11 in (456 cm) TL with a growth rate (Brody growth coefficient K) of
0.089 per year based on Thorson's (1982) data from Lake Nicaragua.
Peverell (2008) calculated that largetooth sawfish from the Gulf of
Carpentaria, Australia grow 1 ft 8.5 in (52 cm) in the first year and 7
in (17 cm) during the fifth year. Maximum size was estimated at 20 ft
11 in (638 cm) TL with a growth rate (Brody growth coefficient K) of
0.08 per year from the von Bertalanffy equation (Peverell, 2008). Kyne
and Feutry (2013) summarize maximum age estimates of 30 years in Lake
Nicaragua and 35 years in the Gulf of Carpentaria. Based on the von
Bertalanffy equation, growth slows at about 35 years or 19 ft 10 in
(606 cm) TL (Kyne and Feutry, 2013).
Reproduction
Largetooth sawfish are thought to reproduce in freshwater
environments (Compagno and Last, 1999; Last, 2002; Compagno, 2002b;
Martin, 2005; Thorburn and Morgan, 2005; Compagno et al., 2006b).
Pupping seems to vary across the range, occurring during the wet season
from May to July in the Indo-Pacific (Raje and Joshi, 2003), and from
October to December in the western Atlantic and Lake Nicaragua
(Thorson, 1976a; Kyne and Feutry, 2013).
The number of pups in a largetooth sawfish litter varies by
location, possibly due to a number of factors. One of the earliest
reproductive studies on largetooth sawfish by Thorson (1976a) reported
the litter sizes of 67 females ranged between 1 to 13 pups and an
embryonic sex ratio for this species is 0.86 males for every 1 female.
Average number of pups is 7 (NMFS, 2010a; Kyne and Feutry, 2013).
Thorson (1976a) also found that both ovaries appeared to be functional,
with the left ovary producing more eggs. Estimates of litter size from
other studies in the Indo-West Pacific (e.g., Wilson, 1999; Moreno
Iturria, 2012; Peverell, 2005) cannot be confirmed (Kyne and Feutry,
2013). Length of gestation for largetooth sawfish is approximately five
months in Lake Nicaragua, with a biennial
[[Page 73986]]
reproduction cycle (Thorson 1976a; NMFS 2010a; Kyne and Feutry, 2013).
In the Indo-West Pacific, largetooth sawfish may reproduce every year
(Peverell, 2008).
Intrinsic rates of population growth vary tremendously throughout
the species' range. Simpfendorfer (2000) estimated that the largetooth
sawfish in Lake Nicaragua had an intrinsic rate of population growth of
0.05 to 0.07 per year, with a potential population doubling time of
10.3 to 13.6 years. Using data from Australia, rates of population
increase for the Indo-Pacific were estimated to be around 0.12 per year
(Moreno Iturria, 2012), with a population doubling time of
approximately 5.8 years and a generation time of 14.6 years. Data from
the western Atlantic Ocean indicate an intrinsic rate of increase of
0.03 per year, with a population doubling time of 23.3 years and a
generation time of 17.2 years (Moreno Iturria, 2012). Annual natural
mortality for the western Atlantic has been estimated at 0.07 to 0.16
(Simpfendorfer, 2000) and 0.14 to 0.15 per year (Moreno Iturria, 2012).
Diet and Feeding
Largetooth sawfish diet is predominantly fish, but varies depending
on geographic area. Small fishes including seer fish, mackerels, ribbon
fish, sciaenids, and pomfrets are likely main diet items of largetooth
sawfish in the Indian Ocean (Devadoss, 1978; Rainboth, 1996; Raje and
Joshi, 2003). Small sharks, mollusks, and crustaceans are also
potential prey items (Devadoss, 1978; Rainboth, 1996; Raje and Joshi,
2003). Taniuchi et al. (1991a) found small fishes and shrimp in the
stomachs of juveniles in Lake Murray, Papua New Guinea, while juveniles
in Western Australia had catfish, cherabin, mollusks, and insect parts
in their stomachs (Thorburn et al., 2007; Whitty et al., 2009; Morgan
et al,. 2010a). Largetooth sawfish have also been found to feed on
catfish, shrimp, croaker, small crustaceans, croaker, and mollusks
(Chidlow, 2007; Thorburn et al., 2007; Morgan et al., 2010a; Morgan et
al., 2010b). Largetooth sawfish captured off South Africa had bony fish
and shellfish as common diet items (Compagno et al., 1989; Compagno and
Last, 1999). In general, largetooth sawfish subsist on the most
abundant small schooling fishes in the area (NMFS, 2010a).
Population Structure
Genetic analyses based on specific sequences of mitochondrial DNA
indicated largetooth sawfish can be found in populations based on ocean
basin: Atlantic, Indo-West Pacific, and Eastern Pacific. There is also
restricted flow of genes in largetooth sawfish between these geographic
areas: Atlantic and Indo-West Pacific; Atlantic and eastern Pacific;
and Indo-West Pacific and eastern Pacific (Faria et al. 2013).
Genetic analyses based on a 480-base pair sequencing of the mtDNA
gene NADH-2 sequence also revealed information indicating largetooth
sawfish subpopulations. West and East Atlantic subpopulations differed
as did samples from Australia and the wider Indian Ocean. Collectively,
a total of 19 haplotypes were identified across largetooth sawfish: One
east Pacific haplotype, 12 western Atlantic haplotypes, two eastern
Atlantic haplotypes, one Indian Ocean haplotype, one Vietnamese-New
Guinean haplotype, and two Australian haplotypes (Faria et al., 2013).
This fine-scale structuring by haplotypes was only partially
corroborated by the regional variation in the number of rostral teeth.
While the rostral tooth count differed significantly in largetooth
sawfish collected from the western and eastern Atlantic Ocean, it did
not vary significantly between specimens collected from the Indian
Ocean and western Pacific (Faria et al., 2013). Largetooth sawfish
collected from the western Atlantic specimens had a higher rostral
teeth count than those collected from the eastern Atlantic. Data from
separate protein and genetics studies indicates some evidence of
distinction among populations of largetooth sawfish in the Indo-
Pacific. At a broad scale, Watabe (1991) found that there was limited
genetic variability between samples taken from Australia and Papua New
Guinea based on lactate dehydrogenase (LDH) isozyme patterns.
Largetooth sawfish might be genetically subdivided within the Gulf of
Carpentaria, Australia, with both eastern and western Gulf populations
(Lack et al., 2009).
Phillips et al. (2011) found that the population of largetooth
sawfish in the Gulf of Carpentaria is different from animals on the
west coast of Australia (Fitzroy River) based on mtDNA. Recent data
(Phillips, 2012) suggests that matrilineal structuring is found at
relatively small spatial scales within the Gulf of Carpentaria region
(i.e., this region contains more than one maternal `population'),
although the precise location and nature of population boundaries are
unknown. The difference in the genetic structuring using markers with
different modes of inheritance (maternal versus bi-parental) suggests
that largetooth sawfish may have male-biased dispersal and females
remaining at, or returning to, their birth place to mate (Phillips et
al., 2009; Phillips, 2012). Phillips (2012) noted that the presence of
male gene flow between populations in Australian waters suggests that a
decline of males in one location could affect the abundance and genetic
diversity of assemblages in other locations.
The genetic diversity for largetooth sawfish throughout Australia
seems to be low to moderate. Genetic diversity was greater in the Gulf
of Carpentaria than in Australian rivers, also suggesting potential
philopatry: Animals return to or stay in their home range (Lack et al.,
2009). Yet, given limited sampling, additional research is needed to
better understand potential population structure of largetooth sawfish
in Australia (Lack et al., 2009; Phillips et al., 2009; Morgan et al.,
2010a; Morgan et al., 2010b).
Distribution and Abundance
Largetooth sawfish have the largest historical range of all
sawfishes. The species historically occurred throughout the Indo-
Pacific near Southeast Asia and Australia and throughout the Indian
Ocean to east Africa. Older literature notes the presence of this
species in Zanzibar, Madagascar, India, and the southwest Pacific
(Fowler, 1941; Wallace, 1967; Taniuchi et al., 2003). Largetooth
sawfish have also been noted in the Eastern Pacific Ocean from Mexico
to Ecuador (Cook et al., 2005) or possibly Peru (Chirichigno and
Cornejo, 2001). In the Atlantic Ocean, largetooth sawfish inhabit warm
temperate to tropical marine waters from Brazil to the Gulf of Mexico
in the western Atlantic, and Namibia to Mauritania in the eastern
Atlantic (Burgess et al., 2009).
Given the recent taxonomic changes for largetooth sawfish, we
examined all current and historic records of P. microdon, P. perotteti,
and P. pristis for a comprehensive overview on distribution and
abundance. We conducted an extensive search of peer-reviewed
publications and technical reports, newspaper, records from the GBIF
Database, and magazine articles. The results of that search are
summarized below by major geographic region.
Indian Ocean
Largetooth sawfish historically occurred throughout the Indian
Ocean; however, current records are rare for many areas. The earliest
record of largetooth sawfish was in 1936 from Grand Lac near the Gulf
of Aden, Indian Ocean (Kottelat, 1985). A second record in 1936 is from
the Mangoky River, Madagascar (Taniuchi et al., 2003).
[[Page 73987]]
Records from the 1960s and 1970s are largely from India and South
Africa. One largetooth sawfish was reported from the confluence of the
Lundi and Sabi Rivers, South Africa in 1960, over 200 miles (mi) inland
(Jubb, 1967). Between 1964 and 1966, several largetooth sawfish were
caught in the Zambesi River, South Africa during a general survey of
rays and skates; largetooth sawfish have also been recorded in the
shark nets off Durban, South Africa (Wallace, 1967). In 1966, a male
(10 ft; 305 cm TL) was captured in a trawl net in the Gulf of Mannar,
Sri Lanka (Gunn et al., 2010). Largetooth sawfish were commonly caught
between 1973 and 1974 in the Bay of Bengal during the wet season (July
and September) but rarely during other times of the year. Largetooth
sawfish were also reported in three major rivers that empty into the
Bay of Bengal: The Pennaiyar, Paravanar, and Gadilam (Devadoss, 1978).
Current reports of largetooth sawfish throughout the Indian Ocean
are isolated and rare. Largetooth sawfish were recorded in South Africa
1992 and 1993 between Nelson Mandela Bay and Cape Town. Eight
additional observations are reported in South Africa but associated
date information was not included (GBIF database). While the species
could not be confirmed, a survey of fishing landing sites and
interviews with 99 fishers in Kenya by Nyingi found 71 reports of
sawfishes over the last 40 years (unpublished report from Dorothy Wanja
Nyingi to J. Carlson, NMFS, 2007). The longest time series of
largetooth sawfish catches is from the swimmer protection beach nets
off Natal, South Africa with a yearly average capture rate of 0.2
sawfish per 0.6 mi (1 km) net per year from 1981 to 1990; since then
only two specimens have been caught (CITES, 2007). Largetooth sawfish
were reported in Cochin, India by the Central Marine Fisheries Research
Institute in 1994, but no information about location, size, or number
of animals is available (Dan et al., 1994). Commercial landings of
elasmobranchs from 1981 to 2000 in the Bay of Bengal were mostly rays
with some largetooth sawfish (Raje and Joshi, 2003). In the Betsiboka
River, Madagascar, four largetooth sawfish were caught in 2001. The
most recent capture of a largetooth sawfish (18 ft; 550 cm TL) in India
occurred on January 18, 2011, between Karnataka and Goa
(www.mangalorean.com).
Indo-Pacific Ocean (Excluding Australia)
Many islands within the Indo-Pacific region contain suitable
habitat for largetooth sawfish, but few reports are available, perhaps
due to the lack of surveys or data reporting. The earliest records of
largetooth sawfish from the Indo-Pacific are from a compilation study
of elasmobranchs in the waters off Thailand that reports a largetooth
sawfish in the Chao Phraya River and its tributaries in 1945
(Vidthayanon, 2002). In 1955, two largetooth sawfish were captured from
Lake Sentani (present day Intan Jaya, Indonesia). Juvenile largetooth
sawfish have also been reported around the same time in a freshwater
river close to Genjem, Indonesia (Boeseman, 1956). In 1956, largetooth
sawfish were recorded in Lake Sentani (present day Intan Jaya,
Indonesia), (Boeseman, 1956; Thorson et al., 1966). In a study by Munro
(1967) in the Laloki River in the southeastern portion of New Guinea,
no sawfish were captured. From 1967 to 1977, five largetooth sawfish
were captured from the Indragiri River, Sumatra (Taniuchi, 2002). The
presence of largetooth sawfish in the Mahakam River, Borneo was
recorded in 1987 (Christensen, 1992). Three largetooth sawfish rostra
were acquired from local fish markets in Sabah in 1996 (Manjaji,
2002a). Additional surveys of local fish markets indicate largetooth
sawfish are still present in these areas, although locals have noticed
a decline in their abundance (Manjaji, 2002a). In 1996, two specimens
were found in Malaysia: One in Palau Nangka and one in Palau Besar
(GBIF Database).
Multiple records of largetooth sawfish have occurred in areas
throughout Papua New Guinea. From 1970 to 1971, Berra et al. (1975)
collected five largetooth sawfish from the Laloki River, Papua New
Guinea. Four largetooth sawfish were recorded in 1975 from the Fly
River system, Papua New Guinea and one in 1979 in the northern part of
Papua New Guinea near new Tangu (GBIF Database). In a survey of the Fly
River system, Papua New Guinea, 23 individuals were captured in 1978
(Roberts, 1978; Taniuchi and Shimizu, 1991; Taniuchi et al., 1991b;
Taniuchi, 2002). There are two reports of largetooth sawfish in the
1980s in Papua New Guinea: One in 1987 and one in 1988 (GBIF Database).
More recently, 36 largetooth sawfish were captured in September 1989 in
Papua New Guinea (Taniuchi and Shimizu, 1991; Taniuchi, 2002).
The scarcity of records from Indo-Pacific led to an increased
effort to document species presence. Anecdotal evidence suggests that
largetooth sawfishes have not been recorded in Indo-Pacific for more
than 25 years (White and Last, 2010). Largetooth sawfish have not been
recorded in the Mekong River, Laos for decades (Rainboth, 1996). In a
comprehensive study compiled by Compagno (2002a), no sawfishes were
found in the South China Sea between the years of 1923 and 1996. Data
from 200 survey days at fish landing sites in eastern Indonesia between
2001 and 2005 recorded over 40,000 elasmobranchs, but only 2 largetooth
sawfish (White and Dharmadi, 2007; Kyne and Feutry, 2013).
Australia
Australia may have a higher abundance of largetooth sawfish than
other areas within the species' current range (Thorburn and Morgan,
2005; Field et al., 2009). Despite their current abundance levels, we
only identified a few historic records from Australia. The first record
of a largetooth sawfish was in 1945 in the Northern Territory (Stevens
et al., 2005). There was a subsequent record in 1947, and two
largetooth sawfish from the Gulf of Carpentaria, Queensland were
reported in 1959 (GBIF Database). Faria et al. (2013) obtained a
rostrum that was collected in Australia in 1960.
Since the 1980s, we found significantly more records of largetooth
sawfish in Australia than other regions. A largetooth sawfish was
captured from the Keep River, Australia in 1981 (Compagno and Last,
1999). Three largetooth sawfish were recorded in 1984 near Marchinbar
Island, Northern Territory (GBIF Database). Blaber et al. (1990) found
that largetooth sawfish were among the top twenty-five most abundant
species in the trawl fisheries of Albatross Bay from 1986 to 1988.
Three largetooth sawfish were reported from the Gulf of Carpentaria,
Queensland: One in 1987 in Walker Creek, one in 1988 in the Gilbert
River, and one in 1991 in Marrakai Creek, a tributary of the Adelaide
River, Northern Territory (GBIF Database). Eight individuals were
captured in the Leichhardt River in 2008 (Morgan et al., 2010b). In a
preliminary survey of the McArthur River, Northern Territory, Gorham
(2006) reported two largetooth sawfish captured between 2002 and 2006.
Surveys (Peverell, 2005; Gill et al., 2006; Peverell, 2008) in the Gulf
of Carpentaria found largetooth sawfish widely distributed throughout
the eastern portion of the Gulf with most catches occurring near the
mouth of
[[Page 73988]]
many rivers (Mitchell, et al., 2005; 2008).
Juvenile largetooth sawfish in Australia use the Fitzroy River and
other tributaries of King Sound (Morgan et al., 2004) as nursery areas
while adults are found more often offshore (Morgan et al., 2010a). In
Western Australia, besides the Fitzroy River and King Sound, the only
other areas where juvenile sawfish have been recently recorded are in
Willie Creek and Roebuck Bay (Gill et al., 2006; Morgan et al., 2011).
Nursery areas for largetooth sawfish are also reported in northern
Australia in the Gulf of Carpentaria (Gorham, 2006). Juvenile
largetooth sawfish have been captured within the Adelaide River,
Australia in 2013 (P. Kyne, Charles Darwin University, pers. comm.,
2013). Abundance estimates for the largetooth sawfish from areas that
support higher human populations may be declining (Taniuchi and
Shimizu, 1991; Taniuchi et al., 1991a; Morgan et al., 2010a). Whitty et
al. (2009) found that the population of juvenile largetooth sawfish in
the Fitzroy River had declined; catch per unit effort was 56.7 sawfish
per 100 hours in 2003 compared to 12.4 in 2009. There were no reported
captures of largetooth sawfish in 2008 from the Roper River system,
which drains into the western Gulf of Carpentaria, Northern Territory
(Dally and Larson, 2008). No adult sawfish were captured in any of the
prawn trawl fisheries in Queensland, Australia during the month of
October 2001 (Courtney et al., 2006).
Outside the northern and western areas of Australia, largetooth
sawfish do occur but reports are less frequent. In southwestern
Australian waters, one female sawfish was captured by a commercial
shark fisherman in February 2003 east of Cape Naturaliste (Chidlow,
2007). Data from the Queensland, Australia Shark Control Program shows
a clear decline in sawfish catch over a 30 year period from the 1960s,
and the complete disappearance of sawfish in southern regions by 1993
(Stevens et al., 2005).
Eastern Pacific
In the eastern Pacific, the historic range of largetooth sawfish
was from Mazatlan, Mexico to Guayaquil, Ecuador (Cook et al., 2005) or
possibly Peru (Chirichigno and Cornejo, 2001). There is very little
information on the population status in this region and few reports of
capture records. The species has been reported in freshwater in the
Tuyra, Culebra, Tilapa, Chucunaque, Bayeno, and Rio Sambu Rivers, and
at the Balboa and Miraflores locks in the Panama Canal, Panama; in Rio
San Juan, Colombia; and in the Rio Goascoran, along the border of El
Salvador and Honduras (Fowler, 1936, 1941; Beebe and Tee-Van, 1941;
Bigelow and Schroeder, 1953; Thorson et al., 1966a; Dahl, 1971;
Thorson, 1974, 1976, 1982a, 1982b, 1987; Compagno and Cook, 1995; all
as cited in Cook et al., 2005). There are 4 records of largetooth
sawfish south of Purto Vallarta, Mexico in 1975, and several reports
from Panama with no associated dates (GBIF Database). The only recent
reports of largetooth sawfish in this area are anecdotal reports from
Colombia, Nicaragua, and Panama (R. Graham, Wildlife Conservation
Society, pers. comm. to IUCN, 2012).
Western Atlantic Ocean
In the western Atlantic Ocean, largetooth sawfish were widely
distributed throughout the marine and estuarine waters in tropical and
subtropical climates and historically found from Brazil through the
Caribbean, Central America, the Gulf of Mexico, and seasonally into
waters of the United States (Burgess et al., 2009). Largetooth sawfish
also occurred in freshwater habitats in Central and South America.
Throughout the Caribbean Sea, the historical presence of the largetooth
sawfish is uncertain and early records might have been misidentified
smalltooth sawfish (G. Burgess, Florida Museum of Natural History,
pers. comm. to IUCN, 2012).
Historic records of largetooth sawfish in the western north
Atlantic have been previously reported in NMFS (2010a). Sawfish were
documented in Central America in Nicaragua as early as 1529 by a
Spanish chronicler (Gill and Bransford, 1877). This species was also
historically reported in Nicaragua by Meek (1907), Regan (1908), Marden
(1944), Bigelow and Schroeder (1953) and Hagberg (1968). Five
largetooth sawfish were reported from a survey of Lake Izabal,
Guatemala from 1946 to 1947, and sawfishes were reported to be
important to inland fisheries (Saunders et al., 1950). There is a
single largetooth sawfish report from Honduras, but the true origin of
the rostrum and the date of capture could not be confirmed (NMFS,
2010a).
In Atlantic drainages, largetooth sawfish has been found in
freshwater at least 833 miles (1,340 km) from the ocean in the Amazon
River system (Manacapuru, Brazil), as well as in Lake Nicaragua and the
San Juan River; the Rio Coco, on the border of Nicaragua and Honduras;
Rio Patuca, Honduras; Lago de Izabal, Rio Motagua, and Rio Dulce,
Guatemala; and the Belize River, Belize. Largetooth sawfish are found
in Mexican streams that flow into the Gulf of Mexico; Las Lagunas Del
Tortuguero, Rio Parismina, Rio Pacuare, and Rio Matina, Costa Rica; and
the Rio San Juan and the Magdalena River, Colombia (Thorson, 1974,
1982b; Castro-Augiree, 1978 as cited in Thorson, 1982b; Compagno and
Cook, 1995; C. Scharpf and M. McDavitt, National Legal Research Group,
Inc., as cited in Cook et al., 2005).
In the United States, largetooth sawfish were reported in the Gulf
of Mexico mainly along the Texas coast east into Florida waters, though
nearly all records of largetooth sawfish encountered in U.S. waters
were limited to the Texas coast (NMFS, 2010a). Though reported in the
United States, it appears that largetooth sawfish were never abundant,
with approximately 39 confirmed records (33 in Texas) from 1910 through
1961.
The Amazon River basin and adjacent waters are traditionally the
most abundant known range of largetooth sawfish in Brazil (Bates, 1964;
Marlier, 1967; Furneau, 1969). Most of the records for which location
is known originated in the state of Amazonas, which encompasses the
middle section of the Amazon River basin along with the confluence of
the Rio Negro and Rio Solimoes Rivers. The other known locations are
from the states of Rio Grande do Norte, Sergipe, Bahia, Espirito Santo,
Rio de Janeiro, Sao Paulo, Para, and Maranhao (NMFS, 2010a). Most
records of largetooth sawfish in the Amazon River (Amazonia) predate
1974. The Magdalena River estuary was the primary source for largetooth
sawfish encounters in Colombia from the 1940's (Miles, 1945), while
other records originated from the Bahia de Cartagena and Isla de
Salamanca (both marine), and Rio Sinu (freshwater) from the 1960's
through the 1980's (Dahl, 1964; 1971; Frank and Rodriguez, 1976;
Alvarez and Blanco, 1985). In other areas of South America, there are
only single records from Guyana, French Guiana, and Trinidad from the
late 1800's and early 1900's. Of the 5 records from Suriname, the most
recent was 1962. Though thought to have once been abundant in some
areas of Venezuela (Cervignon, 1966a, 1966b), the most recent confirmed
records of largetooth sawfish from that country was in 1962.
Many records in the 1970's and 1980's are largely due to Thorson's
(1982a, 1982b) research on the Lake Nicaragua-Rio San Juan system in
Nicaragua and Costa Rica. Bussing (2002) indicated that this species
was known to inhabit the Rio Tempisque and tributaries of the San Juan
basin in Costa Rica. Following Thorson's (1982a, 1982b) studies,
[[Page 73989]]
records of largetooth sawfish in the western North Atlantic decline
considerably. By 1981, Thorson (1982a) was unable to locate a single
live specimen in the original areas he surveyed. There are no known
Nicaraguan records of the largetooth sawfish outside of the Lake
Nicaragua-Rio San Juan-Rio Colorado system (Burgess et al., 2009),
although largetooth sawfish are still captured incidentally by fishers
netting for other species (McDavitt, 2002). Of the known largetooth
sawfish reported from Mexico, most records are prior to 1978 (NMFS,
2010a). Caribbean records are very sparse (NMFS, 2010a). The last
record of a largetooth sawfish in U.S. waters was in 1961 (Burgess et
al., 2009).
Most recent records for largetooth sawfish are in isolated areas.
While many reports of largetooth sawfish from Brazil were from the
1980's and 1990's (Lessa, 1986; Martins-Juras et al., 1987; Stride and
Batista, 1992; Menni and Lessa, 1998; and Lessa et al., 1999), recent
records indicate largetooth sawfish are primarily found in fish markets
near the Amazon-Orinoco estuaries (Charvet-Almeida, 2002; Burgess et
al., 2009). A Lake Nicaragua fisherman reports he encounters a few
sawfish annually (McDavitt, 2002). Other records are rare for the area.
Three recent occurrences were found in Internet searches, one being a
200 lb. (90.7 kg) specimen caught recreationally in Costa Rica (Burgess
et al., 2009). Though reported by Thorson et al. (1966a, 1966b) to be
common throughout the area, there are no recent reports of encounters
with sawfishes in Guatemala. Scientists in Colombia have not reported
any sawfish sightings between 1999 and 2009 (Burgess et al., 2009).
Eastern Atlantic Ocean
Historic records indicate that largetooth sawfish were once
relatively common in the coastal estuaries along the west coast of
Africa. Verified records exist from Senegal (1841-1902), Gambia (1885-
1909), Guinea-Bissau (1912), Republic of Guinea (1965), Sierra Leone
(date unknown), Liberia (1927), C[ocirc]te d'Ivoire (1881-1923), Congo
(1951-1958), Democratic Republic of the Congo (1951-1959), and Angola
(1951). Most records, however, lacked species identification and
locality data and may have been confused taxonomically with other
species. Unpublished notes from a 1950's survey detail 12 largetooth
sawfish from Mauritania, Senegal, Guinea, C[ocirc]te d'Ivoire, and
Nigeria, ranging in size from 35-275 in (89-700 cm) TL (Burgess et al.,
2009).
A more recent status review by Ballouard et al. (2006) reported
that sawfishes, including the largetooth sawfish, were once common from
Mauritania to the Republic of Guinea, but are now rarely captured or
encountered. According to this report, the range of sawfishes has
decreased to the Bissagos Archipelago (Guinea Bissau). The most recent
sawfish encounters outside Guinea Bissau were in the 1990's in
Mauritania, Senegal, Gambia, and the Republic of Guinea. The most
recent documented largetooth sawfish capture was from 2005 in Nord de
Caravela (Guinea Bissau), along with anecdotal accounts from fishers of
captures off of two islands in the same area in 2008 (Burgess et al.,
2009).
In summary, on a global scale, largetooth sawfish appear to have
been severely fragmented throughout their historic range into isolated
populations of low abundance. Largetooth sawfish are now considered
very rare in many places where evidence is available, including parts
of East Africa, India, parts of the Indo-Pacific region, Central and
South America and West Africa. Even within areas like Australia and
Brazil, the species is primarily located in remote areas. Information
from genetic studies indicates that largetooth sawfish display strong
sex-biased dispersal patterns; with females exhibiting patterns of
natal philopatry while males move more broadly between populations
(Phillips et al., 2011). Thus, the opportunity for re-establishment of
these isolated populations is limited because any reduction in female
abundance in one region is not likely to be replenished by movement
from another region (Phillips, 2012).
Natural History of Green Sawfish (Pristis zijsron)
Taxonomy and Morphology
Pristis zijsron (Bleeker, 1851) is frequently known as the
narrowsnout sawfish or the green sawfish. Synonymous names include P.
dubius (Gloerfelt-Tarp and Kailola, 1984; Van Oijen et al., 2007;
Wueringer et al., 2009). An alternative spelling for this species'
scientific name (P. zysron) is found in older literature, due to either
inconsistent writing or errors in translation or transcription (Van
Oijen et al., 2007).
The green sawfish has a narrow saw with 25-32 small, slender
rostral teeth; tooth count may vary geographically (Marichamy, 1969;
Last and Stevens, 1994; Morgan et al., 2010a). Specimens collected
along the west coast of Australia have 24-30 left rostral teeth and 23-
30 right rostral teeth (Morgan et al., 2010a), although other reports
are 23-34 (Morgan et al., 2011). There have been no studies to
determine sexual dimorphism from rostral tooth counts for green
sawfish. The rostral teeth are generally denser near the base of the
saw than at the apical part of the saw (Blegvad and Loppenthin, 1944).
The total rostrum length is between 20.6-29.3 percent of the total
length of the animal and may vary based on the number and size of
individuals. In general, green sawfish have a greater rostrum length to
total length ratio than other sawfish species (Morgan et al., 2010a,
2011).
In terms of body morphology, the origin of the first dorsal fin on
green sawfish is slightly posterior to the origin of pelvic fins. The
lower caudal lobe is not well defined and there is no subterminal notch
(Gloerfelt-Tarp and Kailola, 1984; Compagno et al., 1989; Last and
Stevens, 1994; Compagno and Last, 1999; Bonfil and Abdallah, 2004;
Wueringer et al., 2009; Morgan et al., 2010a; Morgan et al., 2011). The
green sawfish has limited buccopharyngeal denticles and regularly
overlapping monocuspidate dermal denticles on its skin. As a result,
there are no keels or furrows formed on the skin (Deynat, 2005). The
green sawfish is greenish brown dorsally and white ventrally. This
species might be confused with the dwarf or smalltooth sawfish due to
its similar size and range (Compagno et al., 2006c).
Habitat Use and Migration
The green sawfish mostly uses inshore, marine habitats, but it has
been found in freshwater environments (Gloerfelt-Tarp and Kailola,
1984; Compagno et al., 1989; Compagno, 2002b; Stevens et al., 2008;
Wueringer et al., 2009). In the Gilbert and Walsh Rivers of Queensland,
Australia, specimens have been captured as far as 149 miles (240 km)
upriver (Grant, 1991). However, Morgan et al. (2010a, 2011) report
green sawfish do not move into freshwater for any portion of their
lifecycle. Like most sawfishes, the green sawfish prefers muddy bottoms
in estuarine environments (Last, 2002). The maximum depth recorded for
this species is 131 ft (40 m) but it is often found in much shallower
waters, around 16 ft (5 m; Compagno and Last, 1999; Wueringer et al.,
2009). Adults tend to spend more time in offshore waters in Australia,
as indicated by interactions with the offshore Pilbara Fish Trawl
Fishery, while juveniles prefer protected, inshore waters (Morgan et
al., 2010a; Morgan et al., 2011).
[[Page 73990]]
Age and Growth
At birth pups are between 2 ft and 2 ft 7 in (61 and 80 cm) TL. At
age 1 green sawfish are generally around 4 ft 3 in (130 cm) TL (Morgan
et al., 2010a). Peverell (2008) found between ages 1 and 5, green
sawfish measure between 4 ft 2 in and 8 ft 5 in (128 and 257 cm) TL,
based on the vertebral analysis of 6 individuals (Peverell, 2008;
Morgan et al., 2010a; Morgan et al., 2011). A 12 ft 6 in (380 cm) TL
green sawfish was found to be age 8, a 14 ft 4 in (438 cm) TL
individual was found to be age 10, a 14 ft 9 in (449 cm) TL specimen
was found to be age 16, and a 15 ft (482 cm) TL specimen was found to
be age 18 (Peverell, 2008; Morgan et al., 2011).
Adult green sawfish often reach 16 ft 5 in (5 m) TL, but may grow
as large as 23 ft (7 m) TL (Compagno et al., 1989; Grant, 1991; Last
and Stevens, 1994; Compagno and Last, 1999; Bonfil and Abdallah, 2004;
Compagno et al., 2006c; Morgan et al., 2010a). The largest green
sawfish collected in Australia was estimated to be 19 ft 8 in (600 cm)
TL based on a rostrum length of 5 ft 5 in (165.5 cm; Morgan et al.,
2010a; Morgan et al., 2011).
Peverell (2008) completed an age and growth study for green sawfish
using vertebral growth bands. Von Bertalanffy growth model parameters
from both sexes combined resulted in estimated maximum theoretical size
of 16 ft (482 cm) TL, relative growth rate of 0.12 per year and
theoretical time at zero length of 1.12 yrs. The theoretical maximum
age for this species is calculated to be 53 years (Peverell, 2008;
Morgan et al., 2010a).
Reproduction
Last and Stevens (2009) reported size at maturity for green sawfish
at 9 ft 10 in (300 cm) TL, corresponding to age 9. In contrast,
Peverell (2008) reported one mature individual of 12 ft 4 in (380 cm)
TL and estimated its age as 9 yrs. Using the growth function from
Peverell (2008) and assuming length of maturity at 118 in (300 cm),
Moreno Iturria (2012) determined maturation is likely to occur at age
5. Demographic models based on life history data from the Gulf of
Carpentaria indicate the generation time is 14.6 years, the intrinsic
rate of population increase is 0.02 per year, and population doubling
time is approximately 28 years (Moreno Iturria, 2012).
Green sawfish give birth to as many as 12 pups during the wet
season (January through July); Last and Stevens, 1994; Peverell, 2008;
Morgan et al., 2010a, 2011). In Western Australia, females are known to
pup in areas between One Arm Point and Whim Creek, with limited data
for all other areas (Morgan et al., 2010a; Morgan et al., 2011). The
Gulf of Carpentaria, Australia is also a known nursery area for green
sawfish (Gorham, 2006). It is not known where the green sawfish breed
or their length of gestation.
Diet and Feeding
Like other sawfish, green sawfish use their rostra to stun small,
schooling fishes, such as mullet, or use it to dig up benthic prey,
including mollusks and crustaceans (Breder Jr., 1952; Rainboth, 1996;
Raje and Joshi, 2003; Compagno et al., 2006c; Last and Stevens, 2009).
One specimen captured in 1967 in the Indian Ocean had jacks and razor
fish (Caranx and Centriscus) species in its stomach (Marichamy, 1969).
In Australia, the diet of this species often includes shrimp, croaker,
salmon, glassfish, grunter, and ponyfish (Morgan et al., 2010a).
Population Structure
Faria et al. (2013) found no global population structure for green
sawfish in their genetic studies. However, geographical variation was
found in the number of rostral teeth per side, suggesting some
population structure may occur. Green sawfish from the Indian Ocean
have a higher number of rostral teeth per side than those from western
Pacific specimens (Faria et al., 2013).
In Australia, genetic analysis found differences in green sawfish
between the west coast, the east coast, and the Gulf of Carpentaria
(Phillips et al., 2011). Genetic data suggests these populations are
structured matrilineally (from the mother to daughter) but there is no
information on male gene flow at this time. These results may be
indicative of philopatry where adult females return to or remain in the
same area they were born (Morgan et al., 2010a; Morgan et al., 2011;
Phillips et al., 2011). Phillips et al. (2011) also found low levels of
genetic diversity for green sawfish in the Gulf of Carpentaria,
suggesting the population may have undergone a genetic bottleneck.
Distribution and Abundance
The green sawfish historically ranged throughout the Indo-West
Pacific from South Africa northward along the east coast of Africa,
through the Red Sea, Persian Gulf, Southern Asia, Indo-Australian
archipelago, and east to Asia as far north as Taiwan and Southern China
(Fowler, 1941; Blegvad and L[oslash]ppenthin, 1944; Smith, 1945; Misra,
1969; Compagno et al., 2002a, 2002b; Last and Stevens, 2009). Historic
records indicating species presence are available from India, Southeast
Asia, Thailand, Malaysia, Indonesia, New South Wales, and Australia
(Cavanagh et al., 2003; Wueringer et al., 2009; Morgan et al., 2010a;
Morgan et al., 2011). Green sawfish have also been found in South
Africa, the South China Sea, and the Persian Gulf (Fowler, 1941;
Compagno et al., 1989; Grant, 1991; Compagno and Last, 1999; Last,
2002; Compagno, 2002b; Morgan et al., 2010a). To evaluate the current
distribution and abundance of the green sawfish, we conducted an
extensive search of peer-reviewed publications and technical reports,
newspaper, magazine articles, and the GBIF Database. The results are
summarized by geographic area.
Indian Ocean
Green sawfish are widely distributed throughout the Indian Ocean
with the first record coming from Saudi Arabia in 1830 (GBIF Database).
An additional record was reported from the Indian Ocean in the 1850s
(GBIF Database). Several green sawfish were described near the Indian
archipelago in the late 1800s (Van Oijen et al., 2007). Additional
historical records include one female specimen captured in the Red Sea
near Dollfus in 1929. In Egypt, two green sawfish rostra were found in
1938, and an additional rostrum was found on Henjam Island, Gulf of
Oman (Blegvad and Loppenthin, 1994).
Unconfirmed reports of green sawfish are available from the Andaman
and Nicobar Islands, India. In 1963, a male was captured at Port Blair,
Gulf of Andaman (James, 1973). A female was captured in 1967, in the
same area (Marichamy, 1969). One green sawfish was captured in the St.
Lucia estuary, South Africa during a survey between 1975 and 1976
(Whitfield, 1999). In 1984, a green sawfish was observed in Trafalgar,
South Africa (GBIF Database).
Despite historic records, there are few current records of green
sawfish in the Indian Ocean. There are some reports of green sawfish
from Iraq, Iran, South Africa, and Pakistan, but no dates are available
(GBIF Database). We presume green sawfish are extremely rare or
extirpated in the Indian Ocean based on the lack of current records.
Indo-Pacific Ocean (Excluding Australia)
The first description of the green sawfish was based on a rostral
saw (Bleeker, 1851) from Bandjarmasin, Borneo (Van Oijen et al., 2007).
A juvenile male was captured in Amboine, Indonesia in 1856 (Deynat,
2005). An isolated saw from the Gulf of Thailand
[[Page 73991]]
was obtained in 1895 and estimated to be from a green sawfish 4 ft 8 in
(143 cm) TL (Deynat, 2005). Eight specimens were sent to the Wistar
Institute of Anatomy in 1898 from Baram, British North Borneo (Fowler,
1941). One green sawfish was reported from East Sepik, Papua New Guinea
in 1929 (GBIF Database). In 1940, a green sawfish specimen was
collected from Zamboanga, Philippines (GBIF Database).
Many islands within the Indo-Pacific region contain suitable
habitat for sawfish, but few records are available, possibly due to the
lack of surveys or data reporting. Before 1995, there were few local
scientific studies on elasmobranchs, and only two species of freshwater
rays had been recorded in Borneo. As a result, a great effort to
document any unknown species was undertaken by Fowler (2002). Rostra
and records were documented in the study, including several dried
rostra of green sawfish from the Kinabatangan River area in the local
markets of Sabah, Borneo; no collection specifics were provided. Locals
also indicated that this species could often be found in the Labuk Bay
area (Manjaji, 2002a) and in the country's freshwater systems (Manjaji,
2002b); they also reported a decline of sawfish populations overall.
Elsewhere in the Indo-Pacific region, few records of green sawfish
have been reported. This species is currently considered endangered in
Thailand by Vidthayanon (2002) and Compagno (2002a); they also reported
no sawfish species from the South China Sea from 1923 to 1996.
Anecdotal evidence suggests that sawfishes have not been recorded in
Indonesia for more than 25 years (White and Last, 2010). Several
reports of green sawfish exist from Malaysia, Indonesia, and New
Zealand without any associated dates (GBIF Database).
Australia
In Australian waters, the earliest museum collection of the green
sawfish was in 1913 in Llyod Bay, Queensland, Australia (GBIF
Database). The Queensland Museum houses a green sawfish specimen
collected in 1929 that was found in Moreton Bay, Queensland (Fowler,
1941). Two records exist of green sawfish collected in 1936 from
Adeliade, South Australia (GBIF Database). We found very few records
for green sawfish during the middle part of the last century. In the
late 1970s and 1980s, reports of green sawfish began to occur again. In
1978, green sawfish were recorded in the Western Territory by CSIRO
(GBIF Database). There are multiple observations in 1980 of green
sawfish in Australia: two from the Northern Territory, and one from the
Gulf of Carpentaria (GBIF Database). A green sawfish was observed in
the Gulf of Carpentaria in 1981 by CSIRO. Two were observed in Western
Australia, one in 1982 and one in 1983 (GBIF Database). Two green
sawfish were captured from Balgal, Queensland, Australia in 1985
(Beveridge and Campbell, 2005). In the Gulf of Carpentaria, two green
sawfish were recorded in 1986, and one was recorded in 1987 (GBIF
Database).
One green sawfish was caught in the southern portion of the Gulf of
Carpentaria in late 1990 during a fish fauna survey (Blaber et al.,
1994). Alexander (1991) captured a female green sawfish from the west
coast of Australia that was used for a morphological study. Between
1994 and 2010, almost 50 tissue samples were taken from live green
sawfish or dried rostra from multiple areas around Australia, primarily
the Gulf of Carpentaria and northwest and northeast coasts (Phillips et
al., 2011). In 1997, one green sawfish was found at the mouth of
Buffalo Creek near Darwin, Northern Territory (Chisholm and
Whittington, 2000). In a survey from 1999 through 2001 by White and
Potter, (2004), one green sawfish was captured in Shark Bay,
Queensland. In 1999, one green sawfish was captured by CSIRO from the
Gulf of Carpentaria (GBIF Database). Peverell (2005, 2008) noted the
green sawfish was one of the least encountered species in a survey from
the Gulf of Carpentaria. In 2004, one green sawfish was reported near
Darwin, Northern Territory by the European Molecular Biology Lab (GBIF
Database). No green sawfish were captured from the Roper River system
in 2008, which drains into the western Gulf of Carpentaria, Northern
Territory (Dally and Larson, 2008). Some records have been reported for
the east coast of Australia; one female green sawfish was acoustically
tracked for 27 hours in May 2004 (Peverell and Pillans, 2004; Porteous,
2004). Peverell (2005, 2008) noted the green sawfish was one of the
least encountered species in a survey from the Gulf of Carpentaria.
In summary, limited data makes it difficult to determine the
current range and abundance of green sawfish. Nonetheless, given the
uniqueness (size and physical characteristics) of the sawfish, we
believe the lack of records in the areas where the species was
historically found indicates the species is no longer present or has
declined to extremely low levels. Extensive surveys at fish landing
sites throughout Indonesia since 2001 have failed to record the green
sawfish (White pers. comm. to IUCN, 2012). There is some evidence from
the Persian Gulf and Red Sea (e.g., Sudan) of small but extant
populations (A. Moore, RSK Environment Ltd., pers. comm. to IUCN,
2012). Green sawfish are currently found primarily along the northern
coast of Australia, but all sawfish species have undergone significant
declines in Australian waters. The southern extent of the range of
green sawfishes in Australia has contracted (Harry et al., 2011). Green
sawfish have been reported as far south as Sydney, New South Wales, but
are rarely found as far south as Townsville, Queensland (Porteous,
2004).
Natural History of the Non-Listed Population(s) of Smalltooth Sawfish
(Pristis pectinata)
This section includes information from the listed U.S. DPS of
smalltooth sawfish. The U.S. DPS of smalltooth sawfish was listed as
endangered on April 1, 2003 (68 FR 15674). The basis of the U.S. DPS
smalltooth sawfish listing was the significant differences in
management across international borders. We discuss information from
the U.S. DPS of smalltooth sawfish here because there is very little
basic biological information on smalltooth sawfish found outside the
U.S. We believe the information from the U.S. DPS is likely
representative of the non-U.S. population of smalltooth sawfish and is
useful for understanding its biology and extinction risk.
Taxonomy and Morphology
The smalltooth sawfish was first described as Pristis pectinatus,
Latham 1794. The name was changed to the currently valid P. pectinata
to match gender of the genus and species as required by the
International Code of Zoological Nomenclature.
The smalltooth sawfish has a thick body with a moderately sized
rostrum. As with many other sawfishes, tooth count varies by individual
or region. While there is no reported difference in rostral tooth count
between sexes, there have been reports of sexual dimorphism in tooth
shape, with males having broader teeth than females (Wueringer et al.,
2009). Rostral teeth are denser near the apex of the saw than the base.
Most studies report a rostral tooth count of 25 to 29 for smalltooth
sawfish (Wueringer et al., 2009). The saw may constitute up to one-
fourth of the total body length (McEachran and De Carvalho, 2002).
The pectoral fins are broad and long with the origin of the first
dorsal fin over or anterior to the origin of the
[[Page 73992]]
pelvic fins (Faria et al., 2013). The lower caudal lobe is not well
defined and lacks a ventral lobe (Wallace, 1967; Gloerfelt-Tarp and
Kailola, 1984; Last and Stevens, 1994; Compagno and Last, 1999; Bonfil
and Abdallah, 2004; Wueringer et al., 2009). This species has between
228 and 232 vertebrae (Wallace, 1967).
The smalltooth sawfish has buccopharyngeal denticles and regularly
overlapping monocuspidate (single-pointed) dermal denticles on their
skin. As a result, there are no keels or furrows formed on the skin
(Last and Stevens, 1994; Deynat, 2005). The body is an olive grey color
dorsally, with a white ventral surface (Compagno et al., 1989; Last and
Stevens, 1994; Compagno and Last, 1999). This species may be confused
with the narrow or green sawfish (Compagno, 2002b).
Habitat Use and Migration
All research on habitat use and migration has been conducted on the
U.S. DPS of smalltooth sawfish. A summary of recent information (NMFS,
2010b) indicates smalltooth sawfish are generally found in shallow
waters with varying salinity level that are associated with red
mangroves (Rhizophora mangle). Juvenile sawfish appear to have small
home ranges and limited movements. Simpfendorfer et al. (2011) reported
smalltooth sawfish have an affinity for salinities between 18 and at
least 24 ppt, suggesting movements are likely made, in part, to remain
within this salinity range. Therefore, freshwater flow may affect the
location of individuals within an estuary. Poulakis et al. (2011) found
juvenile smalltooth sawfish had an affinity for water less than 3 ft
(1.0 m) deep, water temperatures greater than 86 degrees Fahrenheit (30
degrees Celsius), dissolved oxygen greater than 6 mg per liter, and
salinity between 18 and 30 ppt. Greater catch rates for smalltooth
sawfish less than 1 year old were associated with shoreline habitats
with overhanging vegetation such as mangroves. Poulakis et al. (2012)
further determined daily activity space of smalltooth sawfish is less
than 1 mi (0.7 km) of river distance. Hollensead (2012) reported
smalltooth sawfish activity areas ranged in size from 837 square yards
to 240,000 square yards to approximately 3 million square yards (0.0007
to 2.59 km\2\) with average range of movements of 2.3 yards to 6.67
yards (2.4 to 6.1 m) per minute. Hollensead (2012) also found no
difference in activity area or range of movement between ebb and flood,
or high and low tide. Smalltooth sawfish movements at night suggest
possible nocturnal foraging. Using a combination of data from pop-off
archival transmitting tags across multiple institutional programs,
movements and habitat use of adult smalltooth sawfish were determined
in southern Florida and the Bahamas (Carlson et al., 2013). Smalltooth
sawfish generally remained in coastal waters at shallow depths less
than 32 ft; (10 m) for more than 96 percent of the time that they were
monitored. Smalltooth sawfish also remained in warm water temperatures
of 71.6 to 82.4 degrees Fahrenheit (22 to 28 degrees Celsius) within
the region where they were initially tagged. Tagged smalltooth sawfish
traveled an average of 49 mi (80.2 km) from deployment to pop-off
location during an average of 95 days. No smalltooth sawfish tagged in
U.S. or Bahamian waters have been tracked to countries outside where
they were tagged.
Age and Growth
There is no age and growth data for smalltooth sawfish outside of
the U.S. DPS. A summary of age and growth data on the U.S. DPS of
smalltooth sawfish (NMFS, 2010b) indicates rapid juvenile growth for
smalltooth sawfish for the first two years after birth. Recently,
Scharer et al. (2012) counted bands on sectioned vertebrae from
naturally deceased smalltooth sawfish and estimated von Bertalanffy
growth parameters. Theoretical maximum size was estimated at 14.7 ft
(4.48 m), relative growth was 0.219 per year, with theoretical maximum
size at 15.8 years.
Reproduction
In the eastern Atlantic Ocean, smalltooth sawfish have been
recorded breeding in Richard's Bay and St. Lucia, South Africa
(Wallace, 1967; Compagno et al., 1989; Compagno and Last, 1999).
Pupping grounds are usually inshore, in marine or fresh water. Pupping
occurs year-around in the tropics, but in only spring and summer at
higher latitudes (Compagno and Last, 1999). Records of captive breeding
have been reported from the Atlantis Paradise Island Resort Aquarium in
Nassau, Bahamas; copulatory behavior was observed in 2003 and six
months later the female aborted the pups for unknown reasons (McDavitt,
2006). In October 2012, a female sawfish gave birth to five live pups
at the Atlantis Paradise Island Resort Aquarium in Nassau, Bahamas (J.
Choromanski, Ripley's Entertainment pers. comm to NMFS, 2013).
Several studies have examined demography of smalltooth sawfish in
U.S. waters. Moreno Iturria (2012) calculated demographic parameters
for smalltooth sawfish in U.S. waters and estimated intrinsic rates of
increase at seven percent annually with a population doubling time of
9.7 years. However, preliminary results of a different model by Carlson
et al. (2012) indicates population increase rates may be greater, up to
17.6 percent annually, for the U.S. population of smalltooth sawfish.
It is not clear which of these models is more appropriate for the non-
U.S. population of smalltooth sawfish.
Diet and Feeding
Smalltooth sawfish often use their rostrum saw in a side-sweeping
motion to stun their prey, which may include small fishes, or to dig up
invertebrates from the bottom (Breder Jr., 1952; Compagno et al., 1989;
Rainboth, 1996; McEachran and De Carvalho, 2002; Raje and Joshi, 2003;
Last and Stevens, 2009; Wueringer et al., 2009).
Population Structure
A qualitative examination of genetic sequences revealed no
geographical structuring of smalltooth sawfish haplotypes; however,
variation in the number of rostral teeth per side was found in
specimens from the western and eastern Atlantic Ocean (Faria et al.,
2013).
Distribution and Abundance
Smalltooth sawfish were thought to be historically found in South
Africa, Madagascar, the Red Sea, Arabia, India, the Philippines, along
the coast of West Africa, portions of South America including Brazil,
Ecuador, the Caribbean Sea, the Mexican Gulf of Mexico, as well as
Bermuda (Bigelow and Scheroder, 1953; Wallace, 1967; Van der Elst 1981;
Compagno et al., 1989; Last and Stevens, 1994; IUCN, 1996; Compagno and
Last, 1999; McEachran and De Carvalho, 2002; Monte-Luna et al., 2009;
Wueringer et al., 2009). Yet, reports of smalltooth sawfish from other
than the Atlantic Ocean are likely misidentifications of other sawfish
(Faria et al., 2013). The lack of confirmed reports of smalltooth
sawfish from areas other than the Atlantic Ocean indicates that
smalltooth sawfish are only found in the Atlantic Ocean. In the eastern
Atlantic Ocean, smalltooth sawfish were historically found along the
west coast of Africa from Angola to Mauritania (Faria et al., 2013).
Although smalltooth sawfish were included in historic faunal lists of
species found in the Mediterranean Sea (Serena, 2005), it is still
unclear if smalltooth sawfish occurred as part of the Mediterranean
ichthyofauna or were only seasonal migrants.
To evaluate the current and historic distribution and abundance of
the
[[Page 73993]]
smalltooth sawfish outside the U.S. DPS, we conducted an extensive
search of peer-reviewed publications and technical reports, newspaper,
records from the GBIF Database, and magazine articles. The results of
that search are summarized by major geographic region.
Eastern Atlantic Ocean
Smalltooth sawfish were once common in waters off the west coast of
Africa, but are now rarely reported or documented in the area. The
earliest record of a smalltooth sawfish is a specimen from Namibia in
1874 (GBIF Database). Other records of smalltooth sawfish in Africa
occurred in 1907 from Cameroon, five males and two females. Female
specimens were recorded in the Republic of the Congo in 1911 and 1948.
Other reports from the Republic of Congo include a male and two
females, but dates were not recorded. An undated female specimen from
Mauritania was recorded (Faria et al., 2013). A rostrum from Pointe
Noire, Molez, Republic of the Congo was found in 1958 (Deynat, 2005;
Faria et al., 2013). There are records of smalltooth sawfish from
Senegal as early as 1956 and another rostral saw was recorded in 1959.
Faria et al. (2013) also reports on four other rostra from Senegal, but
no other information is available.
Many records of smalltooth sawfish from the eastern Atlantic Ocean
are reported in the GBIF database during the 1960s, particularly
between 1963 and 1964. The majority of these records are from Nigeria
(118), but others are from Gabon (77), Ghana (51), Cameroon (43), and
Liberia (39). Another online database, Fishbase (www.fishbase.org), has
the same records. It is unclear if these records are duplicative due to
the lack of specific information.
In the 1970s, records of smalltooth sawfish became limited to more
northern areas of West Africa. One rostral saw from Senegal was
recorded in 1975 (Alexander, 1991). Similarly, one rostral saw was
reported from Gambia in 1977, but information about exact location or
sex of the animal was absent (Faria et al., 2013). Faria et al. (2013)
report a record of smalltooth sawfish in Guinea-Bissau in 1983 and a
record of a saw in 1987. For a morphological study, Deynat (2005)
obtained a juvenile female from Cacheu, Guinea-Bissau in 1983, and
another from Port-Etienne, Mauritania, in 1986. Two rostra were
reported from the Republic of Guinea, one in 1980 and one in 1988
(Faria et al., 2013).
In the last 10 years, there has been only one confirmed record of a
smalltooth sawfish in the eastern Atlantic Ocean in Sierra Leone, West
Africa, in 2003 (M. Diop, pers. comm. to IUCN, 2012). Two other
countries have recently reported sawfish (Guinea Bissau, Africa in
2011, and Mauritania in 2010), but these reports did not identify the
species as smalltooth sawfish.
Western Atlantic Ocean (Outside U.S Waters)
Overall, records of smalltooth sawfish in the western Atlantic
Ocean are scarce and show a non-continuous range, potentially due to
misidentification with largetooth sawfish. Faria et al. (2013)
summarized most records of smalltooth sawfish in these areas. Faria et
al. (2013) report the earliest records are a female smalltooth sawfish
from Haiti in 1831 and a female sawfish from Trinidad and Tobago in
1876 (Faria et al., 2013). One smalltooth sawfish was recorded in
Bel[eacute]m, Brazil in 1863 (GBIF Database). Two smalltooth sawfish
saws were reported from Guyana in 1886, and an additional saw was later
recorded in 1900. In Brazil, there is a 1910 report of a female
smalltooth sawfish. In 1914, there is a report of a smalltooth sawfish
in Laguna de Terminos, Mexico (GBIF Database).
In the middle part of the twentieth century, there are reports of
two female smalltooth sawfish from Mexico in 1926. Rostral saws were
found in Suriname in 1943, 1944, and 1963, but no additional location
or specimen information is known. One rostrum was reported from Costa
Rica in 1960 and one rostral saw from Trinidad and Tobago in 1944
(Faria et al., 2013). Several whole individuals and one rostrum were
recorded from Guyana in 1958 and 1960. There are also several other
undated specimens recorded from Guyana from this period (Faria et al.,
2013). There are other records of smalltooth sawfish's presence in the
western Atlantic Ocean but specific information is lacking. For
example, Faria et al. (2013) report that 4 rostral saws came from
Mexico and two from Belize. One female was reported from Venezuela and
two rostra from Trinidad and Tobago. Despite lacking date information,
the GBIF Database and Fishbase have reports of smalltooth sawfish
throughout South and Central America: French Guiana (48), M[eacute]xico
(9), Guyana (6), Venezuela (3), Hait[iacute] (2), and individual
records from Colombia, Nicaragua, and Belize.
In summary, while records are sparse, it is likely the distribution
of smalltooth sawfish in the Atlantic Ocean is patchy and has been
reduced in a pattern similar to largetooth sawfish. Data suggests only
a few viable populations might exist outside the United States. The
Caribbean Sea may have greater numbers of smalltooth sawfish than other
areas given high quality habitats and reduced urbanization. For
example, smalltooth sawfish have been repeatedly reported along the
western coast of Andros Island, Bahamas (R.D. Grubbs, Florida State
University pers. comm. to J. Carlson, NMFS, 2014) and The Nature
Conservancy noted two smalltooth sawfish at the northern and southern
end of the island in 2006. Fishing guides commonly encounter smalltooth
sawfish around Andros Island while fishing for bonefish and tarpon
(R.D. Grubbs pers. comm. to J. Carlson, NMFS, 2014), and researchers
tagged two in 2010 (Carlson et al., 2013). In Bimini, Bahamas,
generally one smalltooth sawfish has been caught every two years as
part of shark surveys conducted by the Bimini Biological Station (D.
Chapman pers. comm.to Carlson, NMFS). In West Africa, Guinea Bissau
represents the last areas where sawfish can be found (M. Diop pers.
comm. to IUCN, 2012). Anecdotal reports indicate smalltooth sawfish may
also be found in localized areas off Honduras, Belize, and Cuba (R.
Graham, Wildlife Conservation Society, pers. comm. to IUCN, 2012).
Peer Review and Public Comments
In December 2004, the Office of Management and Budget (OMB) issued
a Final Information Quality Bulletin for Peer Review pursuant to the
Information Quality Act (IQA). The Bulletin was published in the
Federal Register on January 14, 2005 (70 FR 2664). The Bulletin
established minimum peer review standards, a transparent process for
public disclosure of peer review planning, and opportunities for public
participation with regard to certain types of information disseminated
by the Federal Government. The peer review requirements of the OMB
Bulletin apply to influential or highly influential scientific
information. The proposed rule and included status review were
considered influential scientific information under this policy and
subject to peer review. Similarly, a joint NMFS/FWS policy (59 FR
34270; July 1, 1994) requires us to solicit independent expert review
from at least three qualified specialists, concurrent with the public
comment period, on the science that is the basis for listing decisions.
To ensure this final rule was based on the best scientific and
commercial data available, we solicited peer review comments from three
scientists familiar with elasmobranchs.
On June 4, 2013, we published a proposed rule to list as endangered
five species of sawfish: Narrow sawfish (A.
[[Page 73994]]
cuspidata), dwarf sawfish (P. clavata), largetooth sawfish (P.
pristis), green sawfish (P. zijsron), and the non-U.S. DPS of
smalltooth sawfish (P. pectinata), that occurs outside U.S. waters, and
opened a 90-day public comment period (78 FR 33300). In the proposed
rule, we stated that we were not proposing to designate critical
habitat for any of the five species because they occur outside U.S.
waters. During our comment period we received a request to extend the
public comment period by 45 days. On August 7, 2013, we published a
notice extending the public comment period by 45 days (78 FR 48134). We
received a total of four public comments.
In the following sections of the document we summarize and respond
to the comments received from the public and peer reviewers on the
proposed rule.
Peer Review Comments
Comment 1: One commenter noted that the section of the proposed
rule addressing protective efforts did not include details on the
Sawfish Conservation Strategy developed by the IUCN Shark Specialist
Group. The commenter stated that the strategy is a protective effort
and will improve the conservation status of sawfishes worldwide. The
commenter predicted a medium to high certainty that the actions
identified in the Conservation Plan, when implemented, will be
effective.
Response: We have included the IUCN Sawfish Conservation Strategy
in the Protective Efforts section of this final rule. The Services
established two basic criteria in the PECE for evaluating conservation
efforts: (1) The certainty that the conservation efforts will be
implemented, and (2) the certainty that the efforts will be effective.
We evaluated the IUCN Sawfish Conservation Strategy and determined it
does not meet either criterion identified in the PECE. The strategy
identifies actions for countries to develop regulations or adopt
management actions to implement the strategy. However, the strategy
does not legally bind any country to enact laws or regulations, fund
conservation actions, or otherwise implement the strategy. We believe
there is considerable uncertainty that the actions identified in the
strategy will be adopted by the various countries within the range of
the five species of sawfish, and that resources are limited to support
these actions. Therefore, we cannot find that the strategy will
decrease extinction risk for any of the species.
Comment 2: One commenter stated that the Protective Efforts section
of the proposed rule did not include national protective efforts except
for the Convention on International Trade of Endangered Species of Wild
Fauna and Flora (CITES). The commenter stated that sawfish protections
in Australia were likely effective, but protections in India were
likely ineffective.
Response: We updated the Protective Efforts section of the rule and
included the new information on sawfish protections and conservation
efforts in Australia from the Australian Government's recently
published 2014 Draft Recovery Plan for Sawfish and River Sharks
(Department of Environment, 2014). We also included updated information
on existing laws in Australia and India designed to protect sawfishes
into the Inadequacy of Existing Regulatory Mechanisms section of this
final rule.
Comment 3: It was suggested we use information in Kyne et al.
(2013) to update the occurrence information for P. clavata.
Response: We appreciate the new information and updated the
occurrence information in the preceding sections. The information did
not impact our evaluation of the status of P. clavata.
Comment 4: We received a question about the origin of the 1996
record of dwarf sawfish from the Mekong River Basin, Laos.
Response: We cite Rainboth (1996) for this report from the early
1900s that assumed the dwarf sawfish was from the Mekong River Basin,
Laos. We acknowledge no specimen exists to confirm this report.
Comment 5: The validity of narrow sawfish reports from Tasmania by
Deynat (2005) was questioned in one comment given the cold, temperate
waters that do not support sawfish. The commenter suggested the record
of the sawfish specimen in the fish collection of CSIRO in Hobart,
Tasmania was erroneous.
Response: We reviewed the literature and agree with the commenter.
We removed the reference to reports of narrow sawfish in Tasmania.
Public Comments
Comment 1: One commenter requested we cite a more recent reference
for the information on the supply and demand of sawfish than the 1996
reference in the proposed rule. Specifically, the commenter questioned
the statement that ``sawfishes are in high demand throughout the world
for display'' and suggested that sawfishes are no longer in high demand
for display in aquaria.
Response: We updated our information on the aquaria trade of
sawfishes on current supply and demand of sawfishes in the Scientific
and Educational Uses section and removed the statement cited by the
commenter. Although we believe that sawfish are still in high demand in
the aquaria trade, we recognize that the recent inclusion of all
sawfishes under CITES Appendix I limits the use of sawfish for display
and requires acquisition of animals for aquaria from captivity or
captive breeding.
Comment 2: Several commenters stated that they were concerned about
the impacts of including ``injuring or killing a captive sawfish
through experimental or potentially injurious veterinary care or
conducting research or breeding activities on captive sawfish, outside
the bounds of normal animal husbandry practices'' in the list of
activities that could result in a violation of the ESA Section 9
prohibitions. The concerns relate to the impacts on captive propagation
and rearing programs being conducted by aquaria, and on the use of the
latest advanced technological techniques available for captive held
animals. The commenters requested clarification that fish care and
husbandry techniques could continue to be used by aquaria.
Response: As stated in the proposed rule, sawfish held in captivity
at the time of listing are afforded all of the ESA protections and may
not be killed or injured or otherwise harmed, and, therefore, must
receive proper care. We realize that the care of captive animals
necessarily entails handling or other manipulation and we do not
consider such activities to constitute injury or harm to the animals so
long as adequate care, including veterinary care, is provided. Such
veterinary care includes confining, tranquilizing, and anesthetizing
sawfishes when such practices, procedures, or provisions are necessary
and not likely to result in injury.
On the effective date of a final listing, ESA Section 9 take
prohibitions automatically apply for species listed as endangered and
any `take' of the species is illegal unless that take is authorized
under a permit or through an incidental take statement. Incidental take
statements result from ESA Section 7 consultations on the effects of
federal activities. ESA Section 10 permits can authorize directed take
(e.g., for scientific research or enhancement of the species) or
incidental take during an otherwise lawful activity that would not be
subject to ESA section 7 consultation. ESA Section 10 permits are
issued to entities or persons subject to the
[[Page 73995]]
jurisdiction of the United States. We encourage institutions with
captive sawfish who are considering activities outside the bounds of
normal animal husbandry (e.g., breeding or research) to contact NMFS
Office of Protected Resources, Permits and Conservation Division, to
determine if an ESA Section 10 permit is required to authorize the
proposed activity. We do not have information regarding emerging
advances in fish care and animal husbandry for sawfish held in
captivity so we cannot determine at this time if they are outside the
bounds of normal care for captive animals.
Comment 3: Several commenters requested clarification of the
meaning of the terms ``non-commercial'' and ``non-commercially'' as
those terms are used in the section titled Identification of those
Activities that Would Constitute a Violation of Section 9 of the ESA.
Response: Section 3 of the ESA defines the term ``commercial
activity'' to mean ``all activities of industry and trade, including
but not limited to, the buying and selling of commodities and
activities conducted for the purposes of facilitating such buying and
selling: Provided, however, That it does not include exhibitions of
commodities by museums or similar cultural or historical
organizations.'' NMFS will use the definition of ``commercial
activity'' to evaluate whether an activity is ``non-commercial'' or a
sawfish is being held ``non-commercially'' in captivity.
Our listing determinations and summary of the data on which it is
based, with the incorporated changes, are presented in the remainder of
this document.
Species Determinations
We first consider whether the narrow sawfish (A. cuspidata), dwarf
sawfish (P. clavata), largetooth sawfish (P. pristis), green sawfish
(P. zijsron), and of the non-U.S. DPS of smalltooth sawfish (P.
pectinata) meet the definition of ``species'' pursuant to section 3 of
the ESA. Then we consider if any populations meet the DPS criteria.
Consideration as a ``Species'' Under the Endangered Species Act
Based on the best available scientific and commercial information
described above in the natural history sections for each species, we
have determined that the narrow sawfish (A. cuspidata), dwarf sawfish
(P. clavata), largetooth sawfish (P. pristis), and green sawfish (P.
zijsron) are taxonomically-distinct species and therefore eligible for
listing under the ESA. The largetooth sawfish (P. pristis) now includes
the formerly recognized species P. microdon and the previously listed
P. perotteti. The decision to list P. pristis will replace our 2011
listing determination for P. perotteti.
Distinct Population Segments
In order to determine if the petitioned and currently non-listed
population segment of smalltooth sawfish (P. pectinata) constitutes a
``species'' eligible for listing under the ESA, we evaluated it under
our joint NMFS- USFWS Policy regarding the recognition of distinct
population segments (DPS) under the ESA (61 FR 4722; February 7, 1996).
We examined the three criteria that must be met for a DPS to be listed
under the ESA: (1) The discreteness of the population segment in
relation to the remainder of the species to which it belongs; (2) the
significance of the population segment to the remainder of the species
to which it belongs; and (3) the population segment's conservation
status in relation to the Act's standards for listing (i.e., Is the
population segment, when treated as if it were a species, endangered or
threatened?).
A population may be considered discrete, if it satisfies one of the
following conditions: (1) It is markedly separated from other
populations of the same taxon as a consequence of physical,
physiological, ecological, or behavioral factors; or (2) it is
delimited by international governmental boundaries within which
differences of control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the ESA.
We previously determined that smalltooth sawfish in the United
States merited protection as a DPS and listed the U.S. DPS of
smalltooth sawfish as endangered (68 FR 15674; April 1, 2003). At that
time, there was no information available to indicate smalltooth sawfish
in U.S. waters interact with those in international waters or other
countries, suggesting that the U.S. population may be effectively
isolated from other populations. However, there were few scientific
data on the biology of smalltooth sawfish, and it was not possible to
conclusively subdivide this species into discrete populations on the
basis of genetics, morphology, behavior, or other biological
characteristics. Because there were no identified mechanisms regulating
the exploitation of this species anywhere outside of the United States,
we considered that lack of protection as directly relevant to the
inadequacy of existing regulatory mechanisms and a basis for
considering the U.S. population as discrete across international
boundaries.
We now evaluate the non-U.S. population of smalltooth sawfish to
determine if it meets the discreteness criteria of the joint DPS
policy. First, we determine whether the non-U.S. population of
smalltooth sawfish is discrete from the U.S. population because it is
delimited by international governmental boundaries within which
differences of control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the ESA. Because we have
designated critical habitat for the U.S. DPS population of smalltooth
sawfish, there is a significant regulatory mechanism for protecting
smalltooth sawfish and their habitats in the United States that does
not exist for the non-U.S. population of smalltooth sawfish. Movement
data from smalltooth sawfish tagged in U.S. and Bahamian waters also
indicate no movement to countries outside where they were tagged. This
information provides support that the non-U.S. population is discrete
from the already-listed U.S. DPS on the basis of being markedly
separate as a consequence of ecological factors, in addition to our
previous determination that the U.S. DPS is discrete on the basis of
international boundaries and significant differences in regulatory
mechanisms. For smalltooth sawfish outside the U.S., we have no
information regarding genetic or other biological differences that
would provide a strong basis for further separating the non-U.S.
smalltooth sawfish population into smaller, discrete units. We,
therefore, conclude that the non-U.S. population of smalltooth sawfish
meets the discreteness criterion of the joint DPS policy and we
consider this population as a single potential DPS.
We next must consider whether the non-U.S. population of smalltooth
sawfish meets the significance criterion. The joint DPS policy gives
examples of potential considerations indicating the population's
significance to the larger taxon. Among these considerations is
evidence that the discrete population segment would result in a
significant gap in the range of the taxon. Smalltooth sawfish are
limited in their distribution outside of the United States to West
Africa, the Caribbean, Mexico, and Central and South America. Loss of
this group of smalltooth sawfish would result in a significant gap in
the range of this species and restrict distribution to U.S. waters.
Because the loss of smalltooth sawfish in areas outside the United
States would result in a
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significant gap in the range of the species, we conclude the non-U.S.
population of smalltooth sawfish is significant as defined by the DPS
policy.
Based on the above analysis of discreteness and significance, we
conclude that the non-U.S. population of smalltooth sawfish (P.
pectinata) meets the definition of a DPS and is eligible for listing
under the ESA, and hereafter refer to it as the non-U.S. DPS of
smalltooth sawfish.
Extinction Risk
Our updated extinction risk analysis provides a more detailed
discussion of the extinction risk analysis process that we used to
determine the risk of extinction for narrow sawfish, dwarf sawfish,
green sawfish, largetooth sawfish, and the non-U.S. DPS of smalltooth
sawfish to determine whether the species are threatened or endangered
per the ESA's definitions. We used an adaptation of the approach,
including the primary concepts, developed by Wainwright and Kope (1999)
to organize and summarize our findings. This approach was originally
developed for salmonids and has been adapted and applied in the review
of many other species (Pacific salmonid, Pacific hake, walleye pollock,
Pacific cod, Puget Sound rockfishes, Pacific herring, and black
abalone) to summarize the status of the species according to
demographic risk criteria. The approach is useful when there is
insufficient quantitative data to support development of population
viability models to investigate extinction risk and it allows the
incorporation of sparse and qualitative data. Wainwright and Kope
(1999) identified key demographic parameters that have a strong bearing
on extinction risk, with a focus on risks to small populations from
genetic effects and population dynamics. Using these concepts, adapted
to the biology of these sawfishes and our available data, we estimated
the extinction risk, based on demographic factors, for each of the five
species under both current threats and threats expected in the
foreseeable future. We also performed a threats assessment by
identifying the severity of threats that exist now and in the
foreseeable future.
We defined the ``foreseeable future'' as the timeframe over which
threats, or the species' response to those threats, can be reliably
predicted to impact the biological status of the species. We determined
that the foreseeable future is approximately three generation times,
calculated for each of the species based on the demographic
calculations of Moreno Iturria (2012): Narrow sawfish, 14 years; dwarf
sawfish, 49 years; largetooth sawfish, 48 years; green sawfish, 38
years; and the non-U.S. DPS of smalltooth sawfish, 30 years. After
considering the life history of each species, availability of data, and
type of threats, we concluded that three generations was an appropriate
measure to evaluate threats in the foreseeable future. As a late-
maturing species, with slow growth rate and low productivity, it would
take more than one generation for any conservation management action to
be realized and reflected in population abundance indices. The
timeframe of three generations is a widely used scientific indicator of
biological status, and has been applied to decision making models by
many other conservation management organizations, including the
American Fisheries Society, the CITES, and the IUCN.
We considered three demographic categories in which to summarize
available data and assess extinction risk of each sawfish species: (1)
Abundance, (2) population growth rate/productivity, and (3) genetic
integrity which include the connectivity and genetic diversity of the
species. We determined the extinction risk for each category, for both
now and in the foreseeable future, using a five level qualitative scale
to describe our assessment of the risk of extinction. At the lowest
level, a factor, either alone or in combination with other factors, is
considered ``unlikely'' to significantly contribute to risk of
extinction for a species. The next lowest level is considered to be a
``low'' risk to contribute to the extinction risk, but could contribute
in combination with other factors. The next level is considered a
``moderate'' risk of extinction for the species, but in combination
with other factors contributes significantly to the risk of extinction.
A ranking of ``high'' risk means that factor by itself is likely to
contribute significantly to the risk of extinction. Finally, a ranking
of ``very high'' risk means that factor is considered ``highly likely''
to contribute significantly to the risk of extinction.
We ranked abundance as high or very high risk which is likely to
contribute significantly to the current and foreseeable risk of
extinction for all five species. While it appears the northern coast of
Australia supports the largest remaining groups of dwarf, largetooth,
green, and narrow sawfish in the Pacific and Indian Ocean, data from
the Queensland, Australia Shark Control Program show a clear decline in
sawfish catch (non-species-specific) over a 30-year period from the
1960s. In addition, it shows the complete disappearance of sawfish in
southern regions (Stevens et al., 2005). The available data on
abundance of sawfishes indicates there are still some isolated groups
of sawfish in the western and central Indo-Pacific region, but their
abundance has likely declined from historic levels. Smalltooth sawfish
are still being reported outside of U.S. waters in the Caribbean Sea,
but records are few and mostly insular (e.g., Andros Island) where
habitat is available and gillnet fisheries are not a threat to the
species (see below). There are only four records of largetooth sawfish
in the eastern Atlantic Ocean over the last decade. In the western
Atlantic, recent largetooth sawfish records are from only the Amazon
River basin and the Rio Colorado-Rio San Juan area in Nicaragua.
Wainright and Kope (1999) stated short- and long-term trends in
abundance are a primary indicator of extinction risk. These trends may
be calculated from a variety of quantitative data such as research
surveys, commercial logbook or observer data, and landings information
when accompanied by effort, but there is an absence of long-term
monitoring data for all five sawfishes. We looked at the available data
closely to see if we could support inferences about extinction risk
based on the trends in past observations using the presence of a
particular species at specified places and times (e.g., Dulvy et al.,
2003; Rivadeneira et al., 2009). The available museum records, negative
scientific survey results, and anecdotal reports do indicate the
abundance trend for all five sawfishes is declining and population
sizes are small. Information available on the species' distribution
indicates the species' ranges have also contracted. In many areas where
sawfish still occur, they are subject to commercial and artisanal
fisheries and potential habitat loss. We therefore ranked the risk of
extinction posed by the sawfishes' abundances as high, now and into the
foreseeable future.
We next considered the species' potential growth rates and
productivity as measures of their ability to recover from depleted
levels and provide inherent protection against extinction risk. Sawfish
have historically been classified as having both low reproductive
productivity and low recovery potential. The demography of smalltooth
and largetooth sawfish from the northwest Atlantic Ocean that was
originally investigated using an age-structured life table
(Simpfendorfer, 2000). Using known estimates of growth, mortality, and
reproduction at the time, Simpfendorfer (2000) determined that
intrinsic rates of
[[Page 73997]]
population increase ranged from 8 to 13 percent per year, and
population doubling times were approximately 5 to 8.5 years for both
species. These estimates included assumptions that there was no fishing
mortality, no habitat limitations, no population fragmentation, or
other effects of small population sizes. Simpfendorfer (2006) further
modeled the demography of smalltooth sawfish using a method for
estimating the rebound potential of a population by assuming that
maximum sustainable yield was achieved when the total mortality was
twice that of natural mortality. This demographic model produced
intrinsic rates of population increase that were from two to seven
percent per year for both smalltooth and largetooth sawfish. These
values are similar to those calculated by Smith et al. (2008) using the
same methodology corresponding to elasmobranch species with the lowest
productivity. Musick et al. (2000) noted that species with intrinsic
rates of increase of less than 10 percent were particularly vulnerable
to rapid population declines and a higher risk of extinction.
Some recent studies on the life history of sawfish, however,
indicate they are potentially more productive than originally proposed.
Growth rates (von Bertalanffy ``K'') for some species, like narrow
sawfish, approach 0.34 per year (Peverell, 2008). Data from tag-
recapture studies and analysis of vertebral growth bands from
smalltooth sawfish indicate that the first few years after birth
represent the time when growth is most rapid (e.g., Simpfendorfer et
al., 2008; Scharer et al., 2012). Using updated life history
information, Moreno Iturria (2012) calculated intrinsic rates of
increase for these five species of sawfish and determined values
ranging from a low of 0.02 per year for green sawfish to a high of 0.27
per year for narrow sawfish with dwarf sawfish being second highest at
0.10 per year. Considering this information, and the inferred declining
trend in abundance, we conclude productivity is a moderate risk for the
narrow sawfish but a high risk for the other four species. We also
determined that productivity would remain a moderate risk for the
narrow sawfish and is a high risk for the other four species, in the
foreseeable future.
We also assessed the species' extinction risk, based on genetic
diversity, spatial structure and connectivity. Population structure and
levels of genetic diversity have recently been assessed for the green
sawfish, dwarf sawfish, and largetooth sawfish across northern
Australia using a portion of the mtDNA control region. Phillips et al.
(2011) found statistically significant genetic structure within species
and moderate genetic diversity among these species. These results
suggest that sawfish may be more vulnerable to local extirpation along
certain parts of their range, especially in areas where the population
has been fragmented and movement between these areas is limited.
However, these results do not necessarily suggest a higher risk of
extinction throughout the entire range of the species. Chapman et al.
(2011) investigated the genetic diversity of the U.S. DPS of smalltooth
sawfish that has declined to between one percent to five percent of its
abundance at the turn of the twentieth century, while its core
distribution has contracted to less than 10 percent of its former range
(NMFS, 2009). Surprisingly, given the magnitude of this population
decline and range contraction, the U.S DPS of smalltooth sawfish does
not exhibit any sign of genetic bottlenecks, and it has genetic
diversity that is similar to other, less depleted elasmobranch
populations (Chapman et al., 2011). Given that all five species of
sawfish considered here have suffered similar abundance declines, we
believe this conclusion should serve as a surrogate for the other
sawfish species. Because the U.S. DPS of smalltooth sawfish has not
undergone a genetic bottleneck, we ranked genetic diversity as a
moderate risk for all sawfish species as it is likely, in combination
with other factors, to contribute significantly to the risk of
extinction. However, we determined that the risk of extinction due to
the lack of connectivity was high for all five species, primarily
because all populations have undergone severe fragmentation. While
genetic results provide optimism for the remaining populations of
sawfish, this does not preclude the promotion of management actions to
enhance connectivity among populations that have been historically
fragmented. We are also somewhat optimistic that sawfish populations
may begin to rebuild in some areas and the risk of connectivity was
determined to decrease for smalltooth and the narrow sawfish in the
foreseeable future, although by only a small amount.
After reviewing the best available scientific data and assessing
the extinction risk on the five species of sawfishes based on their
status and demography, we conclude the risk of extinction for all five
species of sawfish is high.
Summary of Factors Affecting the Five Species of Sawfishes
Next we consider whether any of the five factors specified in
section 4(a)(1) of the ESA are contributing to the extinction risk of
these five sawfishes.
The Present or Threatened Destruction, Modification, or Curtailment of
Its Habitat or Range
We identified destruction, modification, or curtailment of habitat
or range as a potential threat to all five species of sawfishes and
determined this factor is currently, and in the foreseeable future,
contributing significantly to the risk of extinction of these species.
Coastal and Riverine Habitats
Loss of habitat is one of the factors determined to be associated
with the decline of smalltooth sawfish in the U.S. (NMFS, 2009). As
juveniles, sawfishes rely on shallow nearshore environments, primarily
mangrove-fringed estuaries as nurseries (e.g., Wiley and Simpfendorfer,
2010; Norton et al., 2012). Coastal development and urbanization have
caused these habitats to be reduced or removed from many areas
throughout the species' historic and current range. Habitat loss was
identified as one of the most serious threats to the persistence of all
species of sawfish, posing high risks for extinction. It is still
unclear how anthropogenic perturbations to habitats affect the
recruitment of juvenile sawfish, and therefore adequate protection of
remaining natural areas is essential. Given the threat from coastal
urbanization coupled with the predicted reduction of mangroves globally
(Alongi, 2008), we believe the risk of habitat loss would significantly
contribute to both the decline of sawfish and their reduced viability.
We expect habitat modification throughout the range of these
sawfishes to continue with human population increases. As humans
continue to develop rural areas, habitat for other species, like
sawfish, becomes compromised (Compagno, 2002b). Habitat modification
affects all five species of sawfish, especially those inshore, coastal
habitats near estuaries and marshes (Compagno and Last, 1999; Cavanagh
et al., 2003; Martin, 2005; Chin et al., 2010; NMFS, 2010). Mining and
mangrove deforestation severely alter the coast habitats of estuaries
and wetlands that support sawfish (Vidthayanon, 2002; Polhemus et al.,
2004; Martin, 2005). In addition, riverine systems throughout most of
these species' historical range have been
[[Page 73998]]
altered or dammed. For example, the potential expansion of the McArthur
River Mine would permanently realign channels that would in turn affect
the number of pools formed during the wet and dry seasons, many of
which are used as refuge areas for dwarf, green, or largetooth sawfish
(Polhemus et al., 2004; Gorham, 2006). In addition to the potential
expansion of the McArthur River Mine, the Nicaragua government is
proposing to build a cross-country canal through habitats currently
used by the remaining largetooth sawfish population in Lake Nicaraugua
(BBC News, Latin America and Caribbean, 2013).
Although the status of habitats across the global range of these
sawfishes is not well known, we expect the continued development and
human population growth to have negative effects on habitat, especially
to nearshore nursery habitats. For example, Ruiz-Luna et al. (2008)
acknowledge that deforestation of mangrove forests in Mexico has
occurred from logging practices, construction of harbors, tourism, and
aquaculture activities. Valiela et al. (2001) reported on mangrove
declines worldwide. They showed that the area of mangrove habitat in
Brazil decreased from 9652 to 5173 square miles (24,999 to 13,398
square kilometers) between 1983 and 1997, with similar trends in
Guinnea-Bissau 1837 to 959 square miles (4758 to 2484 square
kilometers) from 1953 to 1995. The areas with the most rapid mangrove
declines in the Americas included Venezuela, Mexico, Panama, the U.S.,
and Brazil. Along the western coast of Africa, the largest declines
have occurred in Senegal, Gambia, Sierra Leone, and Guinnea-Bissau.
World-wide mangrove habitat loss was estimated at 35 percent from 1980
to 2000 (Valiela et al., 2001). These areas where mangroves are known
to have decreased are within both the historic and current ranges of
these five species.
Hydroelectric and Flood Control Dams
Hydroelectric and flood control dams pose a major threat to
freshwater inflow into the euryhaline habitats of sawfishes.
Alterations of flow, physical barriers, and increased water temperature
affect water quality and quantity in the rivers, as well as adjacent
estuaries that are important nursery areas for sawfish. Regulating
water flow affects the environmental cues of monsoonal rains and
increased freshwater flow for pupping (Peverel, 2008; Morgan et al.,
2011). Changes in siltation due to regulated water flow may also affect
benthic habitat or prey abundance for these sawfishes (Compagno, 2002;
Polhemus et al., 2004; Martin, 2005; Thorburn et al., 2007; Chin et
al., 2010; Morgan et al., 2010a).
New dams being proposed to provide additional irrigation to
farmland upstream may affect sawfish habitat. For example, the Gilbert
River, in Queensland, Australia drains into the Gulf of Carpentaria,
which is the nursery area for green, dwarf, and largetooth sawfish.
Further modification of the McArthur and Gilbert Rivers, along with
increased commercial fishing in coastal waters, will negatively affect
sawfishes by reducing available habitat while increasing bycatch
mortality (Gorham, 2006).
Water Quality
Largetooth sawfish in particular, and likely the other sawfishes,
have experienced a loss of habitat throughout their range due to the
decline in water quality. Agriculture and logging practices increase
runoff, change salinity, and reduce the flow of water into freshwater
rivers and streams that affects the habitat of the largetooth sawfish
(Polhemus et al., 2004; IUCN Red List, 2006); mining seems to be the
most detrimental activity to water quality. Pollution from industrial
waste, urban and rural sewage, fertilizers and pesticides, and tourist
development all end up in these freshwater systems and eventually the
oceans. Pollution from these operations has caused a reduction in the
number of sawfish in these freshwater systems (Vidthayanon, 2002;
Polhemus et al., 2004).
In summary, habitat alterations that potentially affect sawfishes
include commercial and residential development; agricultural,
silvicultural, and mining land uses; construction of water control
structures; and modification to freshwater inflows. All sawfishes are
vulnerable to a host of habitat impacts because they use rivers,
estuaries, bays, and the ocean at various times of their life cycle.
Based on our review of current literature, scientific surveys and
anecdotal information on the historic and current distribution, we find
that destruction, modification, and curtailment of habitat or ranges
are a factor affecting the status of each species. We conclude that
this factor is contributing, on its own or in combination with other
factors, to the extinction risk of all five species of sawfishes.
Overutilization for Commercial, Recreational, Scientific, or
Educational Purposes
We identified overutilization for commercial, recreational,
scientific, or educational purposes as a potential threat to all five
species of sawfishes and determined that it is currently and in the
foreseeable future contributing significantly to their risk of
extinction.
Commercial Fisheries
Commercial fisheries pose the biggest threat to these sawfishes, as
these species are bycatch from many fisheries. Their unusual morphology
and prominent saw makes sawfishes particularly vulnerable to most types
of fishing gear, most notably any type of net (Anak, 2002; Hart, 2002;
Last, 2002; Pogonoski et al., 2002; Cavanagh et al., 2003; Porteous,
2004; Stevens et al., 2005; Gorham 2006; IUCN Red List, 2006; Chidlow,
2007; Field, 2009; Chin et al., 2010; NMFS, 2010; Morgan et al., 2011).
Trawling gear is of particular concern as it is the most common gear
used within the range and habitat of sawfishes (Compagno and Last,
1999; Taniuchi, 2002; Walden and Nou, 2008). In Thailand, all sawfish
fins obtained and sold to markets are a result of bycatch by otter-
board trawling and gillnet fisheries as there are no directed sawfish
fisheries in the country (Pauly, 1988; Vidthayanon, 2002). The Lake
Nicaragua commercial fishery for largetooth sawfish that collapsed
prior to the 1980's was comprised mostly of gillnet boats (Thorson,
1982a), and the commercial small coastal shark fishery in Brazil mainly
uses gillnets and some handlines (Charvet-Almeida, 2002). Subadult and
adult smalltooth sawfish have been reported as bycatch in the U.S. Gulf
of Mexico and south Atlantic shrimp trawl fishery (NMFS SEFSC, 2011);
however, if proper techniques are used, all sawfish species,
particularly adults, are fairly resilient and can be released alive
from most fishing gear (Lack et al., 2009).
Live release of sawfishes from commercial fishing gear does occur
but sawfishes are often retained. The meat is generally consumed
locally, but the fins and rostra are of high value and sold in markets
where these products are unregulated (CITES, 2007). In Brazil, a
captured sawfish is most likely retained because of the value of their
products, as the rostra, rostral teeth, and fins are valued at upwards
of $1,000 U.S. in foreign markets (NMFS, 2010a). The proportion of
largetooth sawfish in these markets is unknown, although as many as 180
largetooth sawfish saws were annually sold at a single market in
northern Brazil in the early 2000's (McDavitt and Charvet-Almeida,
2004). The Trade Records Analysis of Flora and Fauna in Commerce
(TRAFFIC) organization found that meat, liver oil, fins, and skin are
among the most
[[Page 73999]]
preferred sawfish products in Asian markets (Anak, 2002; Vidthayanon,
2002). In the Gulf of Thailand, over 5,291 US tons (4,800 tonnes) of
rays were caught annually from 1976 to 1989; at the same time over
1,102 US tons (1,000 tonnes) of rays were caught in the Andaman Sea
(Vidthayanon, 2002). It is likely that most of these products were sold
in Asian markets because of the high demand for sawfish products.
Reports of sawfish products in various markets throughout Asia are
often inconsistent and inaccurate despite international rules on trade
and possession of sawfish products (Fowler, 2002; Clarke et al., 2008;
Kiessling et al., 2009).
Recreational or commercial fishing gear may be abandoned or lost at
sea. These ``ghost nets'' are an entanglement hazard for sawfishes and
have become an increasing problem in the Gulf of Carpentaria where over
5,500 ghost nets were removed in 2009. Sawfish captures are expected to
occur in regions where no quantitative information about ghost nets
exists (Gunn et al, 2010).
Misidentification, general species-composition grouping, and
failure to record information are all concerns for reporting sawfish
captures in direct or indirect commercial fisheries (Stobutzki et al.,
2002b). With little enforcement of regional and international laws, the
practice of landing sawfishes may continue (NMFS, 2010a). All sawfish
populations have been declining worldwide, partly due to the negative
effects of commercial fishing (Stevens et al., 2000; Peverell, 2008).
Recreational Fisheries
Sawfish are bycatch of many recreational fisheries throughout their
range, even in areas where they are protected, including many
Australian rivers (Walden and Nou, 2008; Field et al., 2009). Peverell
(2008) reports that some sawfish are a target sport fish for
recreational fishermen in the Gulf of Carpentaria, Queensland.
Historical information from the U.S. indicates that recreational hook
and line fishers in Texas sometimes target large sharks as trophy fish
but may capture sawfish (Burgess et al., 2009). Elsewhere in the United
States, the abundance of sawfishes is low and likely never high enough
for recreational fishers to encounter sawfish, much less target it
(NMFS, 2010a). With the increase in human population along the coast,
recreational fishing has the potential to put additional pressure on
sawfish species that use coastal habitats (Walden and Nou, 2008).
Indigenous Take
Due to the large populations of various indigenous people
throughout the range of these five species, and the lack of data on the
animals they harvest, the number of sawfish taken by local peoples is
unknown. Elasmobranchs are caught for consumption throughout the Indo-
Pacific. In some areas, the meat and fins of these animals are of high
market value, and therefore they are sold rather than consumed locally.
Due to this unregulated consumption, removal of elasmobranchs, which
includes sawfishes, is a threat to their population(s) (Compagno and
Last, 1999; Pogonoski et al., 2002; Vidthayanon, 2002; Thorburn et al.,
2007; Peverell, 2008; Morgan et al., 2010a).
Some studies have been conducted on the use and value of
elasmobranch parts to various indigenous groups, particularly those in
eastern Sabah, Malaysia. One study (Almada-Villela, 2002) found the
majority of natives from Pulau Tetabuan and Pulau Mabul only take what
is necessary for subsistence. Sawfish rostra are also valued and kept
as decoration or given as gifts at the expense of the animal (Almada-
Villela, 2002; McDavitt et al., 1996; Vidthayanon, 2002).
Protective Coastal Nets
Protective gillnets to prevent shark attacks on humans is used in
some areas but can have a negative impact due to bycatch. Sawfishes are
highly susceptible to capture in nets because their saws are easily
tangled in nets. The Queensland Shark Control Program in Australia
places nets along beaches during the summer months. From 1970 to 1990,
sawfish bycatch in these nets declined despite relatively constant
effort; likely due to an overall decline in sawfish populations
(Stevens et al., 2005). In South Africa, the first protective gillnets
lined the southeast tip of the continent's coast as early as 1952. By
1990, over 27 mi (44 km) of nets lined the area between Richards Bay
and Mzamba (Dudley and Cliff, 1993). About 350 sharks and rays were
captured in these nets between 1981 and 1990. A high percentage of
entangled sawfish are released alive because of their ability to
breathe while motionless. Dudley and Cliff (1993) reported that 100
percent of largetooth sawfish and 67 percent of smalltooth sawfish
caught during that time were released alive. Still, subsequent
mortality post-release due to stress or injury from the process is
unknown and potentially detrimental given other fishing pressures
(Dudley and Cliff, 1993).
Scientific and Educational Uses
Sawfishes are unique animals that are currently on public display
in many large aquariums. Removal of sawfishes from their natural
habitats has caused some concern for these sawfish species and their
ecosystems. No information is available on the level of mortality that
occurs during the capture and transporting of live sawfish to aquaria.
Removal of female sawfish from the wild could have an effect on the
future reproductive capacity of that population (Anak, 2002; Harsan and
Petrescu-Mag, 2008). Limited information is available regarding the
number of sawfish that have been removed from the wild for display in
aquaria. All sawfish removed from Australian waters for aquaria
collections have been reported as juveniles (S. Olson, Association of
Zoos and Aquariums (AZA), 2013 pers. comm). The two most recent imports
of largetooth sawfish to an Association of Zoos and Aquariums (AZA)
accredited facility were in 2007 and 2008 (S. Olson, AZA, 2013 pers.
comm).
In July 2011, the Australian CITES Scientific Authority for Marine
Species reviewed their 2007 non-detriment finding for the export of P.
microdon and found that it was not possible to conclude with a
reasonable level of certainty that any harvest for export purposes
would not be detrimental to the survival or recovery of the species
(DSEWPaC, 2011). Since then, international trade in freshwater sawfish
from Australia has ceased.
Worldwide, we are not aware of any narrow sawfish in captivity
(Peverell, 2005, 2008). We are aware of 2 dwarf sawfish held in
captivity in Japan (McDavitt, 2006). Largetooth sawfish are the most
common sawfish species in captivity (NMFS, 2010a). Juvenile largetooth
measuring less than 3.5 ft (1 m) TL on average are most often caught
for the aquaria trade as they are easier to transport than adults
(Peter and Tan, 1997).
Globally, scientists are collecting information on sawfish biology.
Research efforts began in 2003 on the U.S. DPS population of smalltooth
sawfish and no negative impacts have been associated with this research
to date.
In summary, while no quantitative data on fishery impacts are
available, we conclude that given the susceptibility of sawfish to
entanglement in gillnets and trawl nets that are commonly used
throughout their range, sawfishes are likely captured as incidental
take. We are not aware of any fisheries
[[Page 74000]]
specifically targeting sawfishes. This impact from fisheries is the
most likely single cause of the observed range contractions and reduced
abundance in many areas of their former range. Trade of sawfish parts
occurs throughout the world. Sawfish have been exploited for their
fins, rostra, and teeth. Sawfish fins have been report in the shark fin
trade since the early 1900s (Mountnorris, 1809). Trade of sawfish parts
occurs on Internet sites such as eBay and Craigslist. Trade of sawfish
parts (e.g., fins, rostral teeth, and rostra) are also ongoing threats
to all five species (Harrison et al., 2014). Therefore, we conclude the
overutilization for commercial, recreational, scientific, or
educational purposes, alone or in combination with other factors as
discussed herein, is contributing significantly to the risk of
extinction of the narrow, dwarf, largetooth, green, and the non-U.S.
DPS of smalltooth sawfish.
Disease and Predation
We have determined that disease and predation are not potential
threats to any of the five species of sawfish and that it is unlikely
that these factors, on their own or in combination with other factors,
are contributing significantly to their risk of extinction of all five
sawfish species.
These species co-occur with other sawfishes and large sharks, but
we are not aware of any studies or information documenting
interspecific competition in terms of either habitat or prey (NMFS
2010a). Thorson (1971) speculated that the Lake Nicaragua bull shark
population may compete with largetooth sawfish, as both were prevalent,
but he offered no additional data. Sawfish have been documented within
the stomach of a dolphin (Tursiops truncatus) near Bermuda (Bigelow and
Schroeder, 1953; Monte-Luna et al., 2009), in the stomach of a bull
shark (C. leucas) in Australia (Thorburn et al., 2004), and evidence of
bite marks from what appeared to be a bull shark (C. leucas) on a
juvenile smalltooth sawfish in the United States have been reported (T.
Wiley-Lescher, Haven Worth Consulting, 2012 pers. comm). Crocodiles
also prey on sawfishes (Cook and Compagno, 2005). There is no evidence
that unusual levels of disease or predation affect any of the five
sawfish species. Based on the information available on disease and
predation for all five species of sawfish, we have determined that
disease and predation on their own, or in combination with other
factors, do not pose an extinction risk to any of these sawfishes.
Inadequacy of Existing Regulatory Mechanisms
We identified inadequacy of existing regulatory mechanisms as a
potential threat to each of the five species of sawfish. We determined
that this factor alone, or in combination with other factors, is
contributing significantly to their risk of extinction.
First, we reviewed general or global regulatory protections for
sawfish. The use of turtle exclusion devices (TEDs) in the nets of
trawl fisheries to conserve sea turtles occurs throughout much of the
range of sawfishes, but TEDs are not efficient in directing sawfish out
of nets because sawfish rostra get entangled (Stobutzki et al., 2002a;
Brewer et al., 2006) prior to reaching the TED. TEDs are often used
when trawling occurs along the sea bottom at depths of 49 ft to 131 ft
(15 to 40 m), areas where sawfish are likely to be found (Stobutzki et
al., 2002a). Most sawfishes show no difference in recovery after going
through a trawl net, regardless of the presence or absence of a TED
(Griffiths, 2006). Stobutzki et al. (2002a) found that large females
are more likely to survive capture after passing through a trawling net
and TED compared to smaller males. Only narrow sawfish were found to
benefit from the presence of TEDs in nets as 73.3 percent escaped
(Brewer et al., 2006; Griffiths, 2006). In general, TEDs tend to have
negligible impact on sawfish that get captured by trawling nets
(Stobutzki et al., 2002a; Griffiths, 2006), but they do provide an
escape route if the animal does not get entangled.
Data reporting agencies (i.e., customs and national fisheries) are
often inconsistent in their reporting of wildlife trade (Anak, 2002).
Reports are often vague and include general descriptions like ``shark
fin'' or ``ray,'' providing practically no information of trading rates
of specific products (Lack and Sant, 2011). Many countries in the Indo-
Pacific do not report bycatch statistics or elasmobranchs taken
illegally (Holmes et al., 2009). In order for effective management
plans to be implemented in fin markets and for sawfish product trade,
data need to be consistent.
Next, we reviewed regional or country specific regulatory
protections for sawfish. Many countries in the Indo-Pacific and the
Middle East do not have formal legislation for management or national
protection of the sawfish that may occur in their waters. Presently,
Thailand has regulated some fisheries, but has no protective
legislation for any elasmobranch in the country except for export of
marine species for aquaria (Vidthayanon, 2002). Among Middle Eastern
countries that fish for sharks, only Iran has implemented an
International Plan of Action for the Conservation and Management of
Sharks (IPOA Shark Plan). Nine Arab countries have recently signed a
Memorandum of Understanding on the Conservation of Migratory Sharks to
improve shark conservation measures under the United Nations
Environment Programme Convention on Migratory Species. Countries in
Africa face similar circumstances as enforcement for sawfish protection
is unknown (NMFS, 2010a). Countries that do have protective legislation
are often unable to effectively patrol their waters, and fishing
restrictions are routinely violated by foreign vessels (Lack. and Sant,
2008). In one study, genetic testing (DNA barcoding) was used to
identify fins from green sawfish confiscated from foreign boats
illegally fishing in northern Australian waters (Holmes, 2009).
The Australian government listed the largetooth, green, and dwarf
sawfishes as vulnerable on their Environmental Protection and
Biodiversity Conservation (EPBC) Act list. The EPBC Act protects these
sawfish and prohibits killing, injuring, taking, trading, keeping, or
moving an individual without a permit. Even with these protections in
place, the Draft Recovery Plan for Sawfish and River Sharks (Department
of the Environment, 2014) reports that these three sawfish species have
experienced substantial population declines.
In summary, several organizations are trying to regulate and manage
sawfish but often these regulations and management initiatives are
inadequate. Illegal exploitation by foreign fishers often occurs when
regulations exist but are not enforced (Kiessling et al., 2009).
Preventative measures on existing fishing mechanisms to avoid sawfish
catch, international monitoring of trade and bycatch, and governmental
influence on fisheries are not presently sufficient to protect
sawfishes. Specific regulation and monitoring of sawfishes by country
would provide better protection (Vidthayanon, 2002; Walden and Nou,
2008). Therefore, we conclude the inadequacy of existing regulatory
mechanisms has and continues to significantly contribute to the risk of
extinction of the narrow, dwarf, largetooth, green, and the non-U.S.
DPS of smalltooth sawfish.
Other Natural or Manmade Factors Affecting Its Continued Existence
In the proposed rule, we determined this was not a factor
contributing
[[Page 74001]]
significantly to the risk of extinction of all five species of sawfish.
We re-evaluated the information for this factor and changed our
conclusion from the proposed rule based on the fact that sawfish life
history traits, which consists of slow growth rates, late maturity,
long life spans, and low fecundity rates. These life history traits do
not enable them to respond rapidly to additional sources of mortality,
such as overexploitation and habitat degradation. Scientific
information available on all five species of sawfish indicates that
other natural or manmade factors are potential threats to all of the
five species of sawfish. We conclude it is likely that these factors,
on their own or in combination with other factors, are contributing
significantly to the risk of extinction for all five sawfish species.
An increase in global sea-surface temperature and sea level may
already be influencing sawfish populations (Clark, 2006; Walden and
Nou, 2008; Chin et al., 2010). Fish assemblages are likely to change
their distribution and could affect the prey base for sawfishes.
Estuaries, including sawfish pupping grounds, may be affected as
climate change changes patterns in freshwater flow due to rainfall and
droughts. Skewed salinities in these areas or extreme tide levels might
discourage adults from making up-river migrations (Clark, 2006).
Saltwater marsh grass and mangrove areas play important roles in
sawfish habitat as well (Simpfendorfer et al., 2010); any disruption to
these areas may affect sawfish populations. There is little agreement,
however, on the effects that climate change will have on sawfish and
their environments specifically (Clark, 2006; Chin et al., 2010).
Red tide is the common name for a harmful algal bloom (HAB) of
marine algae (Karenia brevis) that can make the ocean appear red or
brown. Karenia brevis is one of the first species ever reported to have
caused a HAB and is principally distributed throughout the Gulf of
Mexico, with occasional red tides in the mid- and south-Atlantic United
States. Karenia brevis naturally produces a brevetoxin that is absorbed
directly across the gill membranes of fish or through ingestion of
algal cells. While many HAB species are nontoxic to humans or small
mammals, they can have significant effects on aquatic organisms. Fish
mortalities associated with K. brevis events are very common and
widespread. The mortalities affect hundreds of species during various
stages of development. Red tide toxins can cause intoxication in fish,
which may include violent twisting and corkscrew swimming, defecation
and regurgitation, pectoral fin paralysis, caudal fin curvature, loss
of equilibrium, quiescence, vasodilation, and convulsions, culminating
in death. However, it is known that fish can die at lower cell
concentrations and can also apparently survive in much higher
concentrations. In some instances, mortality from red tide is not
acute, but may occur over a period of days or weeks after exposure to
subacute toxin concentrations. There is no specific information on red
tide effects on sawfish, but a single report exists of a smalltooth
sawfish that was found dead along the west coast of Florida, during a
red tide event (International Sawfish Encounter Database, 2009).
Therefore, we conclude that sawfishes occurring in the U.S. Gulf of
Mexico are vulnerable to red tide, but there is little information
documenting direct mortality resulting from exposure to red tide (NMFS,
2010a). Harmful algal blooms also exist in waters outside of the U.S.
Gulf of Mexico therefore, it is probable that all sawfishes are
vulnerable to harmful algal blooms wherever they occur. Collectively,
these other natural or manmade factors may be affecting the continued
existence of the narrow, dwarf, largetooth, green, and the non-U.S. DPS
of smalltooth sawfish. Based on the results from our extinction risk
analysis and information on other man-made factors affecting all five
species of sawfish, this factor is contributing to their extinction
risk.
Overall Risk Summary
After considering the extinction risks, both threat-based and
demographic, for each of the five species of sawfish, we have
determined the narrow, dwarf, largetooth, and green sawfish and the
non-U.S. DPS of smalltooth sawfish are in danger of extinction
throughout all of their ranges due to (1) present or threatened
destruction, modification or curtailment of habitat, (2)
overutilization for commercial, recreational, scientific, or
educational purposes, (3) inadequacy of existing regulatory mechanisms,
and (4) other natural or manmade factors affecting their continued
existence, and low abundance, lack of connectivity, and genetic
diversity.
Protective Efforts
Section 4(b)(1)(A) of the ESA requires the Secretary, when making a
listing determination for a species, to take into consideration those
efforts, if any, being made by any State or foreign nation to protect
the species. In judging the effectiveness of efforts not yet
implemented, or those existing protective efforts that are not yet
fully effective, we rely on the Services' joint ``Policy for Evaluation
of Conservation Efforts When Making Listing Decisions'' (``PECE''; 68
FR 15100; March 28, 2003). The PECE policy is designed to ensure
consistent and adequate evaluation on whether any conservation efforts
that have been recently adopted or implemented, but not yet proven to
be successful, will result in recovering the species to the point at
which listing is not warranted or contribute to forming the basis for
listing a species as threatened rather than endangered. The purpose of
the PECE policy is to ensure consistent and adequate evaluation of
future or recently implemented conservation efforts identified in
conservation agreements, conservation plans, management plans, and
similar documents when making listing determinations. The PECE provides
direction for the consideration of conservation efforts identified in
these documents that have not yet been implemented, or have been
implemented but not yet demonstrated effectiveness. The policy is
expected to facilitate the development of conservation efforts by
states and other entities that sufficiently improve a species' status
so as to make listing the species as threatened or endangered
unnecessary.
Two basic criteria were established in the PECE to use in
evaluating efforts identified in conservations plans, conservation
agreements, management plans or similar documents: (1) The certainty
that the conservation efforts will be implemented; and (2) the
certainty that the efforts will be effective. When we evaluate the
certainty of whether or not the formalized conservation effort will be
implemented, we may consider the following: Do we have a high level of
certainty that that the resources necessary to carry out the
conservation effort are available? Do the parties to the conservation
effort have the authority to carry it out? Are regulatory or procedural
mechanisms in place to carry out the efforts? If the conservation
effort relies on voluntary participation, we will evaluate whether the
incentives that are included in the conservation effort will ensure the
level of participation necessary to carry out the conservation effort.
In evaluating the certainty that a conservation effort will be
effective, we may consider the following: Does the effort describe the
nature and extent of the threats to the species to be addressed and how
these threats are reduced by the conservation effort? Does the effort
establish specific conservation objectives? Does the effort
[[Page 74002]]
identify the appropriate steps to reduce the threats to the species?
And does the effort include quantifiable performance measures to
monitor both compliance and effectiveness? Overall, we need to be
certain that the formalized conservation effort improves the status of
the species at the time we make a listing determination. The PECE
Policy also states that last-minute agreements (i.e., those that are
developed just before or after a species is proposed for listing) often
have little chance of affecting the outcome of a listing decision.
Last-minute efforts are also less likely to be able to demonstrate that
they will be implemented and effective in reducing or removing the
threats to a species. In addition, there are circumstances in which the
threats to a species are so imminent and/or complex that is will be
almost impossible to develop an agreement or plan that includes
conservation efforts that will result in making the listing
unnecessary. A conservation effort that satisfies the criteria for
implementation and effectiveness is considered when making a listing
determination, but may not ultimately change the risk assessment for
the species. Using the criteria identified in our PECE Policy we
evaluated conservation efforts to protect and recover the five sawfish
species that are either underway but not yet fully implemented, or are
only planned.
CITES restricts the trade of live animals to a vast array of
wildlife products derived from them, including food products, musical
instruments, tourist curios and medicines. Many wildlife species in
trade are not endangered, but the existence of an agreement to ensure
the sustainability of the trade is important in order to safeguard
these resources for the future. All sawfishes in the family Pristidae
were listed on Appendix I of CITES at the 14th Conference of the
Parties meeting in 2007. An Appendix I listing bans all commercial
trade in parts (e.g., rostral teeth, rostra, liver, and fins) or
derivatives of sawfish with trade in specimens of these species
permitted only in exceptional circumstances (e.g., for research
purposes). At that time, an annotation to the Appendix I listing
allowed the largetooth sawfish P. microdon (herein P. pristis) to be
treated as Appendix II ``for the exclusive purpose of allowing
international trade in live animals to appropriate and acceptable
aquaria for primarily conservation purposes.'' The annotation was
accepted on the basis that Australian populations of P. microdon were
robust relative to other populations in the species' range, and that
the capture of individuals for aquaria was not likely to be detrimental
to the population. Later, at the CITES 16th Annual Conference of the
Parties meeting in March of 2013, Australia proposed the transfer of P.
microdon from Appendix II to Appendix I, and the measure was adopted
and became effective on 12 June 2013. Therefore, live trade of P.
pristis (P. microdon) is currently banned and all commercial trade of
all sawfishes is banned per CITES Appendix I listing.
The recent banning of all trade of P. pristis (P. microdon) for
aquaria trade is a good conservation measure for the species and meets
all of the criteria for implementation and effectiveness. The recently
adopted CITES Appendix I listing for largetooth sawfish only bans the
live trade of the fish from Australia to approved foreign aquaria, all
other trade was banned with the 2007 listing. Only 11 largetooth
sawfish were approved for aquaria trade when the largetooth sawfish was
listed under CITES Appendix I with the annotation for aquaria trade.
The recent CITES Appendix I listing for largetooth sawfish is not
likely to significantly affect the species outside of the limited area
(Australia) where they were removed from the wild for aquaria display.
Given live trade of P. pristis (P. microdon) for aquaria use is not a
threat leading to the extinction risk of the species, we conclude the
full CITES Appendix I listing may satisfy the PECE policy's standards
for implementation and effectiveness, but the impact of this measure is
considered insignificant. Australia may be effective at enforcing trade
policies, but the recent Appendix I listing of P. microdon (largetooth
sawfish) alone, is not sufficient to protect the species throughout its
range.
The IUCN Shark Specialist Group, in collaboration with a large
number of the national and international stakeholders in sawfish
conservation, developed A Global Strategy for Sawfish Conservation
(Harrison and Dulvy, 2014). The strategy identifies the actions
required to achieve recovery for all sawfishes. The strategy outlines
seven objectives that are necessary to achieve recovery of all
sawfishes: Fisheries management, species protection, habitat
conservation, trade limitation, strategic research, education and
communication, and responsible husbandry. We evaluated the certainty of
whether or not the strategy would be implemented and determined that
(1) the strategy does not have a high level of certainty that the
resources necessary to carry out the conservation effort are available,
(2) that the strategy team members do not have the authority to carry
out all of the objectives, (3) regulatory or procedural mechanisms are
not in place to carry out the objectives, (4) and the conservation
efforts rely on voluntary participation that does not have incentives
that are included in the conservation effort that will ensure the level
of participation necessary to effectively carry out the conservation
effort. Based on the lack of certainty that the conservation efforts
will be implemented we determined the strategy does not satisfy the
PECE policy's standards for certainty of implementation and
effectiveness.
The Australian Government, Department of the Environment, published
a Draft Recovery Plan for Sawfish and River Sharks (Plan) in 2014
(Department of Environment, 2014). The Draft Plan covers three sawfish
species (P. pristis, P. zijsron, and P. clavata). The Plan identifies
specific actions and objectives necessary to stop local decline of
sawfish and river sharks and promotes their recovery. The goal of the
Draft Plan is to assist with the recovery of sawfish in Australian
waters in two ways: (1) Improving the population status leading to the
removal of the sawfish from the protected species list of EPBC; and (2)
ensuring anthropogenic actives do not hinder the recovery in the near
future, or impact the conservation status of the species in the future.
We evaluated the certainty of whether or not the Draft Plan would be
implemented. We determined that the strategy has a high level of
uncertainty regarding implementation because: (1) The Draft Plan does
not have dedicated funding so the resources necessary to carry out the
conservation efforts may not be available, and (2) the Draft Plan is
dependent on the participation of voluntary groups or organizations
(e.g., indigenous community groups and non-governmental organizations)
to carry out some of the actions. Based on the lack of certainty that
the Draft Plan will be implemented, we determined the Draft Plan does
not satisfy the PECE policy's standards for certainty of implementation
and effectiveness.
Listing Determinations
Section 4(b)(1) of the ESA requires that we make listing
determinations based solely on the best scientific and commercial data
available after conducting a review of the status of the species and
taking into account those efforts, if any, being made by any state or
foreign nation, or political subdivisions thereof, to protect and
conserve the species. We have reviewed the best available scientific
and commercial information including the petition, and the information
in the
[[Page 74003]]
review of the status of the five species of sawfishes, and we have
consulted with species experts.
We are responsible for determining whether narrow sawfish (A.
cuspidata), dwarf sawfish (P. clavata), largetooth sawfish (P.
pristis), green sawfish (P. zijsron), and the non-U.S. DPS of
smalltooth sawfish (P. pectinata) are threatened or endangered under
the ESA (16 U.S.C. 1531 et seq.). We have followed a stepwise approach
as outlined above in making this listing determination for these five
species of sawfish. We have determined that narrow sawfish (A.
cuspidata); dwarf sawfish (P. clavata); largetooth sawfish (P.
pristis); green sawfish (P. zijsron); and the non-U.S. DPS of
smalltooth sawfish (P. pectinata) constitute species as defined by the
ESA. We have conducted an extinction risk analysis and concluded that
the risk of extinction for all five species of sawfish is high, now and
in the foreseeable future. We have assessed the threats affecting the
status of each species using the five factors identified in section
4(a)(1) of the ESA and concluded the narrow, dwarf, largetooth, green,
and the non-U.S. DPS of smalltooth sawfish face ongoing threats from
habitat alteration, overutilization for commercial and recreational
purposes, inadequacy of existing regulatory mechanisms, and other
natural or manmade factors affecting their continued existence
throughout their ranges. Therefore, we find that all five species of
sawfishes are in danger of extinction throughout all of their ranges.
After considering efforts being made to protect these sawfishes, we
could not conclude the proposed conservation efforts would alter the
extinction risk for any of these five sawfishes.
Effects of Listing
Conservation measures provided for species listed as endangered or
threatened under the ESA include recovery actions (16 U.S.C. 1533(f));
Federal agency requirements to consult with NMFS and to ensure its
actions do not jeopardize the species or result in adverse modification
or destruction of critical habitat should it be designated (16 U.S.C.
1536); designation of critical habitat if prudent and determinable (16
U.S.C. 1533(a)(3)(A)); and prohibitions on taking (16 U.S.C. 1538). An
additional benefit of listing beyond these legal requirements is that
the recognition of the species' plight through listing promotes
conservation actions by Federal and state agencies, foreign entities,
private groups, and individuals.
Recovery Plans
NMFS may develop a recovery plan or plans for these species after
considering the conservation benefit to the species per ESA sections
4(f)(1) and 4(f)(1)(A). Section 4 (f)(1) of the ESA directs NMFS to
develop and implement recovery plans for the conservation and survival
of listed species, unless we find that such a plan will not promote the
conservation of the species. Section 4(f)(1)(A) further directs us, to
the maximum extent practicable, to give priority in developing plans to
those species that will most likely benefit from such plans.
Identifying Section 7 Consultation Requirements
Section 7(a)(2) (16 U.S.C. 1536(a)(2)) of the ESA and NMFS/USFWS
regulations require Federal agencies to consult with us to ensure that
activities authorized, funded, or carried out are not likely to
jeopardize the continued existence of listed species or destroy or
adversely modify critical habitat. The requirement to consult applies
to these Federal agency actions in the United States and on the high
seas. The five sawfishes all occur in the waters of foreign nations,
where there would be no consultation requirement. It is possible, but
highly unlikely, that the listing of the five species of sawfish under
the ESA may result in a minor increase in the number of Section 7
consultations for high seas activities.
Critical Habitat
Critical habitat is defined in Section 3 of the ESA (16 U.S.C.
1532(5)) as: (1) The specific areas within the geographical area
occupied by a species, at the time it is listed in accordance with the
ESA, on which are found those physical or biological features (a)
essential to the conservation of the species and (b) that may require
special management considerations or protection; and (2) specific areas
outside the geographical area occupied by a species at the time it is
listed upon a determination that such areas are essential for the
conservation of the species. Critical habitat shall not be designated
in foreign countries or other areas outside U.S. jurisdiction (50 CFR
424.12 (h)).
The best available scientific and commercial data show that the
geographical areas occupied by the narrow sawfish (A. cuspidata), dwarf
sawfish (P. clavata), green sawfish (P. zijsron), largetooth sawfish
(P. pristis), and the non-U.S. DPS of smalltooth sawfish (P. pectinata)
are entirely outside U.S. jurisdiction, so we cannot designate critical
habitat for these species in their occupied range.
We can designate critical habitat in unoccupied areas in U.S.
jurisdiction, if we determine the areas are essential for the
conservation of the species. Only the largetooth sawfish (P. pristis,
formerly P. perotteti) has a range that once included occasional use of
U.S. waters, with approximately 39 confirmed records (33 in Texas) from
1910 through 1961. All records of P. pristis in U.S. waters were
adults, mostly during the summer months. U.S. waters were a limited
part of the historic range, likely used for periodic, seasonal foraging
movements. There is no evidence of U.S. waters supporting any other
biological functions like breeding or nursery areas. Therefore, we
believe reestablishment back into U.S. waters is not required for the
recovery of P. pristis. Based on the best available information we have
not identified unoccupied areas in U.S. jurisdiction that are essential
to the conservation of any of the five sawfish species. Therefore, we
do not intend to designate critical habitat for the narrow, dwarf,
largetooth, green, or the non-U.S. DPS of smalltooth sawfish.
Identification of Those Activities That Would Constitute a Violation of
Section 9 of the ESA
On July 1, 1994, NMFS and FWS published a policy (59 FR 34272) that
requires us to identify, to the maximum extent practicable at the time
a species is listed, those activities that would or would not
constitute a violation of section 9 of the ESA. Because we are listing
all five sawfishes as endangered, all of the prohibitions of section
9(a)(1) of the ESA will apply to all five species. These include
prohibitions against the import, export, use in foreign commerce, and
``take'' of the species. Take is defined as ``to harass, harm, pursue,
hunt, shoot, wound, kill, trap, capture, or collect, or to attempt to
engage in any such conduct.'' These prohibitions apply to all persons
subject to the jurisdiction of the United States, including in the
United States or on the high seas. The intent of this policy is to
increase public awareness of the effects of this listing on proposed
and ongoing activities within the species' range. Activities that we
believe could result in a violation of Section 9 prohibitions of these
five sawfishes include, but are not limited to, the following:
(1) Take within the U.S. or its territorial sea, or upon the high
seas;
(2) Possessing, delivering, transporting, or shipping any sawfish
part that was illegally taken;
(3) Delivering, receiving, carrying, transporting, or shipping in
interstate or
[[Page 74004]]
foreign commerce any sawfish or sawfish part, in the course of a
commercial activity, even if the original taking of the sawfish was
legal;
(4) Selling or offering for sale in interstate commerce any sawfish
part, except antique articles at least 100 years old;
(5) Importing or exporting sawfish or any sawfish part to or from
any country;
(6) Releasing captive sawfish into the wild. Although sawfish held
non-commercially in captivity at the time of listing are exempt from
certain prohibitions, the individual animals are considered listed and
afforded most of the protections of the ESA, including most importantly
the prohibitions against injuring or killing. Release of a captive
animal has the potential to injure or kill the animal. Of an even
greater conservation concern, the release of a captive animal has the
potential to affect wild populations of sawfish through introduction of
diseases or inappropriate genetic mixing. Depending on the
circumstances of the case, NMFS may authorize the release of a captive
animal through a section 10(a)(1)(a) permit; and
(7) Engaging in experimental or potentially injurious veterinary
care or conducting research or breeding activities on captive sawfish,
outside the bounds of normal animal husbandry practices. Normal care of
captive animals necessarily entails handling or other manipulation of
the animals, and NMFS does not consider such activities to constitute
take or harassment of the animals so long as adequate care, including
adequate veterinary care is provided. Such veterinary care includes
confining, tranquilizing, or anesthetizing sawfishes when such
practices, procedures, or provisions are not likely to result in
injury. Captive breeding of sawfish is considered experimental and
potentially injurious. Furthermore, the production of sawfish progeny
has conservation implications (both positive and negative) for wild
populations. Experimental or potentially injurious veterinary
procedures and research or breeding activities of sawfish may,
depending on the circumstances, be authorized under an ESA 10(a)(1)(a)
permit for scientific research or the enhancement of the propagation or
survival of the species.
We have identified, to the extent known at this time, specific
activities that will not be considered likely to result in a violation
of Section 9. Although not binding, we consider the following actions,
depending on the circumstances, as not being prohibited by ESA Section
9:
(1) Take of a sawfish authorized by a 10(a)(1)(a) permit authorized
by, and carried out in accordance with the terms and conditions of an
ESA section 10(a)(1)(a) permit issued by NMFS for purposes of
scientific research or the enhancement of the propagation or survival
of the species;
(2) Incidental take of a sawfish resulting from Federally
authorized, funded, or conducted projects for which consultation under
section 7 of the ESA has been completed, and when the otherwise lawful
activity is conducted in accordance with any terms and conditions
granted by NMFS in an incidental take statement in a biological opinion
pursuant to section 7 of the ESA;
(3) Continued possession of sawfish parts that were in possession
at the time of listing. Such parts may be non-commercially exported or
imported; however the importer or exporter must be able to provide
sufficient evidence to show that the parts meet the criteria of ESA
section 9(b)(1) (i.e., held in a controlled environment at the time of
listing, non-commercial activity);
(4) Continued possession of live sawfish that were in captivity or
in a controlled environment (e.g., in aquaria) at the time of this
listing, so long as the prohibitions under ESA section 9(a)(1) are not
violated. Again, facilities should be able to provide evidence that the
sawfish were in captivity or in a controlled environment prior to
listing. We suggest such facilities submit information to us on the
sawfish in their possession (e.g., size, age, description of animals,
and the source and date of acquisition) to establish their claim of
possession (see For Further Information Contact);
(5) Provision of care for live sawfish that were in captivity at
the time of listing. These individuals are still protected under the
ESA and may not be killed or injured, or otherwise harmed, and,
therefore, must receive proper care. Normal care of captive animals
necessarily entails handling or other manipulation of the animals, and
we do not consider such activities to constitute take or harassment of
the animals so long as adequate care, including adequate veterinary
care is provided. Such veterinary care includes confining,
tranquilizing, or anesthetizing sawfish when such practices,
procedures, or provisions are not likely to result in injury; and
(6) Any importation or exportation of live sawfish or sawfish parts
with all accompanying CITES import and export permits and an ESA
section 10(a)(1)(a) permit for purposes of scientific research or the
enhancement of the propagation or survival of the species.
Section 11(f) of the ESA gives NMFS authority to promulgate
regulations that may be appropriate to enforce the ESA. Future
regulations may be promulgated to regulate trade or holding of sawfish,
if necessary. The public will be given the opportunity to comment on
future proposed regulations.
Policies on Peer Review
In December 2004, the Office of Management and Budget (OMB) issued
a Final Information Quality Bulletin for Peer Review establishing a
minimum peer review standard. Similarly, a joint NMFS/FWS policy (59 FR
34270; July 1, 1994) requires us to solicit independent expert review
from qualified specialists, concurrent with the public comment period.
The intent of the joint peer review policy is to ensure that listings
are based on the best scientific and commercial data available. We
formally solicited expert opinion of three appropriate and independent
specialists regarding the scientific and commercial data or assumptions
related to the information considered for listing.
We considered peer reviewer comments in making our determination.
We conclude that these experts' reviews satisfy the requirements for
``adequate [prior] peer review'' contained in the Information Quality
Bulletin for Peer Review and the joint NMFS/FWS policy (59 FR 34270;
July 1, 1994).
References
A complete list of the references used in this final rule is
available on the Internet at https://sero.nmfs.noaa.gov/protected_resources/sawfish/.
Classification
National Environmental Policy Act
The 1982 amendments to the ESA, in section 4(b)(1)(A), restrict the
information that may be considered when assessing species for listing.
Based on this limitation of criteria for a listing decision and the
opinion in Pacific Legal Foundation v. Andrus, 675 F. 2d 825 (6th Cir.
1981), NMFS has concluded that ESA listing actions are not subject to
the environmental assessment requirements of the National Environmental
Policy Act (NEPA) (See NOAA Administrative Order 216-6).
Executive Order 12866, Regulatory Flexibility Act, and Paperwork
Reduction Act
As noted in the Conference Report on the 1982 amendments to the
ESA, economic impacts cannot be considered when assessing the status of
a species. Therefore, the economic analysis
[[Page 74005]]
requirements of the Regulatory Flexibility Act are not applicable to
the listing process. In addition, this final rule is exempt from review
under Executive Order 12866. This final rule does not contain a
collection-of-information requirement for the purposes of the Paperwork
Reduction Act.
Executive Order 13132, Federalism
In accordance with E.O. 13132, we determined that this final rule
does not have significant Federalism effects and that a Federalism
assessment is not required.
List of Subjects in 50 CFR Part 224
Administrative practice and procedure, Endangered and threatened
species, Exports, Imports, Reporting and recordkeeping requirements,
and Transportation.
Dated: December 8, 2014.
Samuel D. Rauch, III,
Deputy Assistant Administrator for Regulatory Programs, National Marine
Fisheries Service.
For the reasons set out in the preamble, 50 CFR part 224 is amended
as follows:
PART 224--ENDANGERED MARINE AND ANADROMOUS SPECIES
0
1. The authority citation for part 224 continues to read as follows:
Authority: 16 U.S.C. 1531-1543 and 16 U.S.C 1361 et seq.
0
2. In Sec. 224.101, paragraph (h), amend the table by:
0
A. Removing the ``Sawfish, largetooth'' and the ``Sawfish, smalltooth
(United States DPS)'' entries.
0
B. Adding entries for five new sawfish species in alphabetic order by
Scientific name under ``Fishes'':
Sec. 224.101 Enumeration of endangered marine and anadromous species.
* * * * *
(h) The endangered species under the jurisdiction of the Secretary
of Commerce are:
----------------------------------------------------------------------------------------------------------------
Species\1\
-------------------------------------------------------------------- Citation(s) for Critical
Description of listing habitat ESA rules
Common name Scientific name listed entity determination(s)
----------------------------------------------------------------------------------------------------------------
* * * * * * *
Fishes
* * * * * * *
Sawfish, dwarf................ Pristis clavata.. Entire species.. [Insert Federal NA NA
Register
citation] 12/12/
2014.
Sawfish, green................ Pristis zijsron.. Entire species.. [Insert Federal NA NA
Register
citation] 12/12/
2014.
Sawfish, largetooth........... Pristis pristis Entire species.. [Insert Federal NA NA
(formerly Register
Pristis citation] 12/12/
perotteti, 2014.
Pristis pristis,
and Pristis
microdon).
Sawfish, narrow............... Anoxypristis Entire species.. [Insert Federal NA NA
cuspidata. Register
citation] 12/12/
2014.
Sawfish, smalltooth (Non-U.S. Pristis pectinata Smalltooth [Insert Federal NA NA
DPS). sawfish Register
originating citation] 12/12/
from non-U.S. 2014.
waters.
* * * * * * *
----------------------------------------------------------------------------------------------------------------
\1\ Species includes taxonomic species, subspecies, distinct population segments (DPSs) (for a policy statement,
see 61 FR 4722, February 7, 1996), and evolutionarily significant units (ESUs) (for a policy statement, see 56
FR 58612, November 20, 1991).
* * * * *
[FR Doc. 2014-29201 Filed 12-11-14; 8:45 am]
BILLING CODE 3510-22-P