# 78 FR 33300: Endangered and Threatened Wildlife and Plants; 12-Month Finding and Proposed Endangered Listing of Five Species of Sawfish Under the Endangered Species Act

> Federal · Regulations · In force

URL: https://www.frixlaw.com/law-library/statutes/FR_PRORULE_2013-13170

## Section

- **Citation:** 78 FR 33300
- **Heading:** Endangered and Threatened Wildlife and Plants; 12-Month Finding and Proposed Endangered Listing of Five Species of Sawfish Under the Endangered Species Act
- **Jurisdiction:** Federal
- **Kind:** Regulations
- **Status:** In force
- **Text as of:** August 14, 2026
- **Source:** Compiled text
- **Location:** Federal Register / Vol. 78 / 78 FR 33300

## Text

DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration 50 CFR Part 224 [Docket No. 101004485-3501-02] RIN 0648-XZ50 Endangered and Threatened Wildlife and Plants; 12-Month Finding and Proposed 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:
Proposed rule; 12-month petition finding; request for comments.

SUMMARY:
We, NMFS, have completed comprehensive status reviews under the Endangered Species Act (ESA) of five species of sawfishes in response to a petition to list six sawfish species. In our 90-day finding we determined that Pristis pristis, as described in the petition, was not a valid species and began our status review on the remaining five species ( Anoxypristis cuspidata; Pristis clavata; Pristis microdon; Pristis zijsron; and all non-listed population(s) of Pristis pectinata ). During our status review, new scientific information revealed that three previously recognized species ( P. microdon, P. pristis, and P. perotteti ) were in fact a single species, Pristis pristis. We had previously listed P. perotteti as an endangered species (July 12, 2011). We therefore also considered the information from our 2010 status review of P. perotteti, herein P. pristis. We have determined, based on the best scientific and commercial data available and after taking into account efforts being made to protect the species, that the narrow sawfish ( A. cuspidata ); dwarf sawfish ( P. clavata ); largetooth sawfish (collectively P. pristis; formerly P. pristis, P. microdon, and P. perotteti ); green sawfish ( P. zijsron ); and the non-listed population(s) of smalltooth sawfish P. pectinata meet the definition of an endangered species. We also include a change in the scientific name for largetooth sawfish in this proposed rule to codify the taxonomic reclassification of P. perotteti to P. pristis
ooth sawfish (collectively P. pristis; formerly P. pristis, P. microdon, and P. perotteti ); green sawfish ( P. zijsron ); and the non-listed population(s) of smalltooth sawfish P. pectinata meet the definition of an endangered species. We also include a change in the scientific name for largetooth sawfish in this proposed rule to codify the taxonomic reclassification of P. perotteti to P. pristis. We are not proposing to designate critical habitat because the geographical areas occupied by the species are entirely outside U.S. jurisdiction and we have not identified any unoccupied areas that are currently essential to the conservation of any of these species. We are soliciting information that may be relevant to these listing and critical habitat determinations, especially on the status and conservation of these species.

DATES:
Comments on this proposed rule must be received by August 5, 2013. Public hearing requests must be made by July 19, 2013.

ADDRESSES:
You may submit comments, identified by the following document number, NOAA-NMFS-2011-0073, by any of the following methods:
• Electronic Submissions: Submit all electronic public comments via the Federal eRulemaking Portal. Go to www.regulations.gov/#!docketDetail;D=NOAA-NMFS-2011-0073. click the “Comment Now” icon, complete the required fields, and enter or attach your comments.
• Fax: 727-824-5309; Attn: Assistant Regional Administrator for Protected Resources.
Instructions: You must submit comments by one of the above methods to ensure that we receive, document, and consider them. Comments sent by any other method, to any other address or individual, or received after the end of the comment period may not be considered. All comments received are a part of the public record and will generally be posted for public viewing on http://www.regulations.gov without change
s: You must submit comments by one of the above methods to ensure that we receive, document, and consider them. Comments sent by any other method, to any other address or individual, or received after the end of the comment period may not be considered. All comments received are a part of the public record and will generally be posted for public viewing on http://www.regulations.gov without change. All personal identifying information (e.g., name, address, etc.) confidential business information, or otherwise sensitive information submitted voluntarily by the sender will be publicly accessible. We will accept anonymous comments (enter “N/A” in the required fields if you wish to remain anonymous). Attachments to electronic comments will be accepted in Microsoft Word, Excel, or Adobe PDF file formats only.
You can obtain the petition, the proposed rule, and the list of references electronically on our NMFS Web site at http://sero.nmfs.noaa.gov/pr/pr.htm.
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 to 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 A. cuspidata, P. clavata, P. microdon, P. zijsron, and the non-listed population(s) of P. pectinata. Information in our records indicated that P
or threatened under the ESA; or alternatively to 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 A. cuspidata, P. clavata, P. microdon, P. zijsron, and the non-listed population(s) of P. pectinata. Information in our records indicated that P. p ristis 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. During the comment period we received five public comments.
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.” Section 3 of the ESA further 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,” on the other hand, 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)
tion. A “threatened species,” on the other hand, 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
In making listing determinations for these five species, we first determine whether each petitioned species meet the ESA definition of a “species”. Next, using the best available information gathered during the status reviews, we complete an extinction risk assessment using the general procedure of Wainwright and Kope (1999). We then assess the threats affecting the status of each species using the five factors identified in section 4(a)(1) of the ESA.
Once we have determined the threats, we assess efforts being made to protect the species to determine if these conservation efforts were adequate to mitigate the existing threats. We evaluate 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 their certainty of implementation and effectiveness for future or not yet fully implemented conservation efforts. Finally, we re-assess the extinction risk of each species in light of the existing conservation efforts.
Status Reviews
In order to conduct a comprehensive review, NMFS Southeast Region Protected Resources Division and NMFS Southeast Fisheries Science Center, Panama City Laboratory, staff members collaborated to identify the best available information. Unlike some previous 12-month findings from this agency, we have not developed a separate status review report
the existing conservation efforts.
Status Reviews
In order to conduct a comprehensive review, NMFS Southeast Region Protected Resources Division and NMFS Southeast Fisheries Science Center, Panama City Laboratory, staff members collaborated to identify the best available information. Unlike some previous 12-month findings from this agency, we have not developed a separate status review report. Instead, we present all information available for these species in this Federal Register notice; we first discuss background information relative to all five species and then 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) and Class Chondrichthyes (cartilaginous fish), also commonly known as elasmobranchs. 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)
e 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).
All 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, 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
ment 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 the 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). Per Compagno and Cook (1995) and Compagno (1999) these are A. cuspidata (Latham 1794), P. microdon Latham 1794, P. perotteti Muller & 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
these are A. cuspidata (Latham 1794), P. microdon Latham 1794, P. perotteti Muller & 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 these questions regarding the taxonomy of pristids, Faria et al. (2013) used historical taxonomy, external morphology, and mitochondrial DNA (mtDNA) sequences (NADH-2 loci) to hypothesize that the sawfishes comprise 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 at this time.
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, and 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)
o the head in adults, and 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 lateral canals in its rostrum that are not present in other sawfishes. These canals contain an additional connection to the ampullae of Lorenzini located on the underside of the rostrum (Wueringer et al., 2009).
Rostral tooth count varies for this species between 18-22 (Last and Stevens, 1994), 24-28 (Hussakof, 1912), and 27-32 (Miller, 1974). Total number of teeth has been found to vary by individual, region, and sex. Some studies report males having fewer rostral teeth than females, and others 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). Because these denticles first appear on neonate sawfish at 25.6-28 in (65-71 cm) total length (TL), they are developed post-natally
, 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). Because these denticles first appear on neonate sawfish at 25.6-28 in (65-71 cm) total length (TL), they are developed post-natally. 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 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
t 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). At an age of one year, saw length is 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)
sh 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).
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 sexually mature, mating may not occur until the rainy season in March-May (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 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, 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 subpopulations (Faria et al., 2013). 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) populations
Field et al., 2009) and, likely, 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 subpopulations (Faria et al., 2013). 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) populations. In a qualitative analysis when data were pooled, four haplotypes were identified: et al., 2013). A morphological distinction in narrow sawfish between the Indian Ocean and western Pacific Ocean subpopulations occurs in the number of rostral teeth (Faria et al., 2013). Specimens collected from the Indian Ocean had a higher number of rostral teeth per side than those collected from the western Pacific.
Field et al. (2009) examined the primary chemical components 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). However, we realize that using isotopes to separate elasmobranch populations is in its infancy and, coupled with the limited number of samples, it is not yet 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
d 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 pers. comm. to IUCN, 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. The result of that search is summarized below 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)
om 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 (Faria et al., 2013). Four juveniles were recorded in Pakistan waters in 1975; two females and two males.
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). 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 eleven more records off India, but no dates were recorded. From 1982-1994, 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 1981-2000, in the Bay of Bengal, total elasmobranch landings records are dominated by rays, but 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 pers. comm. to IUCN, 2012). The last published record of narrow sawfish from the western edge of the range, in the Straits of Hormuz, was in 1997 (A. Moore pers. comm. to IUCN, 2012)
landings records are dominated by rays, but 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 pers. comm. to IUCN, 2012). The last published record of narrow sawfish from the western edge of the range, in the Straits of Hormuz, was in 1997 (A. Moore 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. The first records of narrow sawfish were for juvenile males in 1852 and 1854 (Faria et al., 2013). In 1952, two females were captured from Batavia, Semarang, Indonesia along with a third female without a rostrum (Van Oijen et al., 2007). Both a female and male were recorded in 1867. Prior to 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 1930's. A female was reported in 1931 in Indonesia (no specific location), and a male 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).
Records of narrow sawfish throughout the Indo-Pacific continue to be scattered and infrequent throughout the 1950's. Faria et al. (2013) recorded rostra from Papua-New Guinea; two from 1955, 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 1970's, but there are no recent records (Taniuchi et al., 1991b)
rostra from Papua-New Guinea; two from 1955, 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 1970's, but there are no recent records (Taniuchi et al., 1991b). In a comprehensive literature search for the period 1923-1996 on the biodiversity of elasmobranchs in the south China Sea, Compagno (2002a) found no records of sawfishes. However, fresh dorsal and caudal fins of narrow sawfish were found during a survey of fish markets from 1996-1997 in Thailand (Manjaji, 2002b).
There are even fewer records of narrow sawfish from the Indo-Pacific over the last few decades. The only known specimen in the 21st century is a single report from New Guinea in 2001 (L. Harrison pers. comm.).
Australia
Australia may have larger populations of narrow sawfish than any other area within the species range (Peverell, 2005). The earliest record of narrow sawfish is from 1926 from Sydney (Pogonoski et al., 2002). We found no 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 3 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). A survey of fisheries data and records identified 74 offshore and 37 inshore records of narrow sawfish in the Gulf of Carpentaria (Peverell, 2005)
wfish 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). A survey of fisheries data and records 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 2007-2010 (Davies, 2010). Data from the Kimberley, Australia (R. McAuley pers. comm.to C. 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 northern area
ery 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 northern area. For example, a bycatch reduction device study conducted in 2001 reported narrow sawfish in the Gulf of Carpentaria, a similar study conducted off the eastern coast did not capture a single specimen (Courtney et al., 2006). 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 1969-2003 shows 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 small size and 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 the smalltooth sawfish or largetooth sawfish species complex (Last and Stevens, 1994; Cook et al., 2006; Morgan et al., 2010a) given the lack of distinct characters. Ishihara et al. (1991a) provides the most concise review of the physical characteristics of the dwarf sawfish.
The dwarf sawfish is olive brown in color dorsally with a white underside
n 1906; however it has often been confused with the smalltooth sawfish or largetooth sawfish species complex (Last and Stevens, 1994; Cook et al., 2006; Morgan et al., 2010a) given the lack of distinct characters. Ishihara et al. (1991a) provides the most concise review of the physical characteristics of the dwarf sawfish.
The dwarf sawfish is olive brown in color dorsally with a white underside. The rostrum of this species is quite short, with 19-23 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 (19-23 teeth) and females (20-23 teeth) (Ishihara et al., 1991a; Thorburn et al., 2008; Morgan et al., 2010a; Morgan et al., 2011). This species can be distinguished from largetooth sawfish based on tooth morphology as described by Thorburn et al. (2007). The rostrum makes up 21-26 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. 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)
he 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. 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 (2-3 m) deep that is often turbid and influenced heavily by tides. 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).
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 (Gorham, 2006). No adults or juveniles were found in freshwater areas of the river during the time of the study. However, physical characteristics such as salinity, temperature, and turbidity may limit the seasonal movements of the dwarf sawfish (Blaber et al., 1989).
Age and Growth
While small compared to other sawfishes, the maximum size of dwarf sawfish has been reported as: 4 ft 11 in (1.5 m) TL (Grant, 1991), 4 ft 7 in (140 cm) TL (Last and Stevens, 1994; Rainboth, 1996; Compagno and Last, 1999), 10 ft (306 cm) TL (Peverell, 2005), and 11.5 ft (350 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
arf sawfish has been reported as: 4 ft 11 in (1.5 m) TL (Grant, 1991), 4 ft 7 in (140 cm) TL (Last and Stevens, 1994; Rainboth, 1996; Compagno and Last, 1999), 10 ft (306 cm) TL (Peverell, 2005), and 11.5 ft (350 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 (24cm) 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 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 dwarf sawfish move into estuarine or fresh waters to breed during the wet season (Larson et al., 2006), however 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
een 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 population doubling time of 7.2 years.
Diet and Feeding
Dwarf sawfish, like other sawfishes, uses its 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, mullet, herring, and croaker (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 the west coast of 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; Morgan et al., 2011).
Later, Phillips et al
s 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; Morgan et al., 2011).
Later, Phillips et al. (2011), using additional samples 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). Genetic analysis of dwarf sawfish on the northern coast of Australia determined that they were distinct from those in other areas (Phillips et al., 2011). 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). 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). In an effort to gather more information on the historic and current range and abundance, we conducted an extensive search of peer-reviewed publications and technical reports, newspaper, and magazine articles. A summary of those findings is presented below by major geographic region
st 1999, Last and Stevens, 2009), there are questions regarding records outside of Australian waters (DSEWPaC, 2011). In an effort to gather more information on the historic and current range and abundance, we conducted an extensive search of peer-reviewed publications and technical reports, newspaper, and magazine articles. A summary of those findings is presented below 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 dwarf sawfish from the Indian Ocean: a female from the Reunion Islands, a female from an unidentified location in the Indian Ocean, and a male from India. 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, there has been two rostral saws in 1910, and 5 other rostra without date or length information.
Although reported historically, dwarf sawfish have not been reported from most other areas in the Indo-Pacific in over a decade. The most recent report of a dwarf sawfish in Thailand was in the Mekong River Basin, Laos in 1996. No sawfish species, including the dwarf sawfish, were reported from the South China Sea from 1923-1996 (Compagno, 2002a).
Pacific Ocean
Very few reports of the dwarf sawfish have been recorded in the western Pacific Ocean. Deynat (2005) reported on two skin samples from a juvenile female found in Tasmanian waters, and Faria et al. (2013) reported on two additional specimens but no specifics were provided.
Australia
Australia likely represents the center of the range of dwarf sawfish
23-1996 (Compagno, 2002a).
Pacific Ocean
Very few reports of the dwarf sawfish have been recorded in the western Pacific Ocean. Deynat (2005) reported on two skin samples from a juvenile female found in Tasmanian waters, and Faria et al. (2013) reported on two additional specimens but no specifics were provided.
Australia
Australia likely represents the center of the range of dwarf sawfish. Dwarf sawfish have been reported from Cairns to the east through the Gulf of Carpentaria in the north and through Kimberley to the west (Compagno and Last, 1999, Last and Stevens, 2009).
Most records for dwarf sawfish are from the north and northwest areas of Australia. The earliest record of this species is from 1877 (Faria et al., 2013). A single rostrum from a dwarf sawfish was found in 1916, but no other information was recorded. In 1946, a number of dwarf sawfish were reported (Faria et al., 2013).
Most records over the last 30 years have been from north and northwest Australia. Five female and five male dwarf sawfish (32-55 in; 82-140 cm TL) were captured in 1990 in the Pentecost River using gillnets (Taniuchi and Shimizu, 1991; Taniuchi, 2002). Between 1994 and 2010, almost 75 tissue samples were taken from live dwarf sawfish or dried rostra from the 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). 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 (eastern Australia) is preserved at the Harvard Museum of Comparative Zoology (Fowler, 1941)
rwin, Australia. One dwarf sawfish was captured in 1998 in the upper reaches of the Keep River estuary (Larson, 1999; Gunn et al., 2010). 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 (eastern Australia) is preserved at the Harvard Museum of Comparative Zoology (Fowler, 1941).
In a comprehensive survey of the Gulf of Carpentaria from 2001-2002, Peverell (2005; 2008) indicated dwarf sawfish were concentrated in the western portion of the Gulf of Carpentaria; twelve males and ten 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).
In northwestern Australia within specific riverine basins, 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, the intense fishing pressures within the range has likely caused the dwarf sawfish to become extirpated from much of the Indo-Pacific region and the species appears to be extirpated from eastern 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 territory and Pilbara region (western Australia) may be the last significant remaining areas for dwarf sawfish (P. Kyne pers. comm
a. 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 territory and Pilbara region (western Australia) may be the last significant remaining areas for dwarf sawfish (P. Kyne pers. comm. to IUCN, 2012).
Natural History of the Largetooth Sawfish ( Pristis pristis )
Taxonomy and Morphology
Many 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 (Günther, 1870; Garman, 1913; Fowler, 1936; Poll, 1951; Dingerkus, 1983; Daget, 1984; Séret and McEachran, 1986; McEachran and Fechhelm, 1998; Carvalho et al., 2007). The recent analysis by Faria et al. (2013) used mtDNA and contemporary genetic analysis to argue 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 1.7-2 times the width between the second anterior-most rostral teeth). Rostral teeth number is 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
en the two posterior-most rostral teeth is 1.7-2 times the width between the second anterior-most rostral teeth). Rostral teeth number is 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 64-68 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. Rostrum length can vary between males and females (Wueringer et al., 2009)
etooth 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. Rostrum length can vary between males and females (Wueringer et al., 2009).
Habitat Use and Migration
Largetooth sawfish are commonly found in coastal, 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 deep water 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 common in deeper holes, occupying muddy or sandy bottoms (NMFS, 2010a).
Adults more often utilize 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)
often utilize 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). Within the Gulf of Mexico, America, mature largetooth sawfish have historically moved as far north as Texas (NMFS, 2010a).
The physical characteristics of habitat strongly influence the movements and areas utilized 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 utilize extremely shallow areas (0-1 ft 7 in or 0-0.49 m) up to 80 percent of the time, mostly to avoid predators et al., 2007). Juveniles and adult largetooth sawfish also utilize rivers (Compagno, 2002b; Gorham, 2006) and can be found in areas up to 248.5 miles (400 km) upstream (Chidlow, 2007). Activity space of 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. At birth, largetooth sawfish are between 2 ft 6 in and 3 ft (76 and 91 cm) TL, 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 35-40 cm per year (NMFS, 2010a). 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)
h are between 2 ft 6 in and 3 ft (76 and 91 cm) TL, 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 35-40 cm per year (NMFS, 2010a). 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 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). Thorson (1982) estimated age of maturity to be 10 years at 9 ft 10 in (300 cm) TL in Lake Nicaragua (NMFS, 2010a). 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; Last and Stevens, 1994; Rainboth, 1996; Peter and Tan, 1997; Compagno and Last, 1999; Thorburn and Morgan, 2005; Compagno et al., 2006b; Chidlow, 2007; NMFS, 2010a). The largest largetooth sawfish recorded in Kimberley, Queensland measured 21 ft 6 in (656 cm) TL (Morgan et al., 2011). In other areas of Australia, the 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).
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 was calculated at 11 ft 11 in (363 cm) TL with a relative growth rate of 0.066 per year. Based on these calculations, it was determined that largetooth sawfish grow around 7 in (18 cm) in the first year and 4 in (10 cm) by the tenth year
1) who generated a von Bertalanffy growth model for specimens collected from Papua New Guinea and Australia. For both sexes combined, the theoretical maximum size was calculated at 11 ft 11 in (363 cm) TL with a relative growth rate of 0.066 per year. Based on these calculations, it was determined that largetooth sawfish grow around 7 in (18 cm) in the first year and 4 in (10 cm) by the tenth year. Thorson (1982a) estimated an early juvenile growth rate of 13-15 in (35-40 cm) per year and annual adult growth rate of 1 in (4.4 cm) per year based on largetooth from Lake Nicaragua. Peverell (2008) calculated a theoretical maximum size of 20 ft 11 in (638 cm) TL with a relative growth rate of 0.08 per year. The theoretical maximum age estimated for this species has been calculated to be 80 years (Morgan et al., 2010a).
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) from May to July (Raje and Joshi, 2003). The number of pups in a largetooth sawfish litter varies by location, and possibly due to other factors. One of the earliest reproductive studies on largetooth sawfish by Thorson (1976a) indicated litter size ranged between 1 to 13 pups, with an average of 7 pups per cycle (NMFS, 2010a). Thorson (1976a) also found that both ovaries appeared to be functional, though the left seemed to be larger and carry more ova (NMFS, 2010a). Length of gestation for largetooth sawfish is approximately five months, with a biennial reproductive cycle (NMFS, 2010a). Chidlow (2007) reported largetooth sawfish had litters with up to 12 pups.
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 population doubling time of 10.3 to 13.6 years
nial reproductive cycle (NMFS, 2010a). Chidlow (2007) reported largetooth sawfish had litters with up to 12 pups.
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 population doubling time of 10.3 to 13.6 years. Using data from Australia, rates of population increase were estimated to be around 0.12 per year (Moreno Iturria, 2012), with a population doubling time of approximately 5.8 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 (Moreno Iturria, 2012).
Diet and Feeding
Largetooth sawfish diet is predominately fish, but varies depending on study and 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 juvenile sawfish 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, 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)
ed on catfish, shrimp, 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 a 480 base pair sequencing of the mtDNA gene NADH-2 sequence revealed information indicating largetooth sawfish subpopulations. Evidence of restricted gene flow has also been found with largetooth sawfish among these geographic areas: Atlantic and Indo-West Pacific; Atlantic and eastern Pacific; and Indo-West Pacific and eastern Pacific. 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 of sub-populations 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 sub-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
h count than those collected from the eastern Atlantic. Data from separate protein and genetics studies indicates some evidence of distinction among sub-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 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 rivers in Australia, also suggesting potential philopatry (Lack et al., 2009). However, 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 historic 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. 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)
stribution and Abundance
Largetooth sawfish have the largest historic 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. 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). Older literature notes the presence of this species in Zanzibar, Madagascar, India, and the south-west Pacific (Fowler, 1941; Wallace, 1967; Taniuchi et al., 2003).
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, and magazine articles. The result of that search is 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 Mangoky River, Madagascar (Taniuchi et al., 2003).
Records from the 1960's and 1970's 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 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; they have also been recorded in the shark nets off Durban, South Africa (Wallace, 1967)
rom 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 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; they 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 (Devadoss, 1978). Largetooth sawfish are also recorded 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. While the species could not be confirmed, a survey of fishing landing sites and interviews with 99 fishers in Kenya, Nyungi (unpublished report to J. Carlson, NMFS 2007), found 71 reports of sawfishes over the last 40 years. The longest time series of largetooth sawfish catches is from the protective 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 in the last decade (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
ted 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 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 Santani (present day Irian Jaya, Indonesia). Juvenile largetooth sawfish had 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, New Guinea (Boeseman, 1956; Thorson et al., 1966). However, in a study by Munro (1967) in the Laloki River in the southeastern portion of New Guinea, no sawfish were captured (Berra et al., 1975). From 1967 to 1977, five largetooth sawfish were captured from the Indragiri River, Sumatra (Taniuchi, 2002). From 1970 to 1971, Berra et al. (1975) collected five largetooth sawfish from the Laloki River, Papua New Guinea.
More recently, 36 largetooth sawfish were captured in September 1989 in Papua New Guinea (Taniuchi and Shimizu, 1991; Taniuchi, 2002)
wfish were captured (Berra et al., 1975). From 1967 to 1977, five largetooth sawfish were captured from the Indragiri River, Sumatra (Taniuchi, 2002). From 1970 to 1971, Berra et al. (1975) collected five largetooth sawfish from the Laloki River, Papua New Guinea.
More recently, 36 largetooth sawfish were captured in September 1989 in Papua New Guinea (Taniuchi and Shimizu, 1991; Taniuchi, 2002). In a survey of the Fly River system, Papau New Guinea, 23 individuals were captured in 1978 (Roberts, 1978; Taniuchi and Shimizu, 1991; Taniuchi et al., 1991b; 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) and survey indicate largetooth sawfish are still present in these areas, although locals have noticed a decline in their abundance (Manjaji, 2002a).
The scarcity of records from Indonesia led to an increased effort to document species presence (Fowler, 2002). Anecdotal evidence suggests that sawfishes have not been recorded in Indonesia 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
Australian Waters
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). Faria et al. (2013) obtained a rostrum that was collected in Australia in 1960.
The most current reports of largetooth sawfish began in the 1980's
e (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). Faria et al. (2013) obtained a rostrum that was collected in Australia in 1960.
The most current reports of largetooth sawfish began in the 1980's. We found many more records of largetooth sawfish in Australia compared to other countries. A largetooth sawfish was captured from the Keep River, Australia in 1981 (Compagno and Last, 1999). 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. 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 many rivers (Mitchell, Gilbert, Archer, Nassau, Ord, and Staaten).
Juvenile largetooth sawfish in Australia use the Fitzroy River and other tributaries of the King Sound (Morgan et al., 2004) as nursery areas while adults are found more often offshore (Morgan et al., 2010a). Outside of the Fitzroy River and King Sound in western Australia, 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)
hile adults are found more often offshore (Morgan et al., 2010a). Outside of the Fitzroy River and King Sound in western Australia, 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). Despite the abundance of records from northern Australia, no sawfish have recently been captured within the Adelaide River, Australia, and abundance estimates from areas that have 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 have declined in recent years as 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 1960's, 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)
alia Shark Control Program shows a clear decline in sawfish catch over a 30 year period from the 1960's, 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; 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). The only recent reports of largetooth sawfish in this area are anecdotal reports from Columbia, Nicaragua, and Panama (R. Graham 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 U.S. (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 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)
cal presence of the largetooth sawfish is uncertain and early records might have been misidentified smalltooth sawfish (G. Burgess 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 from a survey of Lake Izaba, Guatemala from 1946 to 1947, and sawfishes were reported to be important inland fisheries (Saunders et al., 1950). The lone largetooth sawfish reported from Honduras was acquired from that country, but the true origin of the rostrum and the date of capture could not be confirmed (NMFS, 2010a).
In Atlantic drainages, largetooth sawfish were 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, pers. comm., as cited in Cook et al., 2005).
In the U.S., 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)
1982b; Castro-Augiree, 1978 as cited in Thorson, 1982b; Compagno and Cook, 1995; C. Scharpf and M. McDavitt, pers. comm., as cited in Cook et al., 2005).
In the U.S., 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 U.S., 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, and Sao
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, 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, and Caribbean records are very sparse (NMFS, 2010a)
re 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, and 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 primarily in fish markets at the Amazon-Orinoco estuaries (Charvet-Almeida, 2002; Burgess et al., 2009). A Lake Nicaraguan 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 Columbia 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), 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
fied records exist from Senegal (1841-1902), Gambia (1885-1909), Guinea-Bissau (1912), Republic of Guinea (1965), Sierra Leone (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, Guinea, Cote 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)
e 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 migration from another region (Phillips, 2012).
Natural History of Green Sawfish ( Pristis zijsron )
Taxonomy and Morphology
Pristis zijsron (Bleeker 1851) is frequently known as the narrow snout 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 slim 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; Morgan et al., 2011)
erally 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; Morgan et al., 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 aptly named green sawfish is greenish brown dorsally and white ventrally. This species might be confused with the dwarf or smalltooth et al., 2006c).
Habitat Use and Migration
The green sawfish mostly utilizes 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 its 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)
). However, Morgan et al. (2010a; 2011) report green sawfish do not move into freshwater for any portion of its 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).
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 age 1-5, green sawfish measure between 4 ft 2 in and 8 ft 5 in (128 and 257 cm) TL, based on the vertebral analysis of six 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
d 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; Morgan et al., 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 length of gestation
erell, 2008; Morgan et al., 2010a; Morgan et al., 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 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 genet 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
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 genet 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 Løppenthin, 1944; Smith, 1945; Misra, 1969; Compagno et al., 2002a and 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, and magazine articles. The results are summarized below by geographic area.
Indian Ocean
Green sawfish are widely distributed throughout the Indian Ocean with the first record in 1852 and several green sawfish were described near the Indian archipelago in the late 1800's (Van Oijen et al., 2007). Additional historical records include one female specimen captured in the Red Sea near Dollfus in 1929
magazine articles. The results are summarized below by geographic area.
Indian Ocean
Green sawfish are widely distributed throughout the Indian Ocean with the first record in 1852 and several green sawfish were described near the Indian archipelago in the late 1800's (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
Despite historic records, there are few current records of green sawfish in the Indian Ocean. We presume green sawfish are 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 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).
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 the elasmobranchs, and only two species of freshwater ray 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; no collection specifics were provided
scientific studies on the elasmobranchs, and only two species of freshwater ray 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; 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), and reported a decline of sawfish 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) reported no sawfish species from the south China Sea from 1923 through 1996. Anecdotal evidence suggests that sawfishes have not been recorded in Indonesia for more than 25 years (White and Last, 2010).
Australia
In Australian waters, records indicate green sawfish abundance is higher in the north compared to the south. The earliest record obtained was from the Queensland Museum in 1929 indicating that green sawfish were found in Moreton Bay, Queensland (Fowler, 1941).
We found a paucity of records for green sawfish during the middle part of the last century. Reports of green sawfish occur again in the 1980's when two green sawfish were captured from Balgal, Queensland, Australia in 1985 (Beveridge and Campbell, 2005). 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)
ring 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, Australia (Chisholm and Whittington, 2000) and in a survey from 1999 through 2001 by White and Potter (2004) one green sawfish was captured in Shark Bay, Queensland. Peverell (2005; 2008) noted the green sawfish was the least encountered species in a survey from the Gulf of Carpentaria. In 2008, no green sawfish were captured from the Roper River system, which drains into the western Gulf of Carpentaria, Northern Territory, Australia (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).
In summary, the limited data makes it difficult to determine the current range and abundance of green sawfish. However, 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 likely indicates the species may no longer be present. In Australian waters, based on our review, all sawfish species have undergone significant declines. 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, Australia, but are rarely found as far south as Townsville (Porteous, 2004). Green sawfish are currently found primarily along the northern coast of Australia.
Extensive surveys at fish landing sites throughout Indonesia since 2001 have failed to record the green sawfish (White pers. comm
s in Australia has contracted (Harry et al., 2011). Green sawfish have been reported as far south as Sydney, Australia, but are rarely found as far south as Townsville (Porteous, 2004). Green sawfish are currently found primarily along the northern coast of Australia.
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 pers. comm. to IUCN, 2012). However, lack of data from surveys and commercial fisheries throughout much of the remainder of the range suggests that the abundance of green sawfish has declined significantly and it is currently at only a small fraction of its historic abundance.
Natural History of the Non-listed Population(s) of Smalltooth Sawfish ( Pristis pectinata )
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.
The smalltooth sawfish has a thick body with a moderately sized rostrum. As with many other sawfishes, tooth count may vary 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 pelvic fins (Faria et al., 2013)
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 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 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 et al. (2011) reported electivity analysis on sawfish movements and demonstrated 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 30 degrees Celsius (86 degrees Fahrenheit), 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
le smalltooth sawfish had an affinity for water less than 3 ft (1.0 m) deep, water temperatures greater than 30 degrees Celsius (86 degrees Fahrenheit), 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 7 ft to 20 ft (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. Activity area decreased and range of movement increased at night, indicating 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 e t al., in review). All smalltooth sawfish generally remained in coastal waters at shallow depths (96 percent of their time at depths less than 32 ft; 10 m) and warm water temperatures (22-28 degrees Celsius (71.6-82.4 degrees Fahrenheit) within the region where they were initially tagged, travelling an average of 49 mi (80.2 km) from deployment to pop-off location on an average of 95 days. No smalltooth sawfish tagged within 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 is found in NMFS (2010b) indicates rapid juvenile growth for smalltooth sawfish for the first 2 years after birth
lltooth sawfish tagged within 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 is found in NMFS (2010b) indicates rapid juvenile growth for smalltooth sawfish for the first 2 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
Outside U.S. waters, 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 freshwater, and 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 6 months later the female aborted the pups for unknown reasons (McDavitt, 2006). In October 2012, a female sawfish gave birth to five live pups (J. Choromanski, pers. comm.).
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 7 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. populations of smalltooth sawfish
ic rates of increase at 7 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. populations of smalltooth sawfish.
Diet and Feeding
Smalltooth sawfish often use their rostrum saw in a side-sweeping motion to stun its prey, which may include small fishes, or 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 (NADH-2) sequences revealed no geographical structuring of smalltooth sawfish haplotypes (Faria et al., 2013). However, variation in the number of rostral teeth number per side was found in specimens from the western and eastern Atlantic Ocean (Faria et al., 2013).
Distribution and Abundance
Outside U.S. waters, smalltooth sawfish were thought to be historically found in South Africa, Madagascar, the Red Sea, Arabia, India, the Philippines, along the coast of wes

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