AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
Published online in Wiley Online Library
(wileyonlinelibrary.com). DOI: 10.1002/aqc.2394
Status and the potential for extinction of the largetooth sawfish
Pristis pristis in the Atlantic Ocean
JOANA FERNANDEZ-CARVALHOa,b,*, JOHANNA L. IMHOFFa,c, VICENTE V. FARIAd, JOHN K. CARLSONe
and GEORGE H. BURGESSa
a
Florida Program for Shark Research, Florida Museum of Natural History, University of Florida, Gainesville, USA
b
Centro de Ciências do Mar (CCMAR), Universidade do Algarve, Faro, Portugal
c
Department of Biological Science, Florida State University, Tallahassee, USA
d
Institute of Marine Sciences – Labomar, Federal University of Ceará, Fortaleza, Brazil
e
US National Marine Fisheries Service, Southeast Fisheries Science Center, Panama City Laboratory, Panama City, USA
ABSTRACT
1. Sawfishes currently are among the most threatened elasmobranchs in the world. Only two species inhabit
Atlantic waters: the largetooth sawfish (Pristis pristis) and the smalltooth sawfish (Pristis pectinata), both having
suffered dramatic declines in their ranges.
2. The goal of this study was to evaluate the status of P. pristis in the Atlantic, and estimate local extinction risk
based on historical and recent occurrence records. In order to accomplish these goals, a thorough search for
historical and recent records of P. pristis in the Atlantic was conducted, by reviewing scientific and popular
literature, museum specimens, and contacting regional scientists from the species’ historical range.
3. In total, 801 P. pristis records (1830–2009) document its occurrence in four major regions in the Atlantic:
USA (n = 41), Mexico and Central America (n = 535), South America (n = 162), and West Africa (n = 48).
Locality data were not available for 15 records.
4. Historical abundance centres were the Colorado-San Juan River system in Nicaragua and Costa Rica
(and secondarily Lake Izabal of Guatemala), the Amazon estuary, and coastal Guinea-Bissau.
5. Currently, the species faces drastic depletion throughout its entire former range and centres of abundance. It
appears to have been extirpated from several areas. The probability of extinction was highest in the USA, northern
South America (Colombia to Guyane), and southern West Africa (Cameroon to Namibia).
6. Currently, the Amazon estuary appears to have the highest remaining abundance of P. pristis in the Atlantic,
followed by the Colorado–San Juan River system in Nicaragua and Costa Rica and the Bissagos Archipelago in
Guinea Bissau. Therefore the protection of these populations is crucial for the preservation and recovery of the species.
Copyright # 2013 John Wiley & Sons, Ltd.
Received 10 December 2012; Revised 4 June 2013; Accepted 1 August 2013
KEY WORDS:
conservation evaluation; endangered species; estuary; coastal; brackish; fish
*Correspondence to: J. Fernandez-Carvalho, Centro de Ciências do Mar, Universidade do Algarve, Campus de Gambelas, FCT Ed. 7, 8005-139, Faro
Portugal. Email: jfcarvalho@ualg.pt
Copyright # 2013 John Wiley & Sons, Ltd.
J. FERNANDEZ-CARVALHO ET AL.
INTRODUCTION
Over the last century, significant exploitation and
habitat destruction has led to concerns that
populations of marine fish may be at risk of
extinction (Dulvy et al., 2003; Monte-Luna et al.,
2007). Detecting and determining local extinctions is
important to prevent further loss of the species in
other areas and to establish factors related to the
cause of the extinction. Dulvy et al. (2003) identified
133 local, regional and global extinctions of marine
populations and determined that exploitation caused
most extinctions followed by habitat loss, invasive
species, climate change, pollution and disease.
The most common approach to inferring the
extinction of a particular species is based on the
record of its sightings (Solow and Roberts, 2003).
For marine species, information about past
occurrences has been compiled from historical
sightings and museum records. Records are then
analysed in a probabilistic framework to generate
confidence intervals on the potential time of
extinction given the sighting records (Solow, 1993a,
2005; Burgman et al., 1995; Roberts, 2006). For
example, among elasmobranchs, Luiz and Edwards
(2011) concluded that the Galapagos shark,
Carcharhinus galapagensis has been locally extinct
since 1998 at St. Paul’s Rocks in the Atlantic
Ocean. Smalltooth sawfish, Pristis pectinata, were
determined to be extirpated in US waters from
Virginia to New York state from between 1929 and
1956 (Monte-Luna et al., 2009; NMFS (National
Marine Fisheries Service), 2009b). Reliably estimating
the spatial patterns and/or the timing of extinction
remains a major challenge for scientists, as the danger
of extinction can be overestimated because of
incorrect assumptions on population size.
The largetooth sawfish (Pristis pristis, Linnaeus,
1758) is one of the two species of the family
Pristidae occurring in the Atlantic Ocean. It is
found in tropical and subtropical waters in both the
western and eastern Atlantic, with northern and
southern range extremes largely dictated by
seasonal water temperature regimes. The western
North Atlantic distributional termini of both species
occur in the USA whereas those found in the
eastern Atlantic ranged as far north as Mauritania
in the north and Namibia in the south (Bigelow and
Schroeder, 1953; Faria et al., 2013).
Copyright # 2013 John Wiley & Sons, Ltd.
Like all species of sawfish, P. pristis has
significantly declined owing to overfishing, chiefly
as bycatch, and habitat loss (Fowler et al., 2005).
As a result P. pristis is listed as Critically
Endangered on the International Union for
Conservation of Nature Red List (Fowler et al.,
2005; Kyne et al., 2013) and as Endangered
Globally on the US Endangered Species Act (76 FR
40822). Despite being listed as Critically
Endangered and Endangered, respectively, the
status of largetooth sawfish was determined based
on anecdotal information. Thus there is a need both
to qualify and quantify historical records of
P. pristis in the Atlantic Ocean to provide a better
determination of its current status. Here, we report
on the results of an extensive search of historical
and recent records of P. pristis in the Atlantic
Ocean and provide a region by region analysis of its
current status and risk of extinction.
MATERIAL AND METHODS
Data collection
Sawfish records were collected between July and
September 2009, and records were individually
screened for accuracy. Species identification was
based on taxonomic characters described in
Bigelow and Schroeder (1953) for Pristis perotteti
(e.g. the number of rostral teeth). Following Faria
et al. (2013) all records of Pristis perotteti and
Pristis microdon from the Atlantic were considered
largetooth sawfish and as such are referred to as
Pristis pristis.
Records of P. pristis were gathered by consulting
primary literature, media, museums and local
expert scientists. Primary literature records and
P. pristis distribution information were obtained
by searching card catalogues from the libraries of
the key regional universities (University of
Florida, University of Texas, Texas A&M
University, University of South Alabama) within
the species’ historical US range. Library
databases and online search engines were used for
finding photographs and fishing and historical
blogs pertaining to P. pristis records. In addition,
two online historical newspaper databases
(Proquest Historical Newspapers database and
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
www.NewspaperArcive.com) were used to find
sawfish articles in local and regional newspapers.
Positive identifications from historical newspapers
were only possible when photographs were
available or rostral tooth counts were provided
in the story.
Twenty-seven museums were contacted to
obtain records of archived P. pristis specimens.
Additional specimen records from museums and
private collections were obtained from the
International Sawfish Encounter Database
(ISED) housed at Florida Museum of Natural
History and from Faria V. (unpublished data).
Owing to the lack of recent studies addressing
the status of P. pristis in the majority of its
range, scientists and fisheries managers were
queried directly about any sawfish records and
asked to provide their personal evaluations of the
regional status of the species (only regarding
Atlantic range) using a standardized set of
questions. The inquiry included questions about
their knowledge of the species, the existence of
any records (museum, literature, research, fisheries
data, evaluation of the species, etc.) in their
region, their personal evaluation and possible
colleagues in the region that should be contacted
because of their expertise. Hundreds of
researchers in the Gulf of Mexico and Caribbean
regions were contacted via the list server of the
Gulf and Caribbean Fisheries Institute (GCFI) in
English, Spanish, Portuguese and French.
Furthermore, 43 scientists (representing 13
countries: Aruba, Belize, Brazil, Colombia, Cuba,
Curaçao, Guinea-Bissau, Honduras, Mexico,
Nicaragua, Panama, Puerto Rico, Venezuela) were
contacted directly after being identified as
knowledgeable experts in their locale by colleagues,
though not all replied.
All the scientists who contributed sawfish
information are credited in the acknowledgement
section with their affiliation and country. Scientists
who replied but did not provide sawfish information
(e.g. only recommended other colleagues or said
they had no knowledge about sawfish) were not
included in the acknowledgement section. When
citing the personal communications of the
contributing scientists, the year when the personal
communication was provided is also given.
Copyright # 2013 John Wiley & Sons, Ltd.
Data analysis
Sawfish records were entered into a Microsoft Access
database, the ISED. Duplicate records, such as
museum records that were also cited within the
primary literature or multiple newspaper articles
referring to a single specimen, were screened and
removed. Species identifications were verified by
examination of museum specimens, photographs,
scientific notes, and/or interviews with relevant
observers, unless already recently confirmed by an
expert (Faria V., unpublished data). Dubious P.
pristis records, such as records suspected to be
misidentified P. pectinata or two records that were
suspected (but not confirmed) to be from the same
specimen (Springer and Woodburn, 1960), were
included in the database. However, only confirmed
(and not dubious) P. pristis records were included
in the analysis.
Given the nature of the data available, the
probability that local scale extinction has occurred
was determined using the time series of incidental
observations. The probability that local extinction
has occurred follows a stationary Poisson process:
¼1
t n
c
T
where the number of time intervals in which the
species was observed is n, the total number of
intervals sampled is T, and the number of intervals
up to the time the species was last observed is tc
(Solow, 1993a, b; Burgman et al., 1995; Grogan
and Boreman, 1998). Sensitivity analyses suggest
using minimum probability levels of 0.75 for
declaring a species to be in danger of local
extinction and 0.95 for locally extinct (Grogan and
Boreman, 1998). As estimates of the probability of
extinction are point estimates, a Monte Carlo
simulation was used to assess uncertainty in the
model input parameters. Probability density
functions were developed to describe the number
of time intervals in which the species was observed
and the number of intervals up to the time the
species was last observed. The extinction risk was
determined for six regions throughout the
historical range of P. pristis: USA, Central
America (Mexico to Panama), northern South
America (Columbia to French Guinea), Brazil,
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
northern West Africa (Mauritania to Nigeria) and
southern West Africa (Cameroon to Namibia).
For those areas with a high probability of
extinction (≥ 0.95), the upper bound (the upper
confidence limit at which the extinction probability
reaches the nominal 0.05 value) of the timing of
extinction was calculated using several methods
evaluated by Rivadeneira et al. (2009). The first
method made the strictest assumption that sightings
are a Poisson stationary process (Strauss and
Sadler, 1989; Solow, 1993a; McInerny et al., 2006),
with constant probabilities of occurrence and
sampling through time, and an abrupt collapse to
extinction. The other method (Roberts and Solow,
2003; Solow and Roberts, 2003) is least restrictive
and does not make any distributional assumptions,
although the independence of sightings is still
required (Rivadeneira et al., 2009).
In addition, the likelihood that P. pristis has become
extinct was evaluated for the six distributional areas
using the framework suggested by Butchart et al.
(2006) and used for elasmobranchs by Luiz and
Edwards (2011). This framework categorizes
the level of confidence of extinction based on
observational data by considering the time since the
species was last reported and the evidence for and
against extinction.
RESULTS
In total, 801 records of P. pristis throughout the
Atlantic were compiled during this study. These
comprised 41 US P. pristis records (from 1878 to
1961), 535 Mexico and Central America records
(1877 to 1998), 162 South America records (1830 to
2009) and 48 records from the West coast of Africa
(1841 to 2005) (Figure 1). Locality data were not
available for 15 additional P. pristis records.
United States of America
Records of P. pristis in US waters (n = 41, Table 1)
included 35 records from Texas, two ‘Gulf of
Mexico’ records which were probably from Texas,
Figure 1. Records of Pristis pristis obtained in this study. All countries that have P. pristis records are coloured and are delimited by a thick line.
Underlined country names represent countries for which there are records with only general location (i.e. country or state) available, thus the
records cannot be represented in the map. AM=Amazonas, AP=Amapá, BA= Bahia, ES=Espírito Santo, FL= Florida, LA=Louisiana,
MA=Maranhão, PA=Pará, RN=Rio Grande do Norte, RJ=Rio de Janeiro, SE=Sergipe, SP=São Paulo,TX= Texas.
Copyright # 2013 John Wiley & Sons, Ltd.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
Table 1. List of US Pristis pristis records. Records lacking state data are listed under ‘Gulf of Mexico’
State
County
Year
N
1910
1941
1943-1952
1
1
1
FL
FL
FL
Monroe
Monroe
Martin
LA
N/A
1918
1
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
TX
Aransas
Aransas
Brazoria
Cameron
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Galveston
Kleberg/Kennedy
Nueces
Nueces
Nueces
Nueces
Nueces
Nueces
Nueces
N/A
N/A
N/A
1940
1943
1942
1925
1929
1938
1938
1940
1940
1940
1942
1943
1943
1943
1951
1951
1957
1947
1917
1935
1938
1943
1943
1947
1961
1940
1948
-
1
1
1
1
1
1
1
1
1
1
6
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
GOM
GOM
N/A
N/A
1878
-
1
1
TL (cm)
Sex
Source
Mus. Cat. Num./ Reference
MUS
LIT
LIT
AMNH 11
Baughman (1943)
Bigelow and Schroeder (1953)
488
MUS/MED
SUI 17512
570
500
LIT
LIT
LIT
MED
MED
MED
MED
MED
MED
MED
LIT
LIT
LIT
LIT
MED
MUS
MED
MUS
MUS
MED
MED
MED
MED
LIT
LIT
MED
MED
MUS
OTH
MED
Baughman (1943)
Baughman (1943)
Baughman (1943)
Hoover (2008)
MUS
LIT
ANSP 17388
Bigelow and Schroeder (1953)
520
528
594
450
450
427
528
470
430
450
460
700
519
F
M
F
610
549
500
457
498
531
380
457
F
Baughman (1943) (Coll. by Reid, E. F.)
Baughman (1943)
Baughman (1943)
Baughman (1943)
MRAC A4-45-P
UAIC 4138.01-2
BMNH 1867.10
Hoover (2008)
Baughman (1952); Baughmann (1943)
Baughman (1952)
TAMUCC Uncat.
TX = Texas, LA = Louisiana, FL = Florida and GOM = Gulf of Mexico. ENC = records from encounter reports, LIT = Literature records,
MED = Media records, MUS = specimens from museums or other collections.
one record from Louisiana, and three records from
Florida. All the documented records correspond to
large animals ranging from 427 to 610 cm in total
length (TL). An unmeasured Galveston Beach
specimen documented in a 1951 home movie was
estimated to have been of 700 cm TL.
Several records documented sawfishes being
captured by commercial and recreational fishing
gear. Specifically, fishing gear information was
available for 21 records. Sawfishes captured in
commercial fishing gear were associated with
shrimp fisheries and caught either with trawl
(14%), seine (5%), shrimp net (10%) or unclassified
net (9%), while sawfishes captured recreationally
were caught using rod and reel (62%). It can
Copyright # 2013 John Wiley & Sons, Ltd.
be inferred that most captures occurred in
shallow inshore waters off the coast or
occasionally within bays, since most of the shrimp
trawling occurred near shore, and recreational
fishing was shore-based.
In Texas, P. pristis was found primarily in three
almost equidistant regions, from north-east to
south-west: Galveston–Texas City–Freeport (n = 22),
Corpus Christi–Port Aransas (n = 9) and Padre
Island–Laguna Madre (n = 2). Captures occurred
only in warm water months from May to October.
Pristis pristis was last recorded in the USA in Texas
in 1961. The estimated probability of extinction in US
waters was 0.99 (±0.16; Table 2). Depending on
methodology, the upper limit of time of extinction
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
Table 2. Probability and timing of extinction for Pristis pristis from six subregions from their historic range. Numbers in parentheses are the 95%
confidence intervals for probability of extinction. Abbreviations for extinction time indicate the method used: S&S = Strauss and Sadler (1989),
SOL = Solow (1993a), MCY = McInerny et al. (2006), S&R = Solow and Roberts (2003) and R&S = Roberts and Solow (2003)
US
Decade
1830
1840
1850
1860
1870
1880
1890
1900
1910
1920
1930
1940
1950
1960
1970
1980
1990
2000
2010
N
Mexico and
Central America
Time Extinct
N
Northern South
America
N
Time
Extinct
Brazil
Northern
West Africa
N
N
N
Time
Extinct
2
1*
1
1
1
3
2
2
2*
1*
1 + 3*
1
3
3
2
4
23
3
1
1 + 2*
1
1
1*
1
1963 SOL,MCY
1967 S&S
9
5*
496 + 4*
1 + 2*
1994 R&S
10
6
1 (years ago)
2017 S&R
1 (nowdays)
1*
1*
1976 SOL,MCY
1 + 1*
1 + 1*
4
1
8
2 + 3*
1
49 +1*
43
10
1
6 + 3*
2014 SOL, MCY
2020
2024 S&R
2025 R&S
2030
2040
…
2080
…
2320
Probability
Extinction
Southern
West Africa
2047 S&S
2044 S&S
2083 S&R
2326 R&S
0.99 (±0.16)
0.64 (±0.29)
0.99 (±0.06)
0.11 (±0.21)
0.25 (±0.20)
0.99(±0.77)
*Only date of citation was available, thus the date of the record could be earlier.
ranged from 1963 to 1994, although most methods
estimated dates in the 1960s. In addition, qualitative
evidence regarding population extinction did provide
evidence for local extinction (Table 3).
Mexico and Central America
Records of P. pristis from the Mexico and Central
America region (n = 535) are primarily from
Nicaragua and Costa Rica, with only 11
documented records of P. pristis found outside of
those two countries (Table 4). The documented
records include from north to south, Mexico
(n = 5), Guatemala (n = 5), Honduras (n = 1),
Copyright # 2013 John Wiley & Sons, Ltd.
Nicaragua (n = 483) and Costa Rica (n = 37).
Additionally, four old records were found
with no year information and attributed to
‘Caribbean Sea’.
Four Mexican historical records of P. pristis have
been documented for Tamaulipas, Veracruz, Tabasco,
and Campeche, all locations in the south-western Gulf
of Mexico (Zarur-Ménez, 1962 in Castro-Aguirre,
1978, 1999). Although precise dates are not
available, these records (particularly those from
Campeche) are from the 1950s and sawfishes are no
longer registered in literature from the l970s and
1980s (J. C. P. Jiménez, pers. comm., 2009). The
last record of P. pristis from Mexico was a 540 cm
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
Table 3. Evaluation of the qualitative evidence for and against local extinction of Pristis pristis from six subregions from their historic range in the
Atlantic Ocean, using the framework of Butchart et al. (2006)
Observational data
Types of evidence for extinction
For species with recent last records, the decline has been
well documented
Severe threatening processes are known to have occurred
The species possesses attributes known to predispose
taxa to extinction
Recent surveys have been apparently adequate given
the species’ ease of detection, but have failed to
detect the species
Types of evidence against extinction
Recent field work has been inadequate
The species is difficult to detect
There have been reasonably convincing recent local
reports or unconfirmed sightings
Suitable habitat remains within the species’ known
range, and/or allospecies or congeners may survive
despite similar threatening processes
?
US
Mexico and
Central America
Northern
South America
Brazil
Northern
West Africa
Southern
West Africa
N/A
Yes
N/A
Yes
Yes
N/A
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes
Yes?
Yes
Yes
No?
No?
No
No
No?
No
No
No
Yes?
No
Yes
Yes?
No
No
Yes
No
Yes
Yes
No
Yes
Yes
No
No
Yes
Yes
Yes
Yes
Yes
Yes
Existence of uncertainty owing to the scarcity of available information for the region.
TL specimen landed at Mujeres Island, Quinta Roo
in 1997 (J. L. Castillo-Geniz, pers. comm., 2009).
All Guatemalan records (n = 5) are the result of
Robert Rush Miller’s scientific collecting in Lake
Izabal in 1946–1947, with specimens catalogued
and curated at the National Museum of Natural
History (Table 4). In addition, Lake Izabal was
described by Saunders et al. (1950) as the only
Guatemalan lake where sawfishes were present,
being described as the most important inland
fishes of the country. No records were found
for any other Guatemalan Lake while conducting
this study. A historical P. pristis rostrum (UF
uncat.) lacking date and location information
originated from Honduras, and is the only record
from that country.
Historically, the Lake Nicaragua–San Juan
River–Colorado River system, in Nicaragua and
Costa Rica, was of particular importance for P.
pristis and the species was recorded from that
system in several studies (Gill and Bransford,
1877; Meek, 1907; Marden, 1944; Bigelow and
Schroeder, 1953; Thorson, 1982b). However,
neither historical nor modern records were found
from the Atlantic coast or drainages in Nicaragua,
other than the records of Lake Nicaragua and San
Juan River system (Montoya and Thorson, 1982;
Copyright # 2013 John Wiley & Sons, Ltd.
H. Guzmàn, pers. comm., 2009). Besides San Juan
and Colorado Rivers, P. pristis is known in Costa
Rica also from the Tempisque River and
tributaries of the San Juan River basin (Bussing,
2002). In addition, one isolated rostrum examined
in this study (UF uncat.) is from a specimen
caught in Tortuguero during the 1960s.
In total, 384 specimens caught by commercial
longliners were reported captured near the source of
the San Juan River at Lake Nicaragua, Nicaragua
(Thorson, 1976, 1982b). Nineteen of these were large
individuals, ranging in length from 324 to 394 cm
TL (Thorson, 1982b). In addition, 34 specimens
were reported taken by commercial longline fishers
in the lower reaches of the Colorado River near
Barra del Colorado (Costa Rica) between June 1966
and February 1977 (Thorson, 1982b). Two of these
animals were juveniles (127 and 130 cm TL).
Recent records of P. pristis in Nicaragua and
Costa Rica include immature animals. One neonate
was caught in the San Carlos region (near the
origin of the San Juan River on the southernmost
part of Lake Nicaragua), offering tangible evidence
that at least some pupping or nursery areas still
occur in the region. Additionally small specimens
(30 to 60 cm long) have been encountered during
Adpesca investigations in the western part of the
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
Table 4. Records of Pristis pristis from Mexico and Central America. Countries are ordered from north to south. MEX = Mexico, GTM = Guatemala,
HND = Honduras, NIC = Nicaragua and CRI = Costa Rica. CAR = Caribbean Sea. ENC = records from encounter reports, LIT = Literature records,
MED = Media records, MUS = museums or collections, PLIT = specimens from private collections cited in the literature
N
Country
State / Loc
Year
1
1
1
1
1
MEX
MEX
MEX
MEX
MEX
Quintana Roo
Campeche
Tamaulipas
Veracruz
Tabasco
2
1
1
1
GTM
GTM
GTM
GTM
Izabal
Izabal
Izabal
Izabal
1
HND
-
-
1
1
1
1
1
1
1
36
25
1
6
3
1
384
6
3
1
1
2
1
1
1
1
1
1
1
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
NIC
Granada
R.San Juan/Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Rivas
Granada/ Lk. Nicaragua
Granada/ Lk. Nicaragua
Granada/ Lk. Nicaragua
Granada/ Lk. Nicaragua
Granada/ Lk. Nicaragua
R.San Juan/Lk. Nicaragua
1877
1942
1943
1943
1943
Late 1960s
1960
1968
1968
1968
1968
1968
1969
1966-1977
1980
1980
1985
1991
1998
1998
1998
"Nowdays"
"years ago"
-
1
34
2
CRI
CRI
CRI
2
1
1
CAR
CAR
CAR
1997
Pre-1962
Pre-1978
Pre-1978
Pre-1978
1946
1947
1947
1947
Rivas/ Lk. Nicaragua
Rivas/ Lk. Nicaragua
Rivas/ Lk. Nicaragua
Chontales/ Lk. Nicaragua
Granada/ Lk. Nicaragua
Lk. Nicaragua
Rivas/ Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Lk. Nicaragua
Limon
540
Sex
Source
Mus. Cat. Number/Reference
F
ENC
LIT
LIT
LIT
LIT
Castro-Aguirre
Castro-Aguirre
Castro-Aguirre
Castro-Aguirre
MUS
MUS
MUS
MUS
USNM 00111423
USNM 00111443
USNM 00393616
USNM 00146543
MUS
UF Uncat.
LIT
LIT
MUS
MUS
MUS
PLIT
MUS
LIT
LIT
LIT
LIT
LIT
MUS
LIT
LIT
LIT
MED
PLIT
PLIT
PLIT
PLIT
MED
MED
LIT
LIT
LIT
Gill and Bransford (1877)
Thorson (1976)
USNM 00111367
USNM 00111369
USNM 00120468
Faria et al. (2013)
TU 22989
Thorson (1982b)
Thorson (1982b)
Thorson (1982b)
Thorson (1982b)
Thorson (1982b)
AMNH 55624A
Thorson (1976, 1982b)
Thorson (1982b)
Thorson (1982b)
Faria et al. (2013)
Faria et al. (2013)
Faria et al. (2013)
Faria et al. (2013)
www.nicaraguafishing.com
www.nicaraguafishing.com
Gill and Bransford (1877)
Meek (1907)
Bigelow and Schroeder (1953)
MUS
LIT
MED
UF Uncat.
Thorson (1976, 1982b)
www.jjphoto.dk
LIT
MUS
MUS
Bigelow and Schroeder (1953)
BMNH 1923.11.13.1
BMNH 1843.2.1.8
137.6
140
F
732
137.6
M
Juv
Juv
Juv
Adult
Adult
F
M
F
M
M
226-283
226-284
M
F
100
1960
1966-1977
-
lake (no adults have been observed) (Moncada,
2006). Finally, two records for Costa Rica
correspond to two isolated rostra (one of them
probably from a young-of-the-year).
Overall for Mexico and Central America, the
probability of extinction from this area is 0.64
(±0.29; Table 2). The qualitative analysis showed
Copyright # 2013 John Wiley & Sons, Ltd.
TL (cm)
F
F
(1978, 1999)
(1978, 1999)
(1978, 1999)
(1978, 1999)
evidence both for and against extinction in the
region, highlighting that although there are some
recent local records the status of the species seems
very threatened (Table 3). The existence of
adequate field surveys is uncertain owing to the
lack of information available for most of the
countries in the region.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
Table 5. Records of Pristis pristis from South America. Countries are ordered geographically from north to south. COL = Colombia, VEN = Venezuela,
TTO = Trinidad, GUY = Guyana, SUR = Suriname, GUF = French Guiana. ENC = records from encounter reports, LIT = literature records,
MUS = specimens from museums or other collections
N
Country
State / Loc
1
COL
Atlántico
2
1
2
1
1
COL
COL
COL
COL
COL
Bolívar
Santander
Córdoba
Bolívar
Magdalena
Pre-1945
Pre-1945
Pre-1964
Pre-1976
Pre-1985
1
1
1
1
1
1
VEN
VEN
VEN
VEN
VEN
VEN
Zulia
Zulia
Zulia
Delta Amacuro
-
1894
1900
1903
1903
1962
Pre-1942
1
TTO
-
Pre-1940
1
GUY
-
1
1
1
1
1
SUR
SUR
SUR
SUR
SUR
-
1
1
GUF
GUF
Maroni river
Cayenne river
Year
TL (cm)
Source
1967
600
LIT
LIT
LIT
LIT
LIT
LIT
Martínez [1978] in Grijalba-Bendeck
et al. (2009)
Miles (1945)
Miles (1945)
Mejía and Acero (2002)
Mejía and Acero (2002)
Mejía and Acero (2002)
ENC
MUS
MUS
MUS
LIT
LIT
USNM 00232690
USNM 00232688
USNM 00232689
Cervigón (1966)
Bigelow and Schroeder (1953)
LIT
Ramjohn (1999)
-
MUS
BMNH 1889.11.15.1
1830
1856
1878
1962
-
LIT
LIT
MUS
MUS
MUS
Fowler (1919)
Fowler (1919)
ANSP 17390
RMNH D3079
RMNH D2674
1830
1929
MUS
LIT
USNM 00111169
Puyo (1949)
South America
In total, 162 P. pristis records were found for South
America, with the vast majority of records coming
from Brazil (Tables 5 and 6). Much lower numbers
of P. pristis were recorded in the remaining
countries of the species’ range in South America:
Colombia (n = 8), Venezuela (n = 6), Guyana
(n = 1), Suriname (n = 5), French Guiana (n = 2),
Trinidad (n = 1) and Brazil (n = 139).
Eight records were found for Colombia in the
Caribbean states of Córdoba (n = 2), Bolívar
(n = 3), Atlántico (n = 1) and Magdalena (n = 1),
and the neighbouring inland state of Santander
(n = 1) (Table 5). The only record for which
specific date and size are available is a 600 cm TL
specimen captured in a competition in 1967 in
Barranquilla (Atlántico) (Martínez, 1978 in
Grijalba-Bendeck et al., 2009). Despite this low
number of specific records, several texts document
the species’ occurrence in Colombia (Miles, 1945,
1947; Acero et al., 1986; Flórez-González, 1986;
Mejía and Acero, 2002). Listings of P. pristis (as
Copyright # 2013 John Wiley & Sons, Ltd.
Mus. Cat. Number/Reference
610
250
P. microdon) from the Magdalena River estuary
(Miles, 1945, 1947) reported differences in rostral
tooth counts and dorsal fin placement, clearly
indicating that both P. pristis and P. pectinata
were present. The species was present in the
marine environment (Frank and Rodriguez, 1976
in Mejía and Acero, 2002; Alvarez and Blanco,
1985) and in fresh water (Dahl, 1971; Sinú River).
Miles (1947) referred to a Magdalena River record
‘some 600 km from the mouth’ as P. pectinata, but
a record this far upriver likely refers to P. pristis.
Sawfishes (Pristis spp.) in Colombia have been
described in the literature both as having low
economic value (Dahl, 1971; Grijalba-Bendeck
et al., 2009) and as commercial species with
importance in the exportation to Asian countries
(Salazar, 1978 in Caldas et al., 2009). However, no
sightings or captures of the species have occurred
recently (Mejía and Acero, 2002; Grijalba-Bendeck
et al., 2009) and local scientists state that P. pristis
has been eradicated from Colombian waters long
ago (A. Acero, pers. comm., 2009; F. Gomez, pers.
comm., 2009).
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
Table 6. Records of Pristis pristis from Brazil. States are ordered geographically from north to south and from west to east. ENC = records from
encounter reports, LIT = literature records, MUS = specimens from museums or other collections. PLIT = specimens from private collections cited
in the literature
N
State / Loc
Year
48
40
13
1
2
1
1
1
1
1
1
1
1
3
1
Northern Brazil
Northern Brazil
Northern Brazil
Amazon River
Amapá
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
Amazonas
1999
2000
Pre-1934
Pre-1953
1955
1964
1965
1966
1967
1971
Pre-1974
-
1
1
1
1
2
1
4
Pará
Pará
Pará
Pará
Pará
Pará
Pará
1865
1865
1873
1878
2002
Pre-1998
1
1
1
Maranhão
Maranhão
Maranhão
1998
2009
1983-1986
1
Rio Grande do Norte
1
TL (cm)
Sex
Source
Mus. Cat. Number/Reference
PLIT
PLIT
PLIT
PLIT
PLIT
LIT
LIT
LIT
LIT
LIT
LIT
LIT
LIT
LIT
MUS
Faria et al. (2013)
Faria et al. (2013)
Faria et al. (2013)
Faria et al. (2013)
Faria et al. (2013)
Thorson (1974)
Bigelow and Schroeder (1953)
Thorson (1974)
Thorson (1974)
Thorson (1974)
Thorson (1974)
Thorson (1974)
Thorson (1974)
Thorson (1974)
INPA Uncat.
M
M
F
MUS
MUS
MUS
MUS
MUS
LIT
MCZ-302
MCZ-302
MCZ 668
USNM 00110174
INPA uncat 1 to uncat 2
Carvalho and McEachran (2003)
Charvet Almeida and Almeida (2008);
Faria et al. (2013)
F
F
ENC/LIT
ENC
LIT
Almeida et al. (2006)
MUS/LIT
SU 34468
Sergipe
MUS
ZMB 32533
1
Bahia
MUS/LIT
ANSP 17389
1
1
Espírito Santo
Espírito Santo
MUS
PLIT
MCZ-667
Faria et al. (2013)
1
Rio de Janeiro
LIT
Bigelow and Schroeder (1953)
1
São Paulo
LIT
Bigelow and Schroeder (1953)
154
255
175
235
160
116
225
190
F
86.2
95.6
81
700
700
1913
1865
-
F
1
-
1929
MUS
AMNH 49528
1
-
1978
MUS
NCSM 46131
The earliest of the six P. pristis records from
Venezuela is a rostrum from a large adult. This
specimen (believed to be from Venezuela) is
estimated to have been in the possession of a US
family since 1894. Three of the records were from
the Maracaibo region, an area where the species
was reported to have been abundant and
frequently taken (Cervigón, 1966). The last
documented record of P. pristis in Venezuela was
an individual captured in 1962 (Cervigón, 1966),
Copyright # 2013 John Wiley & Sons, Ltd.
and an additional record from before 1942 has no
specific locality data.
Although the documented records of P. pristis in
Venezuela lack detailed locality data, the species
has been reported to be abundant and frequently
taken in fisheries in the Gulf of Venezuela and
Lake Maracaibo (north-western Venezuela), as
well as the Gulf of Paria (north-eastern Venezuela)
and the area south of Trinidad, but it was rare
and only captured occasionally in the Margarita
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
region (north-eastern Venezuela) (Cervigón, 1966).
Before their decline, sawfishes (Pristis spp.) were
mostly caught in the mouth of the Orinoco River
and nearby waters (R. Tavares, pers. comm., 2009).
They were also reported as frequently caught in
north-west Venezuela, specifically on the coast of
Falcon state and in the Gulf of Venezuela, where
interviewed fishermen still have rostra (R. Tavares,
pers. comm., 2009). According to Cervigón (1993),
the main threat to the species was shrimp trawling.
Both species of sawfishes stopped being caught in
artisanal fisheries in the early 1990s, according to
fishermen, and there is no recent information
about sawfishes in the country (R. Tavares, pers.
comm., 2009).
There is a single 19th century museum record of
P. pristis from Demerara, Guyana. Regarding
Suriname, three museum specimens held in public
collections were located. In addition to these,
Fowler (1910, 1936) noted the species from
Suriname and later recorded an additional
Suriname P. pristis specimen from about 1830
(Fowler, 1919). Two records from French Guiana
were found, an 1830 museum specimen from the
Maroni River, which forms the border between
French Guiana and Suriname, and one 1929
record from the Cayenne River (Puyo, 1949).
Ramjohn (1999), citing Mendes (1940), listed the
P. pristis (as P. microdon) without comment from
Trinidad. However, local scientists haven’t heard
of any sighting of the species in the recent decades
while scientific diving around the Dutch Antilles
and Aruba (M. G. G. Grol, pers. comm., 2009).
In Curaçao, only one sawfish specimen (Pristis sp.)
was collected during scientific sampling from the
country’s largest mangrove bay (Schottegat Bay) at
the turn of the 20th century and another single
specimen was encountered in St. Joris Bay in the
early 1970s, but no species is detailed in either case
(A. O. Debrot, pers. comm., 2009). Although
sawfishes were probably never abundant in
Curaçao, they should now be considered extirpated
in that country (A. O. Debrot, pers. comm., 2009).
According to qualitative analysis there seems to be
more evidence supporting local extinction than
against it for the northern areas of South America,
although there is some uncertainty caused by the
lack of information available for this subregion. The
Copyright # 2013 John Wiley & Sons, Ltd.
reduced number of records found resulted in the
probability of extinction of 0.99 (±0.06; Table 2).
Consequently, the timing of extinction varied
considerably and ranged from 1976–2326.
Brazil’s historical and recent records bearing
location data come from the following states (west
to east and north to south): Amapá (n = 2),
Amazonas (n = 12), Pará (n = 11), Maranhão (n = 3),
Rio Grande do Norte (n = 1), Sergipe (n = 1), Bahia
(n = 1), Espirito Santo (n = 2), Rio de Janeiro (n = 1),
and São Paulo (n = 1). In addition, there are 101
more records from ‘northern Brazil’ for which there
is no state information (Table 6).
As these numbers suggest, P. pristis has been
historically more abundant in the northerly
tropical fraction of its range in Brazil, a region
comprising the Amazon River and the states of
Amazonas, Amapá, Pará and Maranhão. In this
region, neonates, juveniles and adults can be found
(Santos and Val, 1998; Almeida and Carneiro,
1999; Charvet-Almeida, 2002; Nunes et al., 2005;
Charvet-Almeida and Faria, 2008). The Amazon
River basin area and adjacent waters have long
been recognized as an area inhabited by
sawfishes, and P. pristis specifically (Bates, 1964;
Marlier, 1967; Furneau, 1969; Thorson, 1974). In
the Amazonas state, which includes the middle
section of the Amazon basin, P. pristis have been
taken as far inland as 2000 km from the coast
(Santos and Val, 1998) as well as several other
locations downstream to the Amazonas–Pará
state border.
The earliest records from Brazil come from Pará,
which contains the estuary and lower reaches of
the Amazon River. Two relatively recent records
of P. pristis were reported from Maranhão, one
adult female caught by gillnet in 1998 (Almeida
and Carneiro, 1999; Nunes et al., 2005) and
another adult female caught in 2009, which was
reported aborting 20–25 young (Jorge Nunes,
pers. comm., 2009). To the south, documented
reports are far more limited and historic in
nature, with only one record per state. The
southernmost record is a historical specimen from
São Paulo (Ribeiro, 1918 in Bigelow and
Schroeder, 1953; Gadig, 1998). At present, there
is no quantitative information on the current status
of P. pristis in northern Brazil. Fishermen have
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
Table 7. Records of Pristis pristis from Africa. North to south: MRT = Mauritania, SEN = Senegal, SENG = Senegambia, GMB = Gambia,
GNB = Guinea Bissau, GIN = Republic of Guinea, SLE = Sierra Leone, LBR = Liberia, CIV = Côte D’Ivoire, COG = Congo, COD = Democratic
Republic of the Congo, AGO = Angola. LIT = literature, MUS = museum
N
Country
Location
Year
RTL (cm)
Source
Museum Catalogue Number
Nothern West Africa
1
MRT
Tidra
1
1
1
1
1
1
1
1
1
1
1
SEN
SEN
SEN
SEN
SEN
SEN
SEN
SEN
SEN
SEN
SEN
2
SENG
Pre-1885
1
1
1
GMB
GMB
GMB
Pre-1909
Pre-2005
-
1
1
3
2
2
1
1
1
GNB
GNB
GNB
GNB
GNB
GNB
GNB
GNB
1
1
1
1
GIN
GIN
GIN
GIN
Soro
Kamsar
Bongolon
1
1
SLE
SLE
Bonthe
1
1
LBR
LBR
1
1
CIV
CIV
Fadiouth
Casamance
Joal
Joal
Bissagos Arch.
Bolama
Bolama
Bissagos Arch.
Bissagos Arch.
Gêba River, Gêba
Sassandra
Sassandra
1980s
LIT
Robillard and Séret (2006)
LIT
LIT
LIT
LIT
LIT
LIT
LIT
LIT
LIT
MUS
MUS
Bigelow and Schroeder (1953)
Bigelow and Schroeder (1953)
Bigelow and Schroeder (1953)
Bigelow and Schroeder (1953)
Bigelow and Schroeder (1953)
Bigelow and Schroeder (1953)
Robillard and Séret (2006)
Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
MNHN A-9699
IFAN R977
LIT
Bigelow and Schroeder (1953)
LIT
LIT
MUS
Bigelow and Schroeder (1953)
Ballouard et al. (2006); Robillard and Séret (2006)
BMNH 1885.1.31.30
LIT
LIT
LIT/MUS
LIT/MUS
LIT/MUS
LIT
LIT
MUS
Ballouard et al. (2006)
Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
Ballouard et al. (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
BMNH 1912.4.1.1
LIT
LIT
LIT
LIT
Ballouard et al. (2006); Robillard and Séret (2006)
Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
Ballouard et al. (2006); Robillard and Séret (2006)
LIT
MUS
Ballouard et al. (2006)
BMNH 2004.11.27.51
1881
1927
MUS
MUS
RMNH D3075
RMNH D3078
1902
Pre-1923
MUS
LIT
MNHN 1902-255
Bigelow and Schroeder (1953)
Pre-1841
Pre-1861
Pre-1865
Pre-1870
Pre-1882
Pre-1902
1980s
1990
Pre-2005
-
1980
1980´s
1983
2003
2004
2005
Pre-2005
1965
1980´s
1983
1989
2003
-
19
106
105
11
117
92
92
9
86
108
89
74
8
Southern West Africa
3
COG
1
Pointe-Noire
1958
CongoBasin
80-103
MUS
MNHN 2003–2612 to 2613
1951
MUS
MRAC 66639
2
1
1
COD
COD
COD
Banana
Banana
Congo R. estuary
1951
1959
-
MUS
MUS
MUS
MRAC 68038 and 68039
MRAC A4-45-P-12
MRAC-A4-45-P-2
1
1
AGO
AGO
Luanda
Luanda
1951
MUS
MUS
MRAC 74723
ZMB 16109
Copyright # 2013 John Wiley & Sons, Ltd.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
reported a decline in the region beginning as early as
the late 1980s (Charvet-Almeida, 1999, 2002).
Qualitative evidence against population extinction
did not support local extinction for Brazil (Table 3).
It should be noted that although we acknowledge
the existence of some research in the area when
compared with other subregions of the Atlantic,
surveys and field work were considered to be
inadequate in the qualitative analyses due to the
vast extent and complex morphology of the area.
Regarding the quantitative analysis, given the
recent records of P. pristis from the Amazon
estuary, the probability of extinction is 0.11 (±0.21).
Africa is 0.25 (±0.20). Accordingly, qualitative
evidence against population extinction did not
support local extinction for this subregion (Table 3).
However, in southern areas of West Africa
(Cameroon to Namibia) qualitative evidence
seems to be stronger for extinction than against,
although uncertainty because of the absence of
data in this subregion should be acknowledged. It
is likely that P. pristis has been extirpated
(0.99 ± 0.77) but the upper limit of the projected
time for regional extinction calculated was highly
variable, probably a result of the lack of records
(2014–2044).
West Africa
Historical museum specimens and literature records
of P. pristis on the west coast of Africa exist from
Mauritania (n=1), Senegal (n = 11), Gambia
(n = 3), Guinea-Bissau (n = 12), Republic of
Guinea (n = 4), Sierra Leone (n = 2), Liberia
(n = 2), Côte d’Ivoire (n = 2), Congo (n = 3),
Democratic Republic of the Congo (n = 4), and
Angola (n = 2) (Table 7). In addition there is a 1951
record from the ‘Congo Basin’ that could be from
either Congo, Democratic Republic of the Congo
or Angola. Virtually all these records lack definitive
locality and date, and specimens from this period
often have been lost. That said, P. pristis was at one
time relatively common along the west coast of
Africa, particularly in areas with riverine estuaries.
Robillard and Séret (2006) reported the
unpublished historical notes gathered by Jean
Cadenat, founder of the Musée de la Mer in
Gorée (Senegal) in the 1950s. In addition, the
same authors and Ballouard et al. (2006) examined
13 P. pristis rostra from 1965–2005 found at
various locales along the coast of West Africa.
Their reviews of the recent status of sawfishes
from Mauritania to Republic of Guinea revealed
that sawfishes, P. pectinata and P. pristis (referred
to as P. microdon), were relatively common in the
past but are now rarely caught or observed. The
last documented regional capture the authors
found was from 2005 in Nord de Caravela at
Guinea-Bissau. Because of these recent records in
areas around Guinea-Bissau, the likelihood that
P. pristis is extirpated from areas in northern West
Copyright # 2013 John Wiley & Sons, Ltd.
DISCUSSION
Historically, P. pristis ranged from the Gulf of
Mexico to south-east Brazil in the western
Atlantic, and from Mauritania to Angola in the
eastern Atlantic. Over the last century this
distribution has been considerably truncated and
abundance levels appear to have significantly
decreased, based on encounter data, museum
records, literature, personal communication with
scientific colleagues, and quantitative and
qualitative analyses performed. Nevertheless, it
should be acknowledged that even though a
thorough and intensive search for records was
conducted in the present study, there are catches
and sightings that go unreported, especially in
more isolated regions of Central and South
America and of Africa. Even in the regions where
sawfish are still caught or landed periodically,
owing to remoteness and consequent lack of
documentation very few records reach the media
and very often only large specimens are reported
since juveniles receive less attention. As a result,
information reported in this study should be used
with caution.
That said, the decline of P. pristis seems to follow
a similar pattern exhibited by P. pectinata in the
USA, with the extremes of its distributional ranges
having sequentially disappeared in the face of
fishing pressure and habitat loss (NMFS, 2009a, b).
Historical centres of abundance of P. pristis were
generally equatorial and influenced by water
temperatures, but the presence of large rivers and
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
riverine and lacustrine estuaries appears to have
been equally essential. Pristis pristis seems to
prefer freshwater and brackish water habitats
more than its relative P. pectinata, although adults
frequently occur in higher salinity, nearshore
continental waters (Compagno and Cook, 1995;
Zorzi, 1995). As is the case with P. pectinata
(Poulakis and Seitz, 2004; Carlson et al., 2007), in
addition to estuaries and inshore bars, the
presence of mangroves seems to be an indicator of
good habitat for P. pristis, since several records
occurred in areas known to have mangroves (e.g.
Amazon river estuary, Bissagos Archipelago).
Furthermore, it is possible that the habitat of the
Amazon estuary and the Bissagos Archipelago in
Guinea-Bissau, might have served as refuges for
the species, offering protection in the form of
complex coastlines heavily indented by inlets and
extensive mangroves.
United States of America
All P. pristis captured in US waters were large
individuals occurring at the northern extremity of
their distributional range only during warm water
months (May to October). It is apparent that this
species historically was represented in the Gulf of
Mexico by seasonal transients that took advantage of
Texas waters, only occasionally wandering eastward
to Louisiana and Florida. Texas historically has had
more documented P. pristis encounters than any
geographic region outside of Nicaragua and Brazil,
although this probably is partially an artefact of the
maturation of regional journalism and fishing
pressure (both commercial and recreational) between
1917 and 1961. Nevertheless, when considered in
conjunction with its regularity of occurrence before its
marked reduction in abundance, it appears that at
one time the western Gulf of Mexico was an
important area for this species, especially for adults
and subadults.
The highest number of P. pristis records in Texas
was from the north-east coast, followed by the central
coast, with few records from the south-western region
adjacent to the Mexican border. Although it might be
expected that the south-west would have more
records, the lower number of encounters may
correspond with the historical freshwater inflow
Copyright # 2013 John Wiley & Sons, Ltd.
pattern of the region. This pattern has a gradient
from highest inflows in the north-east to lowest along
the south-west coasts of the state (Longley, 1994).
Freshwater inflow is important to productivity of the
region (TDWR (Texas Department of Water
Resources), 1982) and the influence of lower salinity
water is especially important to P. pristis (Thorson,
1974). Thus the presence of higher-flow riverine
estuaries may have played a role in attracting larger
numbers of P. pristis to the Galveston–Texas
City–Freeport and Corpus Christi–Aransas regions,
than areas to the south-west. However, in the absence
of catch-per-unit-effort data, the encounter records
may simply be reflective of higher fishing efforts
associated with higher human densities along the
north-east and central Texas coastline.
The ratio of P. pectinata to P. pristis records
from Texas in the ISED is roughly 1:1, so it is
likely that scientific literature and print media
reports from the region currently unattributed to
species represents both species in approximately
equal numbers. This is consistent with the
observations of fishermen who told Baughman
(1943) that the two species were present in equal
proportions. Perhaps half of the 178 Pristis spp.
records found for Texas could represent P. pristis
records, highlighting the former presence of the
species in Texas waters. Unlike P. pectinata, which
has a refugium in the waters of southern Florida,
P. pristis apparently is extinct in US waters, with
last documented records in Florida in 1941,
Louisiana in 1917, and Texas in 1961. Since the
evidence suggests that P. pristis encountered in US
waters were probably seasonal migrants from core
areas of their range, it is possible that the decline
of the species in this peripheral region of its range
is a result of its decline in Central America, with a
smaller core population translating to a smaller
number of seasonal migrants. In addition,
extensive commercial shrimp trawling activities
and recreational fishing may have played a role in
the decline of the species in US waters.
Mexico and Central America
Given the habitat available along the Mexico and
Central America coastlines, it might have been
expected that more historical records of P. pristis
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
would have been found. However, the potential for
taxonomic confusion with P. pectinata is high, and
scientific collecting and popular reporting during
times of former relative abundance may have
caused P. pristis to be under-represented in
museums and the media. A unifying theme
presented by regional biologists is ‘no information
available’. While this may be partially attributable
to lack of regional data on P. pristis, even those
with good knowledge of local fauna and fisheries
indicate that the species is seldom if ever
encountered. The lack of flowing rivers in some
areas of the region (especially the Yucatán) may
have limited available habitat for the species.
However, it is clear that the decline observed in
Texas has also occurred to the south. The most
recent sawfish capture in Mexico (1998) was the
first encounter in 15 years of observing artisanal
fisheries (J. L. Castillo-Geniz, pers. comm., 2009).
The importance of the rivers of Costa Rica and
Nicaragua to P. pristis cannot be overestimated.
Clearly this was and potentially still is a core area
that provides both nursery and adult habitat critical
to P. pristis (Thorson, 1976, 1982a, b; Bussing,
2002; McDavitt, 2002). It also may very well
have served as a population source for adult
movements to the north and south-east, much as
south Florida has served that function for
P. pectinata movements along the US Atlantic
coast (and probably north-eastern Gulf of Mexico)
(NMFS, 2009b).
The decline in this region has occurred as a direct
result of overfishing as the species was heavily fished
along this lake–river system in the 1970s, reaching
population collapse in the 1980s (Thorson, 1982a;
McDavitt, 2002). As a result, the Nicaraguan
government imposed a temporary moratorium on
targeted fishing for sawfish in Lake Nicaragua in
the early 1980s (Thorson, 1982a). However,
despite the ban sawfishes continued to be killed as
bycatch in gillnets (McDavitt, 2002). For example,
a local artisanal fisher indicated he usually
captures four to six sawfishes per year (1998
interview, McDavitt, 2002). Protection was
strengthened in 2006 with a Nicaraguan ban on
fishing for sawfish, but only in Lake Nicaragua
(Charvet-Almeida et al., 2007) and this ban still
does not decrease the bycatch in other fisheries. It
Copyright # 2013 John Wiley & Sons, Ltd.
is probably because of the problem of bycatch in
fisheries and the low productivity of sawfish
(Simpfendorfer, 2000) that recovery of P. pristis
will be slow in the region.
South America
There is little indication that P. pristis are found in
anything other than low and declining numbers
across the northern rim of South America from
Colombia to the Guianas or in the continental
islands of Aruba, Curaçao and Trinidad. One
would expect that the Orinoco River in Colombia
and Venezuela would have been an important area
but records fail to pinpoint this area as equal in
importance to higher abundance areas to the west
and south, particularly the Amazon River basin.
Nevertheless, the lack of recent P. pristis records
in northern South America may be a partial
artefact of limited reporting, but given the declines
of this species throughout the remainder of its
Atlantic range, and the patterns exhibited by other
pristid species worldwide, it seems safe to interpret
an occurrence pattern lacking in recent observations
as a significant decline in abundance. However,
further studies are needed to permit a more
complete assessment of the area.
By contrast, although information about historical
and current levels of P. pristis abundance in Brazil is
unavailable, recent records, personal accounts and
qualitative analysis (Charvet-Almeida and Faria,
2008; Charvet, pers. comm., 2008/2009; Faria
et al., 2013), indicate that the Amazon estuary is
presently the location with the highest number of
recent records and more information available for
the species in the Atlantic, with the present
distribution being possibly restricted to the states
of Amapá, Pará and Maranhão (and maybe
Amazonas).
The remaining population of P. pristis in northern
Brazil has faced several threats in past decades
(between late 1990s and mid-2000s). The species has
been a major component of bycatch in commercial
and artisanal fisheries (Charvet-Almeida and Faria,
2008) with meat and fins traded in markets
(Charvet-Almeida, 1999, 2002; McDavitt and
Charvet-Almeida, 2004). In addition, small saws
were sold as curios for tourists and as folk medicine
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
to treat asthma (Charvet-Almeida, 1999, 2002) while
large saws brought high prices in international
trade (Charvet-Almeida, 2002; McDavitt and
Charvet-Almeida, 2004; Charvet-Almeida and
Faria, 2008). Additional threats included habitat
destruction and modification and the interest of the
ornamental trade (aquaria) in P. pristis neonates
(Charvet-Almeida and Faria, 2008).
Since the mid-2000s, modifications to Brazilian
legislation may have enabled some protection to
P. pristis in northern Brazil. In 2004, P. pristis
became a protected species under the Brazilian
legislation (IN – MMA # 5), and capture or trade
of any kind is prohibited. In addition, at least
three subsequent new rules regulating gillnet
fisheries (IBAMA # 166; 2007), and banning
ornamental trade (IN- MMA # 202; 2008) and
shark finning (INI - MPA # 14; 2012) may also
provide further protection. Finally, several marine
protected areas in northern Brazil may offer
protection to the species (Charvet-Almeida and
Faria, 2008). In summary, the current protection
measures in place for P. pristis in Brazil seem to be
stronger than in the preceding decades, although
enforcement is expected to be difficult in that
region. For example, recently P. pristis (as P.
perotteti) meat was on sale in two fish markets of
Pará labelled as ‘shark’ fillets even though its sale in
Brazil is prohibited (Melo Palmeira et al., 2013).
West Africa
The west coast of Africa encompasses a large
geographic area and though museum records and
recent directed sawfish literature are available,
reports are particularly rare in this region.
Historically, however, certain parts of the west coast
of Africa yielded higher catches of sawfishes,
especially Tidra (Mauritania), Casamance (Senegal),
the Bissagos Archipelago (Guinea-Bissau) and
Kamsar (Republic of Guinea), although currently
the species is largely restricted to the Bissagos
Archipelago (Guinea-Bissau) (Ballouard et al., 2006;
Robillard and Séret, 2006). Unfortunately, the
lack of size data for sawfish records prevent
speculation on any migrations that might occur in
the eastern Atlantic.
Copyright # 2013 John Wiley & Sons, Ltd.
Fishers indicated that sawfishes were once
common (‘a few decades ago’) on the west coast of
Africa, with catches of one or two specimens per net
made regularly and even up to 10 specimens
captured occasionally (Robillard and Séret, 2006).
Fisheries, human population increase along rivers
and habitat degradation have been described as the
causes of the drastic disappearance of sawfishes in
the region within the last three decades (Ballouard
et al., 2006; Robillard and Séret, 2006). Although
there have been sporadic reports of sawfish captures
in 2006 and 2008 from the islands of Orango and
Formosa in Guinea-Bissau, there are no pictures or
evidence to identify the sawfish species (J. Huet,
pers. comm., 2009). The fact that Guinea-Bissau’s
coastal populations have no strong fishing tradition
when compared with the neighbouring countries
may have, in part, helped protect the species in the
region. However, in the last two decades an
increasing number of fishermen from other nearby
countries have been attracted to the rich Bissagos
Archipelago, with most participating in the
shark fishing fleet (Campredon and Cuq, 2001).
Thus, urgent protection for sawfish is needed
for Guinea-Bissau, and in particular for the
Bissagos Archipelago.
CONCLUSIONS
Pristis pristis appears to be threatened in all of its
six distribution areas in the Atlantic. Extinction
risk was detected quantitatively for three of these
areas: the USA, northern South America and
southern West Africa. The US population is now
probably extinct, since the best scenario for
estimated date of extinction was the present
decade and the qualitative evidence supported
local extinction in this region. Regarding northern
South America and southern West Africa, the
results must be taken with caution, as even though
quantitative analyses were robust enough to
indicate extinction risks, the estimated date of
extinction encompassed a wide time-span or the
qualitative evidence for extinction was not strong.
Although P. pristis has been extirpated from
several areas, the species can still be found in
some regions of the Atlantic. The Amazon
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
estuary in Brazil appears to have the highest
remaining abundance of P. pristis in the Atlantic,
although there are indications of low presence of
the species in the Colorado River–San Juan River
system in Nicaragua and Costa Rica, and in the
Bissagos Archipelago in Guinea-Bissau. Evidence
from the literature suggests that the decline of
P. pristis in the Atlantic is mostly associated with
fisheries activities, both direct (target) and indirect
fisheries (bycatch), principally commercial
(industrial) and artisanal trawling and longline
fisheries. In addition, habitat degradation may
also have played a role in this decline, since it has
been associated as a secondary cause of depletion
of other sawfish species that occur in the
Atlantic (e.g. P. pectinata).
Conservation and management implications
The depletion of P. pristis throughout its Atlantic
range is of concern. The situation is even more
dramatic when considered in the context of the
biological limitations of the species: long life span,
late sexual maturity, and low fecundity, which in
combination result in low reproductive potential
and reduced capacity for recovery following
population depletion (Thorson, 1976, 1982a, b;
Simpfendorfer, 2000). The recovery of the related
P. pectinata in the USA is estimated to require
100 years even with zero fishing mortality and
concerted action to conserve critical habitat during
that long time period (NMFS, 2009b). Although
P. pristis shares similar biological attributes,
conservation efforts are predicted to produce an
even slower recovery for P. pristis than for P.
pectinata populations owing to its lower intrinsic
rate of increase caused by their smaller litter size
(Simpfendorfer, 2000). In addition, given that
some of the remaining refugia of P. pristis are
located in areas with currently minimal to no legal
protection and difficult enforcement, a similar
recovery time frame for this species would
probably be over-optimistic. Even though it is
very encouraging that international recognition of
the need for conservation and management
measures already exists for the species (e.g.
CITES, IUCN Red List), it is crucial that
measures at national or regional levels are created
Copyright # 2013 John Wiley & Sons, Ltd.
and that legal protection is enforced soon, if the
extinction of the species in the Atlantic Ocean is
to be prevented.
The three areas that still have potentially viable
populations of P. pristis (i.e. Amazon estuary,
Colorado River–San Juan River system and
Bissagos Archipelago) should be considered priority
areas for conservation and law enforcement. Areas
with only historical abundance records should also
be taken into consideration in any conservation
actions, as these may serve as refugia for P. pristis,
and may be vital for the potential recovery of
the species.
To our knowledge, only four countries within the
Atlantic range of P. pristis have national protection
to date. In the USA the species has been listed
under the Endangered Species Act since 2011
making it illegal to capture, harass or harm any
specimen (NMFS, 2009a). In Mexico, there is a
fishery ban since 2006 on all take of sawfish
(Fordham, 2012). In Nicaragua, a fishery ban on
targeted sawfish on Lake Nicaragua was
implemented in 2006 although it does not address
bycatch (Charvet-Almeida et al., 2007; Fordham,
2012). Finally, in Brazil, a federal law by the
Ministry of Environment (MMA-IN05/2004)
mandates no take of smalltooth and largetooth
sawfish since 2004. No national laws for countries
in the west coast of Africa were found.
Although it is positive that laws protecting
P. pristis already exist in the countries identified
in this study as the areas with the largest
remaining populations of the species (Brazil and
Nicaragua), enforcement is still a critical problem
(Charvet-Almeida et al., 2007; Melo Palmeira
et al., 2013). Although protective laws are the first
step in the conservation of a species, more
conservation measures should follow in order to
achieve the desired recovery of a slow-growing
species such as P. pristis. For example, as was
done in the USA for the smalltooth sawfish
(NMFS, 2009b), it is advised that national
recovery plans be developed and implemented for
P. pristis in order to assess how to minimize
human interactions (injury and mortality),
identify the critical habitat for the species, and
protect (or restore) those habitats to promote
considerable increase in abundance to allow the
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
species to reoccupy areas from which it has been
extirpated. Thus, research on P. pristis should be
encouraged in order to gather information crucial
for developing such recovery plans, namely
attempt to increase the available knowledge about
the basic biology of the species (reproduction,
growth rates, movement patterns, etc.), assess the
threats quantitatively (e.g. fishing effort, landing
data) and identify possible nursery areas which, if
protected, can facilitate adult recruitment. Finally,
public outreach and education would be useful tools
to aid the recovery of this endangered species.
M. G. G. Grol (Department of Animal Ecology and
Ecophysiology, Radboud University, Aruba), J.
Huet ("Poisson-Scie", Noé Conservation/FIBA,
Guinea-Bissau), D. Matos-Caraballo (DNER
Fishery Research Laboratory, Puerto Rico), M. M.
Carranza (ECOSUR, Mexico), J. Nunes (Universidade
Federal do Maranhão, Brazil), F. P. Amargós
(CIEC, Cuba), J. C. P. Jiménez (ECOSUR,
Mexico), L. Py-Daniel (INPA, Brazil), and R.
Tavares (Instituto Nacional de Investigaciones
Agrícolas, Venezuela). Finally, we would like to
thank the anonymous reviewers whose valuable
comments greatly improved this manuscript.
ACKNOWLEDGEMENTS
REFERENCES
Funding to support this project was provided by
the National Marine Fisheries Service. During part
of this study Joana Fernandez-Carvalho was
supported by a grant from FCT, the Portuguese
Foundation for Science and Technology (Ref: BD
60624/2009). We appreciate the assistance provided
by many friends and colleagues, especially D.
Bethea, S. Norton, G. Poulakis, and T. Wiley. The
hard work and patience of FPSR staff members and
volunteers C. Bester, S. Bühler, C. Campbell, F.
Carvalho, R. Coelho, and J. Zill is gratefully
acknowledged. We thank the following curators of
museum, university, and private ichthyological
collections who responded to our requests for
information on their sawfish holdings: A. Bentley,
B. Brown, D. Catania, P. Chakrabarty, P. Charvet,
M. Dressler, N. Douglas, N. Ferre, A. Harold,
K. Hartel, G. Hogue, B. Kuhajda, S. LeCroy, J.
MacLaine, M. McDavitt, R. Norton, C. Opitz,
H. Prestridge, P. Pruvost, N. Rios, M. Rogers,
J. Rosales, M. Sabaj, A. Sanderson, J. Seigel, J.
Seitz, B. Séret, and J. Williams. We thank the
following researchers for their valuable anecdotal
accounts of P. pristis in the state or country in
which they work: A. Acero (Invemar-Instituto de
Investigaciones Marinas y Costeras, Colombia),
R. Burris (MS Dept of Marine Resources),
J.L. Castillo-Geniz (INAPESCA, Mexico), A. O.
Debrot (Carmabi Foundation, Curaçao), H.
Guzmàn
(Smithsonian
Tropical
Research
Institute, Panama), F. Gomez (Departamento de
Biologia Ponteficia Universidad Javeriana, Colombia),
Acero PA, Alvarez-León R, Garzón J. 1986. Peces comerciales
del caribe colombiano, sin incluir los de las islas, cayos y
bajos del caribe occidental (propuesta para la unification de
sus nombres vernaculars). Informe Museo del Mar,
Universidad de Bogota 31: 1–15.
Almeida ZS, Carneiro MC. 1999. Levantamento e ocorrência
de elasmobrânquios capturados pela pesca artesanal no
litoral do Maranhão. Ceuma Perspectivas 3: 122–136.
Almeida ZS, Nunes JLS, Paz AC. 2006. Elasmobrânquios no
Maranhão: biologia, pesca e ocorrência. In Projeto e ações
em Biologia e Química, Costa da Silva A, Martins Bringel
JM. (eds). EDUUEMA: São Luís; 35–57.
Alvarez LR, Blanco RJ. 1985. Composition of fish
communities in the lagoon and estuarine complex of
Cartagena Bay, Cienaga de Tesca and Cienaga Grande of
Santa Marta, Colombian Caribbean. In Fish Community
Ecology in Estuaries and Coastal Lagoons: Towards an
Ecosystem Integration, Yáñez-Arancibia A (ed.).UNAM
Press: Mexico City; 535–656.
Ballouard JM, Robillard M, Yvon C. 2006. Statut et conservation
des poissons-scies et autres chondrichtyens menacés en
Afrique de l’Ouest: Rapport de synthèse. Rapport Missions
2005 et 2006, Poissons-scies. Noé Conservation – CSRP.
Bates M. 1964. The Land and Wildlife of South America. Time
Inc: New York.
Baughman JL. 1943. Notes on sawfish, Pristis perotteti Müller
and Henle, not previously reported from the waters of the
United States. Copeia 1943: 43–48.
Baughman JL. 1952. The marine fishes of Texas. The sawfishes.
Texas Game and Fish 10: 28–29.
Bigelow HB, Schroeder WC. 1953. Sawfishes, guitarfishes,
skates and rays. In Fishes of the Western North Atlantic,
Part two., Tee-Van J, Breder CM, Parr AE, Schroeder WC,
Schultz LP (eds). Memoirs of Sears Foundation for Marine
Research: Yale, University, New Haven; 1–514.
Burgman MA, Grimson RC, Ferson S. 1995. Inferring threat
from scientific collections. Conservation Biology 9: 923–928.
Bussing WA. 2002. Freshwater Fishes of Costa Rica.
Universidad de Costa Rica: San José, Costa Rica.
Butchart SHM, Stattersfield AJ, Brooks TM. 2006. Going or
gone: defining ‘possibly extinct’ species to give a true
Copyright # 2013 John Wiley & Sons, Ltd.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
STATUS OF THE LARGETOOTH SAWFISH IN THE ATLANTIC OCEAN
picture of recent extinctions. Bulletin of the British
Ornithologists’ Club 126A: 7–24.
Caldas JP; Díaz-Trujillo EM; García CB, Duarte LO. 2009.
Revisión Histórica de la Pesca de Tiburones y Rayas en el
Mar Caribe Continental de Colombia. In Avances en el
Conocimiento de Tiburones, Rayas y Quimeras de Colombia,
Puentes V, Navia AF, Mejía-Falla PA, Caldas JP,
Diazgranados MC, Zapata LA (eds). Fundación Squalus,
Ministerios de Ambiente, Vivienda y Desarrollo Territorial,
Instituto
Colombiano
Agropecuario,
Colciencias,
Conservación Internacional, WWF; 99–130.
Campredon P, Cuq F. 2001. Artisanal fishing and coastal
conservation in West Africa. Journal of Coastal
Conservation 7: 91–100.
Carlson JK, Osbourne J, Schmidt TW. 2007. Monitoring the
recovery of smalltooth sawfish, Pristis pectinata, using
standardized relative indices of abundance. Biological
Conservation 136: 195–202.
Carvalho MR, McEachran JD. 2003. Family Pristidae
(sawfishes). In Check List of the Freshwater Fishes of South
and Central America, Reis RS, Kullander SO, Ferraris SJ
(eds), Pontifícia Universidade Católica do Rio Grande do
Sul, Porto Alegre, RS: Brasil; 17–21.
Castro-Aguirre JL. 1978. Catalogo sistemático de los peces
marinos que penetran a las aguas continentales de México
con aspectos zoogeográficos y ecológicos. Dirección General
del Instituto Nacional de Pesca, Serie Científica 19: 1–296.
Castro-Aguirre JL. 1999. Ictiofauna estuarino-lagunar y vicaria de
México. Editorial Limusa S.A. de C.V.: México D.F, México.
Cervigón MF. 1966. Los Peces Marinos de Venezuela, Tomo II.
Estación de Investigaciones Marinas de Margarita,
Fundación La Salle de Ciencias Naturales: Caracas,
Venezuela.
Cervigón MF. 1993. Los peces marinos de Venezuela, Vol 2,
2nd edn. Edit. Fundación Científica Los Roques: Caracas.
Charvet-Almeida P. 1999. General information from sawfishes
in Pará state, north Brazil. Boletim SBEEL 4: 12.
Charvet-Almeida P. 2002. Sawfish trade in the north of Brazil.
Shark News 14: 9.
Charvet-Almeida P, Faria V. 2008. Pristis perotteti. In Livro
Vermelho da Fauna Brasileira Ameaçada de Extinção (Série
Biodiversidade), Machado ABM, Drummond GM, Paglia
AP (eds). Belo Horizonte: Fundação Biodiversitas II; 33–35.
Charvet-Almeida P, Faria V, Furtado M, Cook SF, Compagno
LJV, Oetinger MI. 2007. Pristis perotteti. In IUCN 2010.
IUCN Red List of Threatened Species. Version 2010.4.
(Available from: http//www.iucnredlist.org).
Compagno LJV, Cook SF. 1995. The exploitation and
conservation of freshwater elasmobranches: status of taxa
and prospects for the future. Journal of Aquariculture and
Aquatic Sciences 7: 62–90.
Dahl G. 1971. Los Peces del Norte de Colombia. Ministerio de
Agricultura, Instituto de Desarrollo de los Recursos
Naturales Renovables: Bogota, Colombia.
Dulvy NK, Sadovy Y, Reynolds JD. 2003. Extinction
vulnerability in marine populations. Fish and Fisheries 4: 25–64.
Faria VV, McDavitt MT, Charvet P, Wiley TR, Simpfendorfer CA,
Naylor GJP. 2013. Species delineation and global population
structure of critically endangered sawfishes (Pristidae).
Zoological Journal of the Linnean Society 167: 136–164.
Flórez-González L. 1986. Lista preliminar de las especies ícticas
marinas para el Caribe colombiano. Informe Museo del Mar
32: 1–101.
Copyright # 2013 John Wiley & Sons, Ltd.
Fordham S. 2012. Conservation status of sawfishes. Global
Sawfish Conservation Strategy Workshop, London, May 2012.
Fowler HW. 1910. Notes on batoid fishes. Proceedings of the
Academy of Natural Sciences of Philadelphia 62: 468–475.
Fowler HW. 1919. Notes on tropical American fishes.
Proceedings of the Academy of Natural Sciences of
Philadelphia 71: 128–155.
Fowler HW. 1936. The marine fishes of West Africa: based on
the collection of the American Museum Congo Expedition,
1909–1915. Bulletin of the American Museum of Natural
History 70: 1–606.
Fowler SL, Cavanagh RD, Camhi M, Burgess GH, Cailliet
GM, Fordham S, Simpfendorfer CA, Musick JA. 2005.
Sharks, Rays and Chimaeras: The Status of the
Chondrichthyan Fishes. IUCN/SSC Shark Specialist Group,
IUCN: Gland, Switzerland.
Furneau R. 1969. The Amazon. G. P. Putnam’s and Sons:
New York.
Gadig OBF. 1998. Peixes cartilaginosos da costa do estado de
São Paulo. Revista Ceciliana 8: 41–51.
Gill T, Bransford JF. 1877. Synopsis of the fishes of Lake
Nicaragua. Proceedings of the Academy of Natural Sciences
of Philadelphia 1877: 175–191.
Grijalba-Bendeck M, Acero A, Díaz-Trujillo EM, Gómez F.
2009. Estado del conocimiento de los peces cartilaginosos
del Caribe continental de Colombia. In Avances en el
Conocimiento de Tiburones, Rayas y Quimeras de Colombia,
Puentes V, Navia AF, Mejía-Falla PA, Caldas JP,
Diazgranados MC, Zapata LA (eds). Fundación Squalus,
Ministerios de Ambiente, Vivienda y Desarrollo Territorial,
Instituto Colombiano Agropecuario, Colciencias, Conservación
Internacional, WWF; 41–96.
Grogan CS, Boreman J. 1998. Estimating the probability that
historical populations of fish species are extirpated. North
American Journal of Fisheries Management 18: 522–529.
Hoover JJ. 2008. Searching for sawfish: a history of the hunt.
American Currents 34: 1–15.
Kyne PM, Carlson J, Smith K. 2013. Pristis pristis In: IUCN
2013. IUCN Red List of Threatened Species. Version 2013.1.
www.iucnredlist.org. Downloaded on 29 August 2013.
Longley WL. 1994. Freshwater inflows to Texas bays and
estuaries: ecological relationships and methods for
determination of needs. Texas Water Development Board
and Texas Parks and Wildlife Department: Austin, TX.
Luiz OJ, Edwards AJ. 2011. Extinction of a shark population in
the Archipelago of Saint Paul’s rocks (equatorial Atlantic)
inferred from the historical record. Biological Conservation
144: 2873–2881.
Marden L. 1944. A land of lakes and volcanoes. National
Geographic 86: 161–192.
Marlier G. 1967. Hydrobiology in the Amazon region. In Atas do
Simpósio sobre a Biota Amazónica Vol. 3: Limnologia, Lent H
(ed.). Conselho Nacional de Pesquisas: Rio de Janeiro.
McDavitt MT. 2002. Lake Nicaragua revisited: conversations
with a former sawfish fisherman. Shark News 14: 5.
McDavitt M, Charvet-Almeida P. 2004. Quantifying trade in
sawfish rostra: two examples. Shark News 16: 10–11.
McInerny GJ, Roberts DL, Davy AJ, Cribb PJ. 2006.
Significance of sighting rate in inferring extinction and
threat. Conservation Biology 20: 562–567.
Meek SE. 1907. Synopsis of the fishes of the great lakes of
Nicaragua. Field Columbia Museum Publication 121, Zoology
Series 7: 97–132.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)
J. FERNANDEZ-CARVALHO ET AL.
Mejía LS, Acero AP. 2002. Libro rojo de peces marinos de
Colombia. INVEMAR, Universidad Nacional de Colombia:
Bogota.
Melo Palmeira CA, da Silva Rodrigues-Filho LF, de Luna
Sales JB, Vallinoto M, Schneider H, Sampaio I. 2013.
Commercialization of a critically endangered species
(largetooth sawfish, Pristis perotteti) in fish markets of
northern Brazil: authenticity by DNA analysis. Food
Control 34: 249–252.
Miles C. 1945. Some newly recorded fishes from the Magdalena
River system. Caldasia 3: 453–464.
Miles C. 1947. Los peces del río Magdalena. Ministerio de la
Economía Nacional, Sección de Piscicultura, Pesca y Caza,
Bogotá.
Moncada MJ. 2006. Prohíben pesca de tiburón y pez sierra. La
Prensa (17 January 2006).
Monte-Luna P, Lluch-Belda D, Serviere-Zaragoza E, Carmona
R, Reyes-Bonilla H, Aurioles-Gamboa D, Castro-Aguirre
JL, del Proo SAG, Trujillo-Millan O, Brook BW. 2007.
Marine extinctions revisited. Fish and Fisheries 8: 107–122.
Monte-Luna P, Castro-Aguirre JL, Brook BW, Cruz-Aguero J,
Cruz-Escalona VH. 2009. Putative extinction of two sawfish
species in Mexico and the United States. Sociedade
Brasiliera de Ictiologia: Neotropical Ichthyology 7: 59–512.
Montoya RV, Thorson TB. 1982. The bull shark (Carcharhinus
leucas) and largetooth sawfish (Pristis perotteti) in Lake
Bayano, a tropical man-made impoundment in Panama.
Environmental Biology of Fishes 7: 341–347.
NMFS (National Marine Fisheries Service). 2009a. Listing
Endangered and Threatened Species and Designating
Critical Habitat: Notice of Finding on a Petition To List
the Largetooth Sawfish (Pristis perotteti) as an Endangered
or Threatened Species Under the Endangered Species Act.
Federal Register 74(144): 37671–37674.
NMFS (National Marine Fisheries Service). 2009b. Recovery
Plan for Smalltooth Sawfish (Pristis pectinata). Prepared
by the Smalltooth Sawfish Recovery Team for the
National Marine Fisheries Service. Silver Spring:
Maryland.
Nunes JLS, Almeida ZS, Piorski NM. 2005. Raias capturadas
pela pesca artesanal em águas rasas do Maranhão - Brasil.
Arquivos de Ciências do Mar, Fortaleza 38: 49–54.
Poulakis GR, Seitz JC. 2004. Recent occurrence of the smalltooth
sawfish, Pristis pectinata (Elasmobranchiomorphi: Pristidae),
in Florida Bay and the Florida Keys, with comments on
sawfish ecology. Florida Scientist 67: 227–235.
Puyo J. 1949. Poissons de la Guyane Française. XII: Faune de
l’Empire Français. Office de la Recherche Scientific Outre-Mer,
Librairie Larose: Paris, France.
Ramjohn DD. 1999. Check list of coastal and marine fishes
of Trinidad and Tobago. Marine Fishery Analysis
Unit, Fisheries Division, Ministry of Agriculture, Land
and Marine Resources, Trinidad and Tobago, Fisheries
Information Series 8.
Copyright # 2013 John Wiley & Sons, Ltd.
Rivadeneira MM, Hunt G, Roy K. 2009. The use of sighting
records to infer species extinctions: an evaluation of
different methods. Ecology 90: 1291–1300.
Roberts DL. 2006. Extinct or possibly extinct? Science
312: 997–998.
Roberts DL, Solow AR. 2003. Flightless birds - when did the
dodo become extinct? Nature 426: 245–245.
Robillard M, Séret B. 2006. Cultural importance and decline of
sawfish (Pristidae) populations in West Africa. Cybium
30: 23–30.
Santos GM, Val AL. 1998. Peixe-serra: o falso tubarão da
Amazônia. Ciência Hoje 23: 66–67.
Saunders GB, Holloway AD, Handley CO. 1950. A fish and
wildlife survey of Guatemala. US Fish and Wildlife Service,
Special Scientific Report 5: 1–162.
Simpfendorfer CA. 2000. Predicting population recovery rates
for endangered western Atlantic sawfishes using demographic
analysis. Environmental Biology of Fishes 58: 371–377.
Solow AR. 1993a. Inferring extinction from sighting data.
Ecology 74: 962–964.
Solow AR. 1993b. Inferring extinction in a declining
population. Journal of Mathematical Biology 32: 79–82.
Solow AR. 2005. Inferring extinction from a sighting record.
Mathematical Biosciences 195: 47–55.
Solow AR, Roberts DL. 2003. A nonparametric test for
extinction based on a sighting record. Ecology 84: 1329–1332.
Springer VG, Woodburn KD. 1960. An ecological study of the
fishes of the Tampa Bay area. Florida State Board of
Conservation, Professional Paper Series 1: 1–104.
Strauss D, Sadler PM. 1989. Classical confidence intervals and
Bayesian probability estimates for ends of local taxon ranges.
Mathematical Geology 21: 411–421.
TDWR (Texas Department of Water Resources). 1982. The
influence of freshwater inflows upon the major bays and
estuaries of the Texas Gulf Coast: Executive Summary.
Texas Department of Water Resources: Austin, TX.
Thorson TB. 1974. Occurrence of the sawfish, Pristis perotteti,
in the Amazon River, with notes on P. pectinatus. Copeia
1974: 560–564.
Thorson TB. 1976. Observations on the reproduction of the sawfish,
Pristis perotteti, in Lake Nicaragua, with recommendations for its
conservation. In Investigations of the Ichthyofauna of Nicaraguan
Lakes, Thorson TB (ed.), University of Nebraska, School of Life
Sciences: Lincoln, NE; 641–650.
Thorson TB. 1982a. The impact of commercial exploitation on
sawfish and shark populations in Lake Nicaragua. Fisheries
7: 2–10.
Thorson TB. 1982b. Life history implications of a tagging study
of the largetooth sawfish, Pristis perotteti, in the Lake
Nicaragua-Río San Juan system. Environmental Biology of
Fishes 7: 207–228.
Zorzi GD. 1995. The biology of freshwater elasmobranchs: an
historical perspective. Journal of Aquariculture & Aquatic
Sciences 7: 10–31.
Aquatic Conserv: Mar. Freshw. Ecosyst. (2013)