







































Abstract: Smalltooth saw!sh (Pristis pectinata) are a marine species of high concern since they are listed as critically 
endangered species by the International Union for Conservation of Nature (IUCN) and listed as endangered under 
the U.S. Endangered Species Act (ESA). Smalltooth saw!sh are extremely vulnerable to being caught as bycatch 
due to their long, toothed rostrum that can easily become entangled in !shing nets, especially in shrimp trawling 
!sheries. Several studies have been conducted on smalltooth saw!sh since being listed under the ESA in 2003, 
but their vertical movements and use of depth have still not been thoroughly studied. Until recently, saw!sh were 
thought to stay at a depth of 10 m or less, but studies have now shown that they occupy much  deeper depths. Due 
to these con"icting !ndings in the literature, depth was chosen to be the focus of this study. Furthermore, this study 
investigates whether sex or individual total length has an in"uence on the percentage of time a saw!sh spends at 
a particular depth. Satellite telemetry was used to track the movements of the tagged saw!sh (n=14). #e data 
collected from the pop-o$ archiving satellite tags  (PSATs) showed smalltooth saw!sh spend most of their time in 
shallow water (between 0-8 m), but do frequently occupy deeper depths. Sex was found to be a signi!cant factor, 
while total length was not. Females were found to spend more time at deeper depths than males. Understanding 
how saw!sh use depth is important in order to predict population trends and dynamics. Knowing the factors that 
a$ect a saw!sh’s depth use would be bene!cial to e$orts that manage and conserve this species. With the decline of 
smalltooth saw!sh and their vulnerability to population loss, more research needs to be conducted to create more 
e$ective management and recovery plans.

Aisthesis      Volume 11,  20201

Sex and Length as Predictors of Vertical Movements in 
Smalltooth Saw!sh (Pristis pectinata)

by Taylor Mogavero

Introduction
 Saw!sh are large rays in the Chondrichthyes 
class and are known for their long, toothed “saw” or 
rostrum (Harrison & Dulvy, 2014). #eir rostrum 
is an elongated snout with horizontal rostral teeth 
that is used for feeding and defense (Poulakis & 
Seitz, 2004; Whitty et al., 2009). About 20-28% of a 
saw!sh’s body length comes from its rostrum, and it 
serves a vital role for capturing and detecting prey 
(Harrison & Dulvy 2014). #e rostrum is able to 
!nd and catch prey due to extensive sensory organs 
that detect minute electrical signals emitted by other 
animals. A saw!sh can also use its rostrum to stunt 
or kill !sh by slashing its toothed side into its prey 
(Harrison & Dulvy, 2014). #e species live in coastal 
tropical and subtropical waters in both estuaries and 
freshwater. #ey can be found across the world, but 
the only saw!sh species currently found in the U.S. is 
the smalltooth saw!sh (Pristis pectinata). 
 #e average size at birth for a smalltooth saw!sh 
is 80 cm total length (TL). #e size at maturity is 370 
cm TL for females and 340 cm TL for males (Brame 

et al., 2019). #eir age of maturity is estimated to 
be 7-11 years for males and females (Carlson & 
Simpfendorfer, 2015), with a total lifespan around 
30 years. Smalltooth saw!sh are yolk-sac viviparous, 
meaning young are nourished in utero by an external 
yolk sac and are then born live. It is presumed that 
their reproduction occurs biennially and their average 
litter size is 7-14 pups (Feldheim et al., 2017). Due to 
their long lifespans and late maturity, saw!sh have 
a slow population growth rate, therefore increasing 
their vulnerability to and di%culty recovering from 
population loss. 
 Many of the areas where saw!sh reside are 
highly threatened habitats such as mangroves or 
seagrasses, two habitats that have seen a great decline 
in range over the years. #eir coastal preference also 
tends to overlap with large cities and areas of high 
human population density where more activities 
like !shing occur (Dulvy et al., 2016). #e current 
geographical distribution of P. pectinata is mainly in 
the western Atlantic, but they have also been found 
in the eastern Atlantic (Harrison et al., 2014), and 



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20202

the largest population is found along the southwest 
coast of Florida (Poulakis & Seitz, 2004). Smalltooth 
saw!sh were historically found along the coast of 
the United States as far north as the Carolinas, in 
the Gulf of Mexico, the Caribbean Sea, and along 
the coast as far south as Uruguay (Carlson et al., 
2014; Dulvy et al., 2016; Feldheim et al., 2017; 
Simpfendorfer, 2001; Wiley & Simpfendorfer, 2010). 
#e smalltooth saw!sh is currently found in less 
than 20% of its historic range (Dulvy et al., 2016). 
Now smalltooth saw!sh are only consistently found 
in the coastal waters of southern Florida with a 
slow increase or stable population growth, and an 
estimated population size of only a few thousand 
(Carlson et al., 2014; Feldheim et al., 2017; Wiley & 
Simpfendorfer, 2010). Exact population reduction 
rates are hard to calculate due to limited scienti!c 
data, but it has been estimated that the population 
may have declined as much as 95% from the historic 
stock size (Wiley & Simpfendorfer, 2010). 
 Smalltooth saw!sh tend to stay near or within 
mangroves and seagrass beds as juveniles (Dulvy et 
al., 2016; Wiley & Simpfendorfer, 2010). Saw!sh are 
known to occupy shallow coastal waters typically 10 
m deep or less (Carlson et al., 2014), but they can 
occupy depths deeper than 10 m and can be found 
at depths up to 122 m (Poulakis & Seitz, 2004). It 
has been postulated that smaller and immature 
saw!sh are more commonly found in shallow water 
(Poulakis & Seitz, 2004; Wiley & Simpfendorfer, 
2010; Whitty et al., 2009; Simpfendorfer, 2001). 
#ey also prefer warm water temperatures, mainly 
22–28°C, and their lower thermal tolerance is 
predicted to be around 20°C (Carlson et al., 2014). 
Smalltooth saw!sh o&en feed on schooling !sh such 
as clupeids, carangids, mugilids, elopids, sparids, 
and belonids, as well as dasyatids stingrays (Poulakis 
et al., 2017). #ey obtain their prey by slashing their 
rostrum sideways through a school and impaling the 
!sh on their rostral teeth. A&er caught, they ingest 
their prey whole. Most of their prey are coastal or 
estuarine species that are typically found at or near 
the surface.
 #ere has been evidence for sexual segregation in 
elasmobranchs and saw!sh may be included among 
those. Sexual segregation is the separation of males 
and females of the same species; this separation 
may be spatial as well as temporal in nature, for 
example, occurring only during the non-breeding 

season (Wearmouth & Sims, 2010). It is important 
to understand the habitat use of di$erent sexes in 
order to predict population trends and dynamics. 
#is would provide data that may be useful to the 
successful management and conservation of this 
species, as these spatial dynamics o&en overlap with 
area-focused human activities like !shing. 
 #ere has never been a large-scale !shery that 
directly targeted smalltooth saw!sh, but it is very 
common for a saw!sh to get entangled in !shing nets 
due to their long-toothed rostrum, and therefore 
they are o&en caught as bycatch. #e main threat 
responsible for the decline in smalltooth saw!sh 
has been and remains commercial and recreational 
!sheries. Shrimp trawl !sheries present a major 
concern due to the high bycatch mortality of large, 
mature saw!sh, which could reduce the population’s 
reproductive potential. Fortunately, saw!sh are 
expected to su$er less and recover quicker when 
caught and released by longlines as opposed to trawls 
and gill nets (Brame et al., 2019). 
 Smalltooth saw!sh are said to be one of the 
world’s most vulnerable marine !shes (Dulvy et 
al., 2016; Feldheim et al., 2017). #e United States 
population of smalltooth saw!sh was listed as 
endangered under the U.S. Endangered Species Act 
in April 2003 (Poulakis & Seitz, 2004). Penalties 
such as imprisonment or steep !nes could be given 
to anyone who harasses, harms, or kills any animal 
listed on the U.S. Endangered Species Act. Smalltooth 
saw!sh are classi!ed as critically endangered by the 
International Union for Conservation of Nature 
(IUCN). #is is the highest level of alert—the closest 
to extinction—set by the IUCN and is de!ned by 
them as “a species facing an extremely high risk of 
extinction in the wild.” #e IUCN Red List states 
their population trend is still decreasing (Carlson et 
al., 2013). 
 Pop-o$ archiving satellite tags (PSATs) are a 
relatively new electronic tagging technology. PSATs 
detach from the tagged animal a&er a predetermined 
time, "oat to the surface, and transmit the archived data 
to satellites which provide the data to the researcher. 
#is technology allows tracking the movements of 
pelagic !sh in their natural environment to be much 
more accessible and economical. PSATs can sample 
temperature, depth, and light levels at user-de!ned 
time intervals, and then store and process these data 
(Luo et al., 2006). It has become more common to 
use PSATs to gather data on horizontal and vertical 



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20203

movements of pelagic !shes, especially since the 
vertical movement of PSAT-tagged !shes in sea 
water has a high degree of accuracy and precision 
(depth and temperature resolution are claimed to be 
0.5 m and 0.05°C, respectively) (Luo et al., 2006).

Why we need to study and protect saw!sh
 #e U.S. Endangered Species Act requires by 
law that critical habitat must be designated for any 
listed species. To date, critical habitat has only been 
designated for small juvenile smalltooth saw!sh. 
Satellite tagging of saw!sh adults was done for 
the following reasons: 1) to aid in de!ning critical 
habitat for adults and vertical space use; 2) to 
determine potential aggregation sites for mating 
and areas where males and females overlap in depth; 
and 3) to determine if the U.S. population is distinct 
from adjacent populations (e.g. Bahamas, Cuba) by 
determining if saw!sh frequently traverse deep water 
(up to 800 m depths). With the decline of saw!sh, 
more research needs to be conducted in order to 
create more e$ective status assessments, management 
measures, and recovery plans (Dulvy et al., 2016). 
Fishery management, where it does occur, focuses 
mostly on commercially valuable !sh populations, 
so populations like saw!sh are rarely the main 
concern. It is di%cult to develop recovery strategies 
for species that do not have a su%cient amount of 
scienti!c data concerning their distribution and 
habitat use, especially if they are widely dispersed 
(Wiley & Simpfendorfer, 2010). More research 
must be conducted on saw!sh so adequate recovery 
plans can be developed. #e purpose of this study 
is to assess whether sex or length has an in"uence 
on the percent time a smalltooth saw!sh spends at 
a particular depth. Conducting research on how 
saw!sh use depth could help predict population 
trends and dynamics. Knowing the factors that a$ect 
a saw!sh’s depth would allow for the successful 
management and conservation of this species, so 
it is important that these factors are studied and 
considered. 
      
Methods
 Pop-o$ archival satellite tagging data was 
collected from 2011 to 2017 and forty-seven 
saw!sh were tagged. #e saw!sh were caught using 
a research vessel and a bottom longline of 50 or 
100 16/0 hooks baited with lady!sh, Elops saurus. 
Longline stations were selected based on being 

potential habitats suitable for smalltooth saw!sh and 
historic encounter records. Tagging was conducted 
in the Florida Bay and the Florida Keys. Most 
of the sites had coordinates around 25°N 81°W 
(Figure 1). Satellite telemetry was used to track the 
movements of the tagged saw!sh. Pop-o$ archiving 
satellite tags (PSATs), which record depth (m), 
temperature (°Celsius), time, and light levels, were 
attached externally to the saw!sh !rst dorsal !n. 
#ree di$erent models of PSATs (all manufactured 
by Wildlife Computers, Inc.) were used: Mini-PAT, 
MK10, and PATF.  #ree di$erent models of PSATs 
were used because multiple research institutions 
contributed to tagging the saw!sh. #e tags were 
programmed to detach from the animals a&er a 
certain number of days (depending on the tag type) 
and transmit the archived data to a satellite. #e 
PSATs use light-based geolocation where the spatial 
track is based on the timing of local noon (used to 
estimate longitude) and day length (used to estimate 
latitude) and corrected for temperature (Luo et al., 
2006). Depth data were collected every four hours. 
Each satellite tag or PTT (Platform Transmitting 
Terminal) was assigned a PTT number.
 For this study, data was available for fourteen 
satellite tags (Wildlife Computers, Inc.), spanning a 
deployment period from March 2011 to March 2017. 
Six of the tags were MK10 tags (programmed for 150 
days), !ve were PATF tags (tracked for 60 days), and 
three were Mini-PAT tags (programmed for 105 
days). 

Choosing viable data
 #e GPE2 program from Wildlife Computers 
was opened using iGOR Pro 6.37 so&ware. #e 
graphs that displayed viable data (daily depths 
measurements for at least a duration of two weeks) 
were chosen to examine further. Further examination 
was done by looking at the speci!c depth measures 
in the Excel sheets for each of those tags. Only 
depths with an error of 4 m or less were considered, 
since that was the most accurate reading recorded by 
any tag. Anything with a higher error than 4 m was 
considered to be inaccurate.

Used data
 In total, forty-three satellite tags were inspected 
for data use. Only fourteen of those tags were analyzed 
due to some tags failing to report or record enough 
data to make it viable. Eight of these were males and 



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20204

six were females. Statistics for each analyzed saw!sh 
can be seen in Table 1. Any negative minimum 
depths were regarded as 0 m (at the surface). 

Creating histograms of percentage time-at-depth
 All of the recorded depth measurements, with an 
error of 4 m or less, for an individual saw!sh were 
considered. Depth bins were created in 8 m intervals 
(0-8, 9-16, 17-24, etc.), since the recorded data were 
expressed in increments of 8 m (0, 8, 16, 24, etc.). #e 
percent of time each saw!sh spent in each depth bin 
was calculated and made into a histogram in Excel. 

Average maximum depth
 Excel was used to calculate the average maximum 
depth of each saw!sh. #e average maximum depth 
of each day was found, then all of these calculations 
were averaged together. 

Model choice 
 Akaike’s Information Criteria (AIC) is a log-
likelihood that penalizes any super"uous parameters 
in the model. AIC estimates the quality of each 
entered model, relative to each of the other models. 
AICc has small-sample correction.  A low AIC value 
indicates the model is a good !t; the model with the 
lowest AIC is considered the best !t. If the delta AIC 
is lower than 2, it can be said to have substantial 
support. Various linear mixed-e$ects models were 
tested. #e entered model that was the simplest was 
chosen by comparing AICc values to determine 
which model has a better !t. Due to the principle 
of parsimony, all the factors that did not cause a 
signi!cant increase in deviance were removed from 
the model to form a minimal adequate model. 

Linear mixed-e"ects model 
 R Statistical Computing (R Core Team, 2019) 
and lme4 (Bates, Maechler, Bolker & Walker, 2015) 
were used to perform a linear mixed e$ects analysis 
of the relationship between percent time and 
depth. A mixed-e$ects model was chosen because 
the data had both random and !xed e$ects due to 
temporal pseudoreplication resulting from repeated 
measurements on the same individuals. As the 
random e$ect, the saw!sh PTT number was entered 
(without interaction term) into the model. Intercepts 
for depth and sex were !xed e$ects, as well as by-
subject and by-item random slopes for the e$ect of 

depth. Visual inspection of residual plots did not 
reveal any obvious deviations from homoscedasticity 
or normality for sex. #e function “lmer()” was used, 
as it allows for non-normal errors and non-constant 
variance with the same error as a generalized linear 
model. 

Results
Histograms of percentage time-at-depth
 #e results of the histograms can be seen in 
Figure 2. #ese histograms showed the percentage of 
time a saw!sh spent at a speci!c depth by analyzing 
all of the data recorded on that saw!sh’s tag. Every 
saw!sh was seen at a depth of 0-8 m for the majority 
of the time; however, a few saw!sh (14%) were 
recorded to go as deep as 88 m. Most tagged (n = 12) 
saw!sh spent over 60% of their time at 0-8 m, but 
two saw!sh (one male and one female) spent over 
50% of their recorded time at depths deeper than the 
0-8 m bin. As depth increases, there is no pattern of 
depth use. When a saw!sh is found at a depth deeper 
than 8 m, there is not a favored depth, and their 
preference for a depth did not consistently decrease 
with increasing depth. Sex-dependent trends were 
apparent, however. All of the females spent time in 
at least six other depth bins, while half of the males 
solely spent time in 0-8 m. 

Which sex has a greater maximum depth?
 #is t-test compared the average maximum 
depth each saw!sh occupied each day, which shows 
the deepest depth a saw!sh occupies on average. 
#e average maximum depth occupied by males 
was 5.736 m (standard error = 2.475) and 22.61 m 
(standard error = 5.118) for females. An unpaired 
student t-test was performed and the two-tailed 
p-value equaled 0.0068, suggesting there was a 
signi!cant di$erence between the means (t = 3.3985; 
df = 10 ; standard error of di$erence = 4.965). 

Model choice 
 #e lowest AICc value indicated the most 
parsimonious model, relative to the other model !ts 
with a higher AICc value (best model: lmer(Percent.
Time ~ Depth + Sex + (1|Saw!sh)). #e best model 
to explain the in"uence of percent of time at depth 
indicated sex was relevant, while length was not. #e 
minimal adequate model showed a common slope 
for percent time against depth with two intercepts, 



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20205

one for each sex. #e delta AICc of the best linear 
mixed-e$ects model was between 0-2 and therefore 
can be said to have substantial support. 

Linear mixed-e"ects model 
 #e variables that were tested were depth, length, 
and sex to address the following question: Does total 
length and/or sex have an e$ect on the percent-of-
time a saw!sh spent at a particular depth? Since 
length fell out of the model, it is not considered a 
signi!cant factor. #e percent time each sex stays at 
a speci!c depth is shown in Figure 3. In this !gure, 
it is clear that saw!sh spend more time at shallow 
depths, but de!nitely spend a considerable amount 
of time in deeper water. Due to the model choice 
conducted with AICc, both the male and female 
isoclines were given the same slope. Males spend a 
much greater time at shallower depths than females. 
#is is consistent with the average maximum depth 
t-test results that show females have a signi!cantly 
deeper average depth than males. #is shows that 
females are expected to be found at deeper depths 
than males. 

Discussion
      #e goal of this study was to assess whether sex 
or length in"uences the percent of time a smalltooth 
saw!sh spends at a speci!c depth. Knowing what 
factors determine the depth of saw!sh will help 
further understand the habits of saw!sh and 
could help with conservation management for this 
critically endangered species. #e results concluded 
that both male and female saw!sh spend more time 
in shallow depths over deeper depths, as expected 
from previous saw!sh literature (Carlson et al., 2014; 
Poulakis & Seitz, 2004; Simpfendorfer, 2001; Wiley & 
Simpfendorfer, 2010). #is knowledge is important 
to comprehend in order to understand their 
habitats and apply them to conservation practices. 
Expecting shallower depths to be favored, the main 
focus of this study was to see which sex spent more 
time occupying deeper depths. #e results showed 
females tend to spend more time at deeper depths 
than males, and males spend more time at shallower 
depths than females. Two males did go relatively deep 
(65-72 m), and two females only occupied shallow 
water (0-8 m), but the general trend showed females 
spending more time at deeper depths than males. 
#is was seen in the histograms of percentage time-

at-depth (Figure 2), the t-test conducted for greatest 
maximum depth, and the linear mixed-e$ects model 
(Figure 3). 
 Trends in the histograms (Figure 2) reveal that 
despite sex di$erences, all saw!sh analyzed spent the 
majority of their time at 0-8 m. #erefore, this area 
of the water column should still be the main concern 
for conservation and management practices. 
However, it is important to know that saw!sh do 
also frequently occupy deeper depths, and deeper 
waters still need to be considered for management 
measures to support population increases of this 
critically endangered species. 
 #ese results mean we can expect to !nd females 
at greater depths than males and could potentially 
a$ect the way female, especially pregnant, saw!sh 
are protected. Females are very important for 
population dynamics since they give birth to pups. 
In order to ensure the saw!sh population grows, 
females must be protected. If females have di$erent 
depth preferences than males, this could change the 
way the conservation of female saw!sh is managed. It 
may be more important to ensure female survival, so 
this information can help tailor saw!sh conservation 
more towards females. Further research on whether 
sexual maturity or reproductive state a$ects the 
depth a saw!sh occupies would add to scienti!c 
knowledge and management, helping to ensure 
these individuals are protected, therefore promoting 
the growth of the saw!sh population.
 Improvements to this model could include 
adding factors such as seasonality. Month was 
considered when sorting the data, but seasonality 
was not analyzed. Upon visual inspection of the data, 
it appears saw!sh mostly occupied deeper depths 
during the late summer months. #e deepening of 
the thermocline during the summer months may be 
a reason for saw!sh to go deeper. Further analysis 
regarding month and the breeding season would 
need to be calculated to see if sex is only segregating 
at certain times of the year. Di$erences in depth 
between males and females could also be due to 
the reproductive cycle of females (Carlson et al., 
2014). #is could be counterintuitive since females 
are more o&en found at nurseries, which occur in 
shallow water areas like estuaries, and would not 
support the data showing females spend more time 
at deeper depths than males (Feldheim et al., 2017). 
However, this could potentially be evidence of sexual 



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20206

segregation, where females are actively avoiding 
males during non-reproductive seasons. Further 
studies that track the exact seasons during which 
female saw!sh mate would have to be conducted in 
order to see if this could be a factor. 
 Other factors that may a$ect the distribution 
of sexes could be due to feeding behavior or 
thermoregulation (Carlson et al., 2014). #ere has 
been evidence of sex-speci!c dietary requirements, 
such as females eating more than males of the same 
size. Sex-speci!c temperature preferences may also 
occur if a female is pregnant, as warm waters can 
help an embryo develop (Wearmouth & Sims, 2010). 
Feeding behavior and temperature preference could 
be di$erent among sexes, but more data collection 
and analysis would need to be done to see if these 
factors have an e$ect on percent of time at a particular 
depth. Depth could also depend on the time of day. 
Saw!sh have been recorded to move into shallower 
waters at night and deeper waters during the day, 
so diurnal movements could be a factor (Carlson et 
al., 2014). Total length was not a signi!cant factor 
when considering what depth a saw!sh occupies, 
but modeling length and depth could still be useful 
to see if any visual trends are spotted, especially if 
this work was extended to include juveniles as well as 
adult saw!sh. It’s important to note that the analyzed 
saw!sh in this study did not have much size variety 
(ranging from 279-428 cm); therefore, length may 
be a signi!cant factor, but the study may not have 
had enough statistical power to show any variation. 
Further testing with saw!sh of greater size variety 
could be done to see if increasing the statistical power 
of length creates a di$erence. Overall, there are 
several potentially confounding variables that could 
have in"uenced the results of this study. Additional 
data and analyses would need to be collected and 
completed in order to see if these variables cause a 
signi!cant di$erence. 
 Understanding how saw!sh use depth is 
important in order to predict population trends and 
dynamics. Knowing the factors that a$ect a saw!sh’s 
depth preferences would allow for the successful 
management and conservation of this species, since 
these spatial dynamics o&en overlap with area-
focused human activities like !shing. Until recently, 
saw!sh were thought to stay at a depth of 10 m or 
less (Simpfendorfer, 2001), so data on their depth use 
is limited, but important to know in order to fully 

understand their full vertical range. Understanding 
their depth range could help with the conservation 
and recovery e$orts of this critically endangered 
species. With the decline of saw!sh, more research 
needs to be conducted in order to create more 
e$ective status assessments, management measures, 
and recovery plans (Dulvy et al., 2016). With the 
knowledge from this study, hopefully more e$ective 
recovery plans can be developed and key areas 
of research that still need to be conducted can be 
identi!ed. 

Acknowledgements 
 I want to thank Jasmin Graham, Dr. Dean 
Grubbs, Dr. Janie Wul$ and Dr. Ian MacDonald for 
their guidance and assistance throughout my entire 
thesis project. 

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179-191.

7



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  2020

Figures and Tables

8



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  20209



Sex and length as predictors of vertical movements in smalltooth saw!sh (Pristis pectinata)

Aisthesis      Volume 11,  202010


