Report - Distributions and Movements of Atlantic Shark Species

Distributions and Movements of Atlantic Shark Species:
A 52-Year Retrospective Atlas of Mark and Recapture Data
NANCY E. KOHLER and PATRICIA A. TURNER

Introduction

doi: https://doi.org/10.7755/MFR.81.2.1

pants fishing for sharks with a variety
of gear types. Detailed summary information on the history and methods of
the CSTP has been published previously (Casey, 1985; Casey and Kohler,
1992; Kohler et al., 1998; Kohler and
Turner, 2001) and excerpts from those
reports are included here.
A mark (also defined as tag)/recapture (M/R) approach to studying
sharks is an applied science used by
fishery managers to support sustainable management of shark populations
(Rounsefell and Everhart, 1953; Harden Jones, 1968; Kohler and Turner,
2001; Speed et al., 2010). This requires
knowledge of stock boundaries and
spatial ecology within the geographic
range of the species in question (Sims,
2010; Queiroz et al., 2016). Tagging
data can be used to calculate movement vectors that allow spatially explicit population models (Vandeperre
et al., 2014) to delineate life-stage-specific habitats such as pupping or feeding areas. For example, many sharks
demonstrate sex-specific dispersal or
migratory patterns (Vandeperre et al.,
2014; Secor, 2015) or increasing range

ABSTRACT—The National Marine Fisheries Service (NMFS) Cooperative Shark
Tagging Program (CSTP) was initiated in
1962 as a collaborative effort between recreational anglers, the commercial fishing
industry, and the NMFS. The CSTP data
describe the geographic range, minimum
estimates of longevity, and movements of
coastal and pelagic sharks in the Atlantic
Ocean using conventional mark/recapture
methods. This document summarizes information collected by the CSTP for a 52-year
period through 2013, updating a previous
1998 publication. A total of 229,810 sharks
of 35 species were tagged, and 13,419
sharks of 31 species were recaptured during this period. To characterize the move-

ments and distribution patterns, these data
were summarized by sex for times at liberty and distance traveled. The longest time
at liberty for any individual shark was 27.8
years (sandbar shark). Distances traveled
ranged from negligible movement to 3,997
nautical miles (blue shark). Overall, and in
some cases, seasonal distributions, as well
as movements of tagged sharks, are mapped
with respect to the Atlantic Ocean and marginal seas, state boundaries, the 200 mile
United States Exclusive Economic Zone,
and international and territorial waters of
other countries. Detailed profiles are provided for 14 noteworthy shark species where
the updated data have significantly extended previous ranges and movements.

The National Marine Fisheries Service (NMFS) Cooperative Shark Tagging Program (CSTP) is one of the
largest and longest running shark tagging programs in the world. The CSTP
is a collaborative effort among recreational anglers, the commercial fishing
industry, biologists, and the NMFS to
study the life history of sharks in the
Atlantic Ocean. Initiated in 1962 by
John G. Casey, volunteer participation began with an original group of
74 anglers involved in tagging feasibility studies in 1963. The program expanded in subsequent years to include
volunteers distributed along the entire
North American and European Atlantic coasts including the Gulf of Mexico. The 52-year database represents
the efforts of thousands of particiNancy E. Kohler (retired) and Patricia A. Turner
(retired) were with the National Marine Fisheries Service, NOAA, Northeast Fisheries Science
Center, Narragansett Laboratory, 28 Tarzwell
Drive, Narragansett, RI 02882-1199 (email
[email protected]).

81(2)

with body size (Speed et al.; 2010; Secor, 2015). These data have spurred
more detailed genomic and electronic tagging studies into adaptive significance and contribute to placement
and monitoring of marine reserves or
related conservation measures (McCandless et al., 2007; Portnoy et al.,
2015). Life history parameters such as
age, growth, reproduction, and mortality can be independently estimated
from tagging data providing basic inputs into stock assessment models and
verification of traditional biological
estimates (Brooks et al., 2010). Mark
and recapture may be the most costeffective, reliable, and direct means
to obtain population data for sharks
but there are also specific challenges
(Everhart and Youngs, 1981; Gordon,
1990).
The spatial and temporal scales involved with tracking shark movement can be considerable as many
shark species are moving long distances over whole ocean basins involving multinational fisheries (Kohler et
al., 1998). Other species are difficult
to study because they are inaccessible in oceanic or deepwater habitats,
while some species are rare or with
low population numbers (McLaughlin
and O’Gower, 1971). Even seemingly accessible species, such as those in
coastal habitats, present issues such as
being large, active predators not easily available for any length of time
(Kneebone et al., 2012). Due to these
limiting factors, information on long
distance movements, migratory ranges, and movement patterns is lacking
for many shark species (Ferreira et al.,
2015). The paucity of information hinders assessments of population trends
or spatial management options (Lea
1

et al., 2015; Queiroz et al., 2016) at a
time when data are needed to gain insight into shark migration pathways,
population structure, spatial vulnerability to fisheries, and ecological impacts (Sims, 2010; Campana, 2016).
The collaborative CSTP, with volunteers tagging and recapturing throughout the range of many of these species
over a long time period, is an effective
and efficient approach to address these
challenges and meet these conservation and management goals of sustainability.
This report addresses the critical
need for information on the movements and distribution patterns of Atlantic shark species, particularly over
large spatial and temporal scales. It
updates and replaces a previous atlas
(Kohler et al., 1998), adding an additional 20 years of CSTP M/R information for coastal and pelagic shark
species. Summary statistics on numbers of fish tagged and recaptured,
long distance and time at liberty maximums, and seasonal and transboundary
movements are tabulated. Additionally, 14 species-specific profiles are included to highlight updates for sharks
where new data have extended previous ranges and movements.
Materials and Methods
Tagging methods have remained
consistent during the past 52 years
(Kohler et al., 1998; Kohler and Turner, 2001). The two principal tags are
a dart tag (M tag) and a fin tag (Rototag) (Fig. 1). The dart tags, in use
since 1965, are sent to participants on
self-addressed return postcards for recording tagging information (species,
size, and sex of shark, date and location of tagging, and gear) along with
a tagging needle, tagging instructions,
and shark identification and fisheries
management information. Fin tags are
used primarily by participating biologists. Each M tag has a capsule with a
legend containing contact information
and a request for information. Thus,
when a tagged shark is re-caught similar data to that collected at tagging is
obtained which allows for the calculation of time at liberty, displacement,
2

Figure 1.—The two principal tags,
(A) M dart tag and (B) Jumbo Rototag, used in the NMFS Cooperative
Shark Tagging Program (1962–
2013). The numbered tagging legend is contained within the capsule
at the end of the dart tag.

and growth. In 1988, a hat with an embroidered logo replaced the previous
monetary reward.
Data quality assurance and quality control (QA/QC) is an ongoing
process for the program (Kohler et
al., 1998). The CSTP has continued
to improve its participants’ ability to
correctly identify species (SchulzeHaugen et al., 2003), as well as to contact individuals, when possible, on
missing or suspect information (e.g.,
size estimate too small or large based
on knowledge of species, and outlier locations on land). Other auditing
procedures include rapid follow-up to
the person returning the recapture report to verify data, obtaining pictures,
and generating distribution maps by
species of previous year’s M/R data to
verify locations.
These efforts have been aided, in
particular, by the development of the
NMFS Integrated Mark/Recapture Database System (I-MARK), which was
brought online in 2008 to provide a
platform to keep multi-species tagging
program data in a common format for
management and analysis. I-MARK
is a web application written in PERL
scripting language used for data input and quality control of the M/R
data, which is maintained in an Ora-

cle1 database. I-MARK was designed
to facilitate the tracking of fish (e.g.,
one fish re-tagged several times with
different tags) and tag numbers independently and consists of several web
application modules including an inventory of tags, initial release events,
subsequent recapture events, bulk data
entry of large groups of tags (e.g.,
from research surveys), contact name
and address information, map display,
reports, and statistical queries. Fate of
the animal, fate of the tag, double tags,
and multiple recaptures can be accommodated within the database. Most
importantly, the web application provides extensive QA/QC during the entry and maintenance of the I-MARK
data; standard audits check data type,
outlier locations, and other allowable
values such as maximum sizes. More
complex validations are also included
to check relationships between the fate
of animal, the fate of tag, and event
type. To provide feedback to the participants, the web application generates a letter that includes a map and
information on size, location of tagging and recapture, time at liberty, and
distance traveled of each shark that is
1Mention

of trade names or commercial firms
does not imply endorsement by the National Marine Fisheries Service, NOAA.

Marine Fisheries Review

Table 1.—Summary of tag and recapture data for 35 species of sharks from the NMFS Cooperative Shark Tagging Program (1962–2013).
            Species
Common
Scientific
Atlantic angel shark
Atlantic sharpnose shark
Basking shark
Bigeye thresher
Bignose shark
Blacknose shark
Blacktip shark
Blue shark
Bonnethead
Bull shark
Common thresher shark
Crocodile shark
Dusky shark
Finetooth shark
Galapagos shark
Great hammerhead
Greenland shark
Lemon shark
Longfin mako
Night shark
Nurse shark
Oceanic whitetip shark
Porbeagle
Reef shark
Sand tiger
Sandbar shark
Scalloped hammerhead
Shortfin mako
Silky shark
Smalltail shark
Smooth dogfish
Smooth hammerhead
Spinner shark
Tiger shark
White shark

Number of
Number of
sharks tagged sharks recaptured

Recapture Max. distance
rate (%) traveled (nmi)

Max. time
at liberty (yr)

Squatina dumeril
170
0
0.0
-  
Rhizoprionodon terraenovae 4,977
79 1.6
570 7.3
Cetorhinus maximus
168
0
0.0
-  
Alopias superciliosus
400 12 3.0
2,067 10.5
Carcharhinus altimus
175 11 6.3
1,778 11.2
Carcharhinus acronotus
2,958
35 1.2
226 9.9
Carcharhinus limbatus
10,293
269 2.6
1,183 9.3
Prionace glauca
117,962 8,213
7.0 3,997
15.9
Sphyrna tiburo
5,057
221 4.4
302 7.0
Carcharhinus leucas
2,129
36 1.7
628 6.7
Alopias vulpinus
203 4 2.0
271 8.0
Pseudocarcharias kamoharai
20
0
0.0
-  
Carcharhinus obscurus
8,465
164
1.9 2,052
16.1
Carcharhinus isodon
2,807
60 2.1
365 4.9
Carcharhinus galapagensis
422
18 4.3
1,087 7.1
Sphyrna mokarran
282 5 1.8
649 3.4
Somniosus microcephalus
68 1 1.5
0 1.0
Negaprion brevirostris
3,231 277
8.6 494 10.9
Isurus paucus
106 6 5.7
1,852 5.5
Carcharhinus signatus
289 19 6.6
1,456 13.8
Ginglymostoma cirratum
2,186 175
8.0 385 11.6
Carcharhinus longimanus
643 8 1.2
1,226 3.3
Lamna nasus
1,754
178 10.1
1,216 16.8
Carcharhinus perezii
768
24 3.1
26 9.2
Carcharias taurus
2,019
73 3.6
641 5.3
Carcharhinus plumbeus
35,929 1,474
4.1 2,039
27.8
Sphyrna lewini
3,537
62 1.8
902 9.6
Isurus oxyrinchus
8,525
1,148 13.5
3,043 12.8
Carcharhinus falciformis
1,238
65 5.3
1,288 8.6
Carcharhinus porosus
24
0
0.0
-  
Mustelus canis
1,186
37 3.1
460 6.8
Sphyrna zygaena
269 7 2.6
496 2.1
Carcharhinus brevipinna
1,723
27 1.6
861 6.8
Galeocerdo cuvier
9,772
709
7.3 3,643
11.2
Carcharodon carcharias
55 2 3.6
546 2.5

then sent to both the tagger and recapturer along with the reward.
Data for this atlas are for sharks
that were tagged and/or recaptured
between 1962 and 2013 in the Atlantic Ocean (North and South) and
its marginal and associated seas, including the Gulf of Mexico, Caribbean Sea, and Mediterranean Sea. Only
those tags or recaptures with year and
location information were included
for analysis. A recapture was excluded if a tagged shark was found dead
(e.g., on a beach) or if only the tag
was found. In addition, sharks released
from aquaria and embryos were omitted from the database; these types of
releases may artificially indicate species presence outside of their normal
distribution. Distance traveled is the
great circle distance in nautical miles
(nmi) between tag and recapture locations for a single event. For species
(e.g., sandbar shark) that traveled between the Atlantic and Gulf of Mexico, the great circle distance would
cross land through Florida, a significant underestimate, so the routes be-

tween mark and recapture for these
fish were assumed to pass through two
standard offshore waypoints, southeast
and southwest of Florida (bordering
the Straits of Florida).
Cooperative Shark Tagging Program
M/R data for sharks are displayed by
species in a two-page format with
standard sets of figures (maps and
graphs). For ease in locating data displays, species sections appear in alphabetical order by common name which
are used throughout the text (see Table 1 for scientific names). Maps are
displayed in a longitude/latitude WGS
84 projection with the United States
(U.S.) Exclusive Economic Zone
(EEZ) boundary represented by a dotted-dashed line and 200 meter (m)
contour by a solid line. Latitude and
longitude grids are in 10° increments
for all maps. Data for each species
are presented in similar format. Overall summary information is presented
as total and by sex and includes numbers that were tagged and recaptured,
recapture percentage, mean and maximum distance traveled (nmi), and time

at liberty (years). Yearly summaries include the number of sharks tagged and
recaptured and percent recapture plotted on three distinct line graphs with
the same year scale for comparison.
Percent recapture was calculated as the
number of sharks subsequently recaptured from a particular release year divided by the number of sharks tagged
that year. Maps include

1) Recapture distribution where
M/R information for each species is displayed on a single
map, and only those data with
both mark and recapture locations were displayed. For distances traveled greater than or
equal to 10 nmi, arrows depict
the point of tagging (origin of
arrow) and point of recapture
(arrow head). For distances traveled less than 10 nmi, recaptures
are distinguished by a triangle.
 2) Distribution locations by sex
where all tag and recapture locations of an individually marked

81(2) 3

Figure 2.—Yearly summary of number of sharks tagged, recaptured, and percent recapture rate for all sharks in the NMFS
Cooperative Shark Tagging Program (1962–2013).

fish are plotted to show the total
distribution of a species in a geographic area. On some maps, an
inset was included (e.g., blacktip
shark) to increase clarity for the
majority of the data.
 3) Seasonal distribution of the
western North Atlantic M/R locations during spring (March,
April, May), summer (June, July,
August), fall (September, October, November), and winter (December, January, February). To
clarify and facilitate comparisons among seasons, only the
western North Atlantic M/R locations were displayed on the
same map scale for each season. For species where no western North Atlantic data exist
(e.g., crocodile shark), all data
were displayed by season. An
additional two-page layout was
added for species with large
numbers of tags and recaptures
(blue shark) and/or extensive
geographic ranges (blue shark,
shortfin mako, tiger shark).
Results and Discussion
Totals
Between 1962 and 2013, a total of
229,810 sharks of 35 species were
tagged and 13,419 sharks of 31 species were recaptured (Table 1, Fig. 2).
4

Figure 3.—Summary of tag releases by species in the NMFS Cooperative Shark
Tagging Program (1962–2013).

Fishermen representing 32 countries
have tagged sharks for the CSTP and
59 countries are represented in the tag
returns. Seven species accounted for
85% of the tags, and a single species,
the blue shark, accounted for 51%
of the total tagged sharks (Table 1,

Fig. 3). The number of sharks tagged
ranged from 20 for the crocodile shark
to 117,962 for the blue shark. Most
species (31) had more than 100 sharks
tagged.
Numbers of tag returns by species
ranged from 0 to 8,213. Seven species
Marine Fisheries Review

accounted for 91% of the recaptures,
and the blue shark accounted for 61%
of the recaptured sharks (Table 1, Fig.
4). For most species (25), less than
100 fish were recaptured. The recapture rate ranged from 0.0% (Atlantic
angel shark, basking shark, crocodile
shark, and smalltail shark) to 13.5%
(shortfin mako) with an overall mean
of 5.8%.
Overall, recreational fishermen
(56%), most using rod and reel, accomplished the majority of the tagging (Fig. 5) followed by biologists
(32%) using longline and net gear.
The majority of tag returns were from
commercial fishermen (50%) using longline and net gear and anglers
(43%) using rod and reel (Fig. 6).
Times at Liberty
With the inclusion of the additional
20 years of CSTP data since Kohler et
al. (1998), time at liberty has increased
substantially for 22 species. These increases ranged from months (e.g.,
blacknose shark, dusky shark, night
shark, tiger shark) to many years (e.g.,
blue shark, bonnethead, lemon shark,
porbeagle, spinner shark) (Table 1).
The longest time at liberty for any individual shark remained at 27.8 years.
This record is for a sandbar shark that
was tagged by Narragansett NMFS biologist Charles Stillwell, fishing with
a gill net in Great Machipongo Sound,
VA, in June of 1965, and recaptured
by a commercial shark longline fisherman east of Daytona Beach, FL, in
March of 1993. Overall, individuals
of 11 species were at liberty for more
than 10 years, 14 were at liberty between 5–10 years, and 6 for less than 5
years. Most recaptured sharks were at
liberty for less than 1 year (59%) and
from 1 to 5 years (36%) (Fig. 7).
Knowledge of longevity estimates is
needed for calculation of lifetime fecundity, which is especially critical for
long lived, slow growing animals, such
as sharks. Longevity can be estimated using the oldest aged specimen in a
traditional hardpart ageing study (minimum estimate), calculations from von
Bertalanffy growth function parameter
estimates (e.g., L∞), or using radiocar81(2)

Figure 4.—Summary of tag returns (recaptures) by species in the NMFS Cooperative Shark Tagging Program (1962–2013).

bon dating in combination with band
pair counts (Natanson et al., 2002; Andrews et al., 2011). Maximum time at
liberty records from M/R data serve
as direct evidence of longevity, particularly if size (age) at tagging is taken into account (Casey and Natanson
1992; Frazier et al., 2015). Times at
liberty verify calculated data for certain species (e.g., shortfin mako, blue
shark, sandbar shark), or serve as a
proxy for minimum lifespan estimates
for species with no published age data.
Longevity estimates for shortfin
mako based on analyses of vertebral
centra ranged from 21 to 38 years, for
males and females, respectively (Natanson et al., 2006). Long-term recaptures reported in Natanson et al.
(2006), include a male at liberty for
12.8 years (estimated age at recapture, based on length at tagging, was
21 years) and a female at liberty for

10.5 years (estimate age at recapture
was 32 years). In 2015, a male shortfin mako (not included in this analysis) that was tagged as a 1+ years was
recaptured after 19.1 years at liberty.
This recapture also verifies a 20+ year
longevity for the species. It is notable
that this fish was recaptured only 55
nmi from its original tagging location.
The oldest directly aged blue shark
was 16 and 15 years for males and females, respectively, in a validated age
study of blue sharks in the North Atlantic Ocean (Skomal and Natanson,
2003). Calculated longevities from
that study based on 95% and 99%
of L∞ ranged from 16.5–26.1 years
(Skomal and Natanson, 2003). The
longest time at liberty for a blue shark
in the CSTP is 15.9 years (Table 1).
Based on provided estimates of length
and weight, this male blue shark was
8–11 years of age at tagging and thus

5

would have been between 24+ and 27+
years at recapture; verifying and possibly extending the published longevities for this species.
As noted previously, the overall
CSTP maximum time at liberty record is for a sandbar shark at large for
27.8 years (Table 1). Age at recapture
for this fish was estimated at 33 to 36
years which confirms the 30+ years reported from growth bands in vertebrae
and bomb radiocarbon dating (Andrews et al., 2011).
Distances Traveled

Figure 5.—Summary of tag releases by (A) industry of participants
and (B) gear in the NMFS Cooperative Shark Tagging Program
(1962–2013).

6

Distances traveled ranged from negligible movement to 3,997 nmi (blue
shark) (Table 1). Two peaks in distance
traveled were observed: one at less
than 100 miles (46%) and the second
between 1,000 to 2,000 nmi (17%)
(Fig. 8). Maximum distance traveled
reflects the longest distance between
mark and recapture for a single event
(Table 1). Since Kohler et al. (1998),
maximum distances traveled increased
for 20 species between 10 nmi (reef
shark) to 1,700 nmi (tiger shark) (Table 1). Overall, individuals of three
species traveled distances greater than
3,000 nmi (blue shark, shortfin mako,
tiger shark); three species traveled
distances between 2,000–3,000 nmi
(bigeye thresher, dusky shark, sandbar shark), and eight species traveled
between 1,000–2,000 miles (bignose shark, blacktip shark, Galapagos
shark, longfin mako, night shark, oceanic whitetip shark, porbeagle, and
silky shark). Individuals of seven species traveled distances between 500–
1,000 nmi (Atlantic sharpnose shark,
bull shark, great hammerhead, sand tiger, scalloped hammerhead, spinner
shark, and white shark).
In some instances, a shark may be recovered multiple times after the initial
tagging and is subsequently released
with the original tag in place or retagged with a new tag. For these fish,
the total distance traveled may be larger than the furthest for a single tagging
event (maximum distance travelled)
as is the case for both the nurse shark
(385 nmi to 387 nmi) and sand tiger
(641 nmi to 755 nmi). A nurse shark
Marine Fisheries Review

was originally tagged near Bimini,
Bahamas, and recaptured in the same
area (2 nmi) 2.7 years later; a second
recapture occurred off Cuba (385 nmi
from the first recapture location) after
2.3 years. A sand tiger was recaptured
three times and was at liberty for nearly
3 years. The fish was originally tagged
off the coast of Maryland in early fall
of 2009. Its first recapture was to the
north, off Delaware, in late summer of
2010 (20 nmi); it was again recaptured
to the south, off South Carolina, in late
spring of 2011 (364 nmi from the first
recapture location); and its third recapture was again to the north, off Delaware, in late summer of 2012 (371 nmi
from the second recapture location).
Details of these multiple recaptures
over time also highlight a north-south
seasonal component to the migrations
of this species.
Transboundary Movements
Sharks can be highly mobile, moving long distances often over entire
ocean basins. Detailing these movements with respect to national and
international boundaries is critical
for the sustainable management of
shark populations. Overall, 22 species
crossed the U.S. EEZ boundary (Table
2). Twenty-seven species occurred in
the Gulf of Mexico. Thirteen species
moved into the Gulf of Mexico from
the Atlantic and 11 species moved out
of the Gulf of Mexico into the Atlantic. Twenty species occurred in the
Caribbean Sea. Ten moved into the Caribbean Sea from the Atlantic Ocean;
one moved in from the Gulf of Mexico (tiger shark), and one was recaptured at the entrance of the Caribbean
Sea from the Gulf of Mexico (dusky
shark). Two species (blue shark, tiger
shark) were tagged in the Caribbean
Sea and moved to the Atlantic Ocean.
Ten species occurred in the South Atlantic, and one species, the blue shark,
crossed the equator from the North
Atlantic Ocean to the South Atlantic
Ocean (Table 2).
Seasonal Distribution

Figure 6.—Summary of tag returns (recaptures) by (A) industry of
participants and (B) gear in the NMFS Cooperative Shark Tagging
Program (1962–2013).

Seasonal migrations are common in
many animal taxa and can be defined
81(2)

7

Figure 7.—Times at liberty for all tag returns in the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 8.—Distances traveled by sharks tagged and recaptured in the NMFS Cooperative Shark Tagging Program
(1962–2013).

as movements between distinct habitats where organisms take advantage
of available resources at each location
to enhance fitness (Ramenofsky and
Wingfield, 2007). Sharks, like most
marine species, exhibit distinct thermal preferences (Casey and Kohler,
8

1992) so shifts in the distribution of
species within the North Atlantic can
be expected with seasonal changes in
water temperature. Many of the species
in the CSTP (e.g., Atlantic sharpnose
shark, blacknose shark, bull shark, finetooth shark) are at their most north-

erly range in summer and fall and
are located further south in the colder months of winter and spring. These
species are present on the U.S. mid-Atlantic and northeast continental shelf
(north of Cape Hatteras, NC) during
the warmest months of the year. Some
Marine Fisheries Review

81(2)
Table 2.—Summary of occurrence and transboundary movement for 35 species of sharks from the NMFS Cooperative Shark Tagging Program (1962–2013).
Shark species tagged and/or recaptured in:

NA = Not applicable due to no recaptures.

1

x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x
x
x

x

x

x
x

x
x
x

x
x
x
x

x
x

x
x

x
x

x

x

x

x
x
x
x

x
x

x

x
x
x
x
x

x

x

x

x

x

x

x

x

x
x

na1

na

na

na

na

na

na

na

na

na

x
x

x
x

x
x

x
x
x
x

x
x

x

na

na

na

x

x

x

x

x
x
x

Gulfof Mexico

x
x

x
x
x
x
x
x
x

South Atlantic

x
x

x
x
x
x
x
x
x

Gulfof Mexico

x
x

x
x
x
x
x
x
x

Caribbean Sea

x
x
x
x
x

x

x
x
x
x
x
x
x
x

x

Moved
out:

Moved into:

U.S. EEZ

x
x
x
x
x

x

x

South Atlantic Ocean

x
x
x
x

x
x
x
x

x

x
x

x

x
x

Easterm North Atlantic

x
x
x
x

x
x

x

x
x

x
x

Winter

x
x
x
x

x
x

x
x
x

x
x

Fall

x
x
x
x
x
x
x
x

Summer

x
x
x
x
x
x
x

x

Spring

x
x
x
x
x
x
x
x

Winter

x
x

Fall

x
x

Summer

Fall

x
x

x
x
x
x
x

Moved
across:

Caribbean Sea

Spring

Summer

x
x
x
x
x

x
x
x
x
x
x
x
x
x
x
x

x
x
x
x
x

Gulf of Mexico

Winter

Spring

10
214
5
79
5
67
583
3,894
111
185
15
9
329
19
10
48
0
581
23
22
245
125
181
141
76
1,011
226
447
250
2
59
6
141
1,812
4

Winter

76
947
18
144
17
914
2,096
21,998
759
627
54
0
1,156
467
21
37
45
429
17
48
369
150
768
210
263
5,966
586
1,779
291
9
241
33
434
1,899
8

South of
Cape Hatteras, NC

Fall

40
2,317
74
80
61
1,715
5,968
93,333
3,089
846
122
8
5,908
2,061
397
105
5
1,417
27
100
1,102
139
579
122
1,405
23,507
1,368
6,770
231
8
825
186
898
4,135
38

Summer

44
1,577
71
109
103
295
1,904
6,801
1,316
503
15
3
1,233
318
12
97
19
1,074
45
136
644
237
396
318
347
6,885
1,416
658
530
5
98
51
277
2,613
7

Spring

Winter

Squatina dumeril
Rhizoprionodon terraenovae
Cetorhinus maximus
Alopias superciliosus
Carcharhinus altimus
Carcharhinus acronotus
Carcharhinus limbatus
Prionace glauca
Sphyrna tiburo
Carcharhinus leucas
Alopias vulpinus
Pseudocarcharias kamoharai
Carcharhinus obscurus
Carcharhinus isodon
Carcharhinus galapagensis
Sphyrna mokarran
Somniosus microcephalus
Negaprion brevirostris
Isurus paucus
Carcharhinus signatus
Ginglymostoma cirratum
Carcharhinus longimanus
Lamna nasus
Carcharhinus perezii
Carcharias taurus
Carcharhinus plumbeus
Sphyrna lewini
Isurus oxyrinchus
Carcharhinus falciformis
Carcharhinus porosus
Mustelus canis
Sphyrna zygaena
Carcharhinus brevipinna
Galeocerdo cuvier
Carcharodon carcharias

Fall

Atlantic angel shark
Atlantic sharpnose shark
Basking shark
Bigeye thresher
Bignose shark
Blacknose shark
Blacktip shark
Blue shark
Bonnethead
Bull shark
Common thresher shark
Crocodile shark
Dusky shark
Finetooth shark
Galapagos shark
Great hammerhead
Greenland shark
Lemon shark
Longfin mako
Night shark
Nurse shark
Oceanic whitetip shark
Porbeagle
Reef shark
Sand tiger
Sandbar shark
Scalloped hammerhead
Shortfin mako
Silky shark
Smalltail shark
Smooth dogfish
Smooth hammerhead
Spinner shark
Tiger shark
White shark

Summer

Species

North of
Cape Hatteras, NC

Spring

Total number of tags and
recaptures by season

x

x

x

x

x
x

na

na

x

x

x
x

x

x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x
x

x
x
x
x

x
x
x
x
x
x

x
x
x
x
x
x

x
x
x
x
x
x

x
x
x
x
x
x

x

x

x
x
x
x

x
x
x
x

x
x
x
x
x

x
x
x
x

x
x
x
x

x
x
x
x
x

x
x

x

x
x

x
x
x
x
x

x

x

x

x
x
x
x

x
x
x
x

x
x

x

x

x

x

x
x
x

x
x

x
x

x
x

x

x

x
x

x
x

x
x

x

x

x

x
x
x

x
x
x

x

x
x

x
x
x
x

x
x
x

x
x

x
x

x

x

x
x

x

x
x

x
x
x
x
x
x

x

x
x

x
x

x
x

x

x
x
x
x

x
x
x

x
x

na

na

na

x
x

x

x
x

x
x
x
x
na

na

x
x

9

species (e.g., tiger shark, dusky shark,
shortfin mako, and blue shark) also occur in the adjacent shelf edge and oceanic waters, while others mainly occur
in the waters of the deeper slope areas and canyons (e.g., night shark)
or in oceanic waters (e.g., oceanic
whitetip shark). With the exception of
five species (crocodile shark, Galapagos shark, nurse shark, reef shark, and
smalltail shark), all shark species were
found north of Cape Hatteras, NC, in
the warmer summer months; these five
species are not found or do not generally range this far north along the U.S.
East Coast.
Movement patterns seen in the
CSTP database include latitudinal and inshore-offshore migrations
in response to changing water temperatures and such movements have
been shown for almost all families
of tagged sharks, e.g., heterodontids
(McLaughlin and O’Gower, 1971),
lamnids (Bruce, 1992; Casey and
Kohler, 1992), carcharhinids (Stevens,
1976; Tricas, 1977; Olsen, 1984; Francis, 1988), sphyrnids (Clarke, 1971),
and squalids (Holland, 1957; Jensen et al., 1961; Templeman, 1976).
Monthly variations in water temperatures, changes in the path of the Gulf
Stream, and the presence of variable
oceanic features (e.g., warm and cold
core rings, thermal fronts) can, therefore, alter the seasonal distribution of
species from year to year.
Data Interpretation Challenges
The shark distributions presented
here from the CSTP database (Table
2, Fig. 9–43) are based on presenceabsence data and thus the number of
species during any season can be considered a minimum. There are many
reasons why a few individuals of a
species are tagged. Some species are
naturally uncommon or rare throughout their range (e.g., white shark, longfin mako) and, therefore, a low catch
rate would be expected. Other sharks
may inhabit areas outside of the principal fishing and tagging effort (e.g.,
oceanic whitetip shark) and/or may not
be present during the primary fishing
season. Some may be difficult to iden10

tify (e.g., bignose shark), are not readily caught (e.g., basking shark, whale
shark), or commonly tagged (e.g.,
smooth dogfish) by members of the
program. Some of the smaller species,
such as Atlantic sharpnose sharks,
spiny dogfish, and smoothhounds are
so numerous that only a small percentage are tagged relative to the number
caught, particularly since we discourage use of our tags on sharks less than
3 feet. For these species, the residence
times and seasonal distributions are
not well characterized by the CSTP
data. In addition, fishing effort in
some seasons may be low (e.g., winter, spring) or some species may not
be selected by the fishing gears active
in particular areas/seasons. The data,
therefore, may under-represent the
species present during those months.
A prime example exists for the blue
shark in the Northeast U.S., which
is the main tagging area for this species. The weather limits fishing in winter, thus the season is primarily May
through November; outside this fishing season, total tag numbers may be
low due to reduced tagging effort.
The number of fish tagged and recaptured is influenced by many factors
and does not directly reflect population abundance or correlate with population trends. For example, tagging
effort can vary due to annual changes in fishing effort, weather conditions,
water temperature, number of participants in the CSTP, occurrence of research cruises, opening or closure of
a commercial fishery, and number of
tags available. All these variables are
difficult to measure and may mask any
direct correlation of number of tags
used per year and population size fluctuations. Specifically, the blue shark, is
an abundant species and because of its
low economic value, many are tagged
and released; while the shortfin mako
is prized by both recreational and commercial fishermen, tagged and released
in relatively low number, and yet has
the highest overall percent recapture.
Life history characteristics may also
influence tagging and recapture probability, e.g., a species that stays in an
accessible area for extensive periods

of time. For example, the nurse shark,
is more subject to capture and recapture than a species that is highly migratory. Some species live in deeper
waters or farther offshore and are not
present in areas during the primary
fishing season, or are simply not readily caught. For instance, 168 basking
sharks were tagged by members of the
CSTP, but none have been recaptured.
This is because basking sharks are relatively easy to tag while they are free
swimming but are not often caught on
fishing gear and are not subject to directed commercial or recreational fisheries.
Another data interpretation challenge for the CSTP is the lack of information on tag reporting rates,
which can be critical to estimating
abundance, survival, and exploitation from a tagging program (Pine et
al., 2003). There are several methods
used to estimate reporting rates including the use of high reward tags
and planting tags in specific fisheries
(reviewed in Pollock et al., 2001). The
CSTP has not conducted any studies
to date to estimate reporting rate by
species or tag type, however, the future use of a high reward tag should
be examined.
The CSTP spans more than 50
years and thus reported shark distributions may represent historic ranges
and distributions for some species. Alternatively, range expansions may be
associated with environmental change
(e.g., water temperature), expanding
populations, or movement into new areas to exploit a food supply. Perceived
range changes may be due to increased
tagging by a user group, increased tagging in a particular area, and/or increased access to a species or species
life stage. An example of this is the
expansion of the NMFS Cooperative
Atlantic States Pupping and Nursery
(COASTSPAN) program, which has
substantively increased the tagging effort by biologists in coastal areas of
the Atlantic states used as pupping and
nursery grounds for coastal shark species. The expanded COASTSPAN program has produced substantially more
M/R data for the Atlantic sharpnose
Marine Fisheries Review

shark, blacknose shark, bonnethead, finetooth shark, sand tiger, smooth dogfish, and sandbar shark. In the early
years of the CSTP, many of the sharks
tagged in the southeastern U.S. were
from shore; fishing clubs such as the
Florida Shark Club accounted for
much of the recreational tagging off
Florida. There has been a recent resurgence of shore-based recreational shark fishing in the southeastern
U.S. and an expansion into the northeastern U.S. and more blacktip shark,
bull shark, lemon shark, and other
nearshore species are being tagged as
a result of this increased shore-based
fishing effort.
The long-term data generated by
the CSTP have shown that it is vital to
have a large database to determine the
movements of sharks. Just since the
publication of Kohler et al. (1998), the
increased numbers of sharks crossing
various boundaries and shark species
that have shown differences in movements show the importance having
large numbers of each species tagged.
For example, until the CSTP was in its
34th year, no known tiger shark had
crossed the Atlantic. Using a small
short-term subsample would likely
bias the data; only conventional tagging can provide the data reported in
this study. However, these types of tags
do have limitations. Conventional tagging relies on the physical recapture of
a shark and location data are only taken at tagging and recapture. Additionally, no environmental or behavioral
data are recorded. While satellite tags
also have drawbacks (e.g., limited battery life, expense, and up to 60 nmi error in location data), supplementing
the conventional tag data with electronic tagging methods, such as popup satellite archival tags and acoustic
tag methods, can provide details unavailable to conventional tagging. By
combining the larger database of conventional tagging, with the details of
the electronic tags, we can fill the spatiotemporal gaps in conventional tag
data and add environmental details obtained from electronic tagging to determine not only where the sharks are
going but how and why.

Synopses for Selected Species
With the addition of 20 years of
M/R data (over 123,000 sharks tagged
and 8,000 sharks recaptured) to the
CSTP since the previous atlas (Kohler
et al., 1998), new information has extended, in some cases, species ranges and distributions farther north and
south and illustrated more transboundary movements and migrations into
specific bodies of water (Table 2; Fig.
9–43). In addition, M/R information
was included for two additional species of shark (crocodile shark, smooth
dogfish). Selected species-specific profiles are included to highlight noteworthy updates for sharks where new data
have extended previous ranges and
movements.
Bigeye Thresher
The bigeye thresher is reported as an
oceanic and coastal species, virtually
circumglobal in tropical and temperate seas (Castro, 2011). In the Atlantic,
they are distributed from Cape Cod
(Castro, 2011) to southern Brazil, including the Gulf of Mexico and Caribbean Sea in the west, and from
Portugal to South Africa including the
Mediterranean Sea in the east (Compagno, 2001). This species reportedly ranges from intertidal areas to at
least 500 m deep but is mostly found
at depths below 100 m (Compagno,
2001). In this study, however, all bigeye thresher M/R locations along the
U.S. Atlantic and Gulf coasts were in
deep water, on or outside the 200 m
depth contour (Fig. 12b). Locations
ranged from offshore of Cape Cod,
MA, to Brazil including the Gulf of
Mexico and Caribbean Sea, and across
the Atlantic between the Equator and
44°N latitude. Evidence of bigeye
thresher occurrence in the Caribbean
Sea includes multiple tagging events
and two recaptures. These recaptures
showed movement across the Caribbean Sea (tagged SE Puerto Rico, recaptured off Panama; 816 nmi; 2.4
years at liberty) and movement into
the Caribbean Sea from the mid-Atlantic (Fig. 12a). The latter recapture
is the longest distance traveled (2,067

nmi) with the shark tagged at 09°50N,
32°55W and recaptured off Puerto Cabello, Venezuela (10°56N, 67°55W)
(2,067 nmi; 9.5 years at liberty). Maximum time at liberty was 10.5 years
for a bigeye thresher (Table 1).
Blacktip Shark
The reported western North Atlantic range for the blacktip shark is
from coastal New England to Brazil,
including the Gulf of Mexico, and as
sightings north of Cape Hatteras, NC
(Castro, 2011). The range observed
in the CSTP M/R data is from Delaware Bay to French Guiana, including
the Gulf of Mexico and Caribbean Sea
(Fig. 15b). Seasonal north-south Atlantic coastal migrations occurred with
movements across the Gulf of Mexico from Texas to Mexico, and Bimini,
Bahamas to Cuba. No migrations were
documented between the Atlantic and
Gulf of Mexico (Fig. 15a) supporting
two stock management (NMFS, 2006).
Long distance movements were shown
for two blacktip sharks between St.
John, U.S. Virgin Islands (USVI) (Caribbean Sea) to Cape Canaveral, FL
(1,049 nmi) and Saint Andrew Sound,
GA (1,183 nmi—maximum distance
traveled). Maximum time at liberty
was 9.3 years for a blacktip shark that
was tagged and recaptured by biologists and moved from South Carolina
to Florida.
Blue Shark
The blue shark is cosmopolitan in
subtropical and temperate oceanic waters (Castro, 2011). In the tropics, the
blue shark exhibits tropical submergence where it occurs at greater depths
than other areas (Compagno, 1984).
Blue sharks have been tagged or recaptured by CSTP participants from
Newfoundland and the Gulf of St.
Lawrence, Canada, to Argentina in the
western Atlantic, including the Gulf of
Mexico and Caribbean Sea, and from
south of Iceland (57°20N, 19°56W) to
just south of the Equator in the eastern
Atlantic including the Mediterranean
Sea. Altogether, these areas represent its previously reported range for

81(2) 11

the Atlantic Ocean (Compagno, 2001;
Castro, 2011) (Fig. 16d).
The spatial distribution of tags
and recaptures over vast areas of the
North Atlantic further substantiates
one stock for this widely distributed species. Long distance movements
of the blue shark were observed between the U.S. Atlantic coast to all
parts of the North Atlantic, including
Grand Banks of Newfoundland (Canada), Azores, Europe, Mediterranean
Sea, Africa, Canary Islands, Cape
Verde Islands, South America, Central
America, Caribbean Sea, Cuba, and
south of the Equator (Fig. 16a). Many
of these migrations occurred within
one year of tagging (Fig. 16c). Maximum distance traveled was 3,997 nmi
for a blue shark tagged off Long Island, NY, and recaptured in the South
Atlantic after 8.4 years at liberty (Table 1). In all, ten blue sharks traveled
across the Equator from release locations off the U.S. northeast coast
(five), Canada (one), United Kingdom (one), Portugal (one), and north
of the Equator (two). The southernmost recapture location was for a
blue shark tagged north of the equator that moved 1,279 nmi due south
after 65 days at liberty. Other recaptures of blue sharks tagged outside
of the U.S. EEZ (Fig. 16b) showed
movement between the eastern and
western North Atlantic, from the Caribbean Sea to Long Island, NY, and
from South America to Canada and
the Azores. Multiple fish were tagged
and recovered in an area to the west
of the Azores and south of the Flemish Cap, showing exchange between
this area and all parts of the North
Atlantic. The area to the west of the
Azores and south of the Flemish Cap
has been found to be a discrete central
North Atlantic nursery for this species
where juveniles can reside for up to at
least 2 years (Vandeperre et al., 2014).
Maximum time at liberty increased
from 8.5 to 15.9 years, approaching
maximum age for the species (see details above), for an individual that was
tagged north of Cape Cod, MA, and
recaptured 873 nmi to the southeast in
the open ocean.
12

Along the U.S. Atlantic coast, blue
sharks were caught year round in the
offshore waters from Cape Cod, MA,
to Florida (Fig. 16e). From December
through March, they remained primarily off the continental shelf. In April,
some fish were tagged inshore off New
York and New Jersey. By May, blue
sharks began to move onto the continental shelf from offshore waters and
were common inshore from Virginia
to Cape Cod, MA. During the warmer months of June through September,
they were found closer to shore from
New Jersey northward into the Gulf of
Maine and off Newfoundland, Canada, but were not abundant on Georges
Bank. In the waters off North Carolina and Virginia, this species was more
likely to be caught in deeper shelf and
adjacent oceanic waters during this
time. As the waters cool in October
and November, blue sharks began to
move offshore and to all parts of the
North Atlantic.
Bull Shark
The bull shark is cosmopolitan in
coastal tropical and subtropical seas
ranging in the western North Atlantic from Chesapeake Bay, VA (Castro,
2011) to southern Brazil (Compagno,
1984). The CSTP M/R locations extend this range farther north with multiple tags deployed in Delaware Bay
(Delaware–New Jersey) (Fig. 18b).
Published records farther north are
from single strays reported off New
Jersey and Massachusetts (Bigelow
and Schroeder, 1948). Tagging occurred throughout the Gulf of Mexico
and into the Caribbean Sea. Recapture
data showed limited movement along
the U.S. Atlantic coast, from Bimini,
Bahamas to Cuba, and one fish that
moved from the U.S. Atlantic coast
into the Gulf of Mexico (Fig. 18a).
The latter recapture was tagged by a
biologist off Delray Beach, FL, and
was recaptured west of Naples, FL,
after 6.7 years at liberty (maximum
time at liberty for a bull shark in the
CSTP). In addition, multiple recaptures (6) were from fish tagged in the
U.S. Gulf of Mexico and subsequently moved into Mexican coastal waters.

Maximum distance traveled among all
tagged bull sharks was 628 nmi from
Texas to Veracruz, Mexico, after 2.9
months at liberty.
Common Thresher Shark
The common thresher shark is distributed worldwide in warm and temperate
waters and off the U.S. Atlantic coast
primarily from Newfoundland, Canada,
to Florida; however, this coastal-pelagic
species is reported as not generally found south of Cape Canaveral, FL
(Castro, 2011). In the eastern Atlantic, the common thresher shark ranges
from Norway to South Africa including the Mediterranean Sea (Compagno,
2011). CSTP data shows tagging locations from Maine to South Carolina in
the western Atlantic and Italy and England in the eastern Atlantic (Fig. 19b).
A noteworthy location for a common
thresher shark was on the Grand Banks
of Newfoundland, Canada, tagged by a
NMFS biologist in September of 2012.
Only four common thresher sharks were
recaptured. Two were tagged and recaptured off the northeastern U.S. traveling
less than 100 nmi and at liberty for 4.2
and 8.0 years (maximum time at liberty). The other two recaptures were in
the eastern Atlantic; one moved from
the English Channel to the Bay of Biscay, France, after 2.7 years (271 nmi;
maximum distance traveled), and the
other was tagged and recaptured off Italy in the Adriatic Sea (6.8 years; 66
nmi) (Fig. 19a).
Crocodile Shark
The crocodile shark is a little-known,
oceanic and circumtropical species, reported in the eastern Atlantic from
southeast of the Cape Verde Islands to
Guinea-Bissau, Guinea, Angola, and
South Africa from the surface to at
least 590 m (Compagno, 2001). In the
western North Atlantic, it can be found
in the Caribbean Sea with one specimen recorded off Virginia in the Gulf
Stream (Castro, 2011). Twenty tagged
fish in the CSTP were distributed along
the equator from Brazil to Ghana between 0° and 35°W longitude and 2°S
and 4°N latitude (Fig. 20b). None of
these tagged fish were recaptured.
Marine Fisheries Review

Finetooth Shark
The finetooth shark inhabits the
coastal waters of the western North
Atlantic and is reported from North
Carolina to Florida, Cuba, Gulf of
Mexico, and southern Brazil (Compagno, 1984). Reported specimens
north of Cape Hatteras, NC, are considered rare and suspected of species
misidentification, as are occurrences
in Mexico, Cuba, and the Caribbean
(Castro, 2011). CSTP M/R locations
for this species were in coastal U.S.
waters from Virginia south and into the
U.S. Gulf of Mexico to the Mexican
border with one fish tagged in Mexican Gulf of Mexico waters (Fig. 22b).
Both northernmost (Machipongo, VA)
and southernmost (Tamaulipas, Mexico) tags were positively identified by
NMFS biologists confirming some of
the extremes of the finetooth shark
geographic range. Recaptures showed
movement along the U.S. east coast between North Carolina and South Carolina and between South Carolina and
Florida (Fig. 22a). All of the latter recaptures were tagged by biologists off
South Carolina with one of these fish
moving 365 nmi south from McClellanville, SC, to Jupiter, FL (maximum
distance traveled) after 0.6 years. The
one recapture in the Gulf of Mexico
(tagged by a biologist) showed movement between St. Vincent Island, FL,
and Biloxi, MS (distance of 192 nmi;
2.2 years at liberty). Maximum time
at liberty was 4.9 years for a finetooth
shark and recaptured 5 nmi away after
4.9 years at liberty.

es were in the warmer months in summer (New York) and fall (New Jersey)
with spring (North Carolina) and winter (Florida) locations further south.
Updated tag sites show locations in
the USVI and on the south coast of
Cuba in the Caribbean Sea. Updated
recapture information shows movements in the Gulf of Mexico from
Louisiana to the Dry Tortugas, FL,
(502 nmi) after 3.4 years at liberty
(maximum time at liberty) and from
the Florida west coast to Campeche,
Mexico (649 nmi; maximum distance
traveled) (Fig. 24a).
Longfin Mako
The longfin mako is an oceanic and
tropical species recorded sporadically in the western North Atlantic. Most
commonly reported off Cuba, other
widely scattered records include Florida, the Bahamas, the Gulf Stream System off the eastern U.S., and southern
Brazil (Compagno, 2001). CSTP M/R
locations are found in the offshore waters from Cape Cod, MA, to Brazil including the Gulf of Mexico, Caribbean
Sea, and east to 36°W longitude (Fig.
27b). Recaptures show movement
from the Gulf of Mexico to the Atlantic coast of Florida and to the northern
coast of Cuba (Fig. 27a). Since Kohler
et al. (1998), data were added on two
recaptures showing movement from
the northern coast of South America
(Guyana, Suriname, French Guiana) to
Venezuela (Atlantic Ocean to Caribbean Sea) and to Delaware (1,852 nmi;
5.5 years; maximum distance traveled
and maximum time at liberty).

Great Hammerhead

Nurse Shark

The great hammerhead is reported to be a coastal-pelagic and semioceanic tropical predator (Compagno,
1984). Its previously recorded western North Atlantic range was from
New Jersey (Kohler et al., 1998) to
Uruguay (Compagno, 1984). Data
presented in the current study document a great hammerhead tagged off
Long Island, NY, in late August 2004
at 40°47N, 71°36W and is a northern
range extension for this species (Fig.
24b). The most northerly occurrenc-

The nurse shark is an inshore, demersal shark of the continental and
insular shelves in tropical and subtropical waters (Compagno, 2001).
The western North Atlantic range
is Cape Hatteras, NC, to Brazil
with two strays recorded in Chesapeake Bay and off Rhode Island (Bigelow and Schroeder, 1948). CSTP
M/R locations in this study fall within the reported range for this species,
with occurrences from North Carolina to Brazil (Fig. 29b). Movements

from the Florida Keys to the Florida
east coast (Gulf of Mexico to Atlantic), Texas to Mexico, and from Bimini, Bahamas, to both the Florida
east coast and to the northern coast
of Cuba (385 nmi, maximum distance
traveled) were observed (Fig. 29a).
Maximum time at liberty is 11.6 years
for a nurse shark tagged and recaptured in the Gulf of Mexico.
The CSTP database has records of
two nurse sharks released and recaptured off New Jersey. The two fish
were released on the same day from
a New Jersey aquarium in September; one was recaptured one day later (2 nmi) and the other 9 days later
(66 nmi). Both were heading south to
warmer waters. These two fish (and all
fish released from aquaria) were excluded from this atlas because they
can show artificial distributions and
movements that fall outside of normal
species ranges. These types of records
may account for some of the reported
strays in published literature for this
species.
Scalloped Hammerhead
The scalloped hammerhead has
circumglobal distribution in coastal warm temperate and tropical seas,
ranging in the western Atlantic from
New Jersey to Brazil, including the
Gulf of Mexico and Caribbean Sea
(Compagno, 1984). CSTP M/R locations extend this range farther north
off Long Island, NY, and offshore as
far north as 41°N latitude (Fig. 35b).
Additionally, tagging locations occurred on the Cuban south coast in
the Caribbean Sea. Movements took
place along the U.S. Atlantic coast
with long distance migrations from
Florida to North Carolina, Maryland,
and New York, and from North Carolina to the southern edge of Georges
Bank and to Cuba (Fig. 35a). The latter recapture is the maximum distance
traveled for an individual scalloped
hammerhead (902 nmi) in the CSTP
database. Updated data in the Gulf of
Mexico show movements within the
Gulf (Mississippi to the Dry Tortugas,
FL), between the U.S. and Mexican
Gulf of Mexico (Texas to Tampico

81(2) 13

and Campeche, Mexico), and from
Gulf waters to the Atlantic (Mississippi to the Atlantic northern coast of
Cuba). Maximum time at liberty was
9.6 years for a scalloped hammerhead
tagged off Miami Beach, FL, and recaptured off Cape Lookout, NC.
Shortfin Mako
The shortfin mako is found in all
tropical and warm temperate seas
(Compagno, 2001). In the western
North Atlantic, the species ranges
from northeast of the Grand Banks of
Newfoundland, Canada (as far north
as 50°N) (Castro, 2011) to southern
Brazil and possibly northern Argentina, including Bermuda, Gulf of Mexico, and Caribbean (Compagno, 2001).
In the eastern Atlantic, it ranges from
Norway to South Africa, including the
Azores and the Mediterranean Sea
(Compagno, 2001). Capture locations
in the CSTP M/R data for the shortfin mako in the western Atlantic range
from 50°N to 40°S and confirm locations at the extreme ends of the published range for this species from east
of the Flemish Cap in the north to off
Buenos Aires, Argentina in the south
(Fig. 36d). In the eastern Atlantic, the
shortfin mako was tagged from northern Spain to south of the Equator off
Gabon (45°N to 2°S). Not commonly reported in the Gulf of Maine (Castro, 2011), updated CSTP data show
shortfin mako from Massachusetts to
Nova Scotia, Canada (Fig. 36f) and recapture movements into and out of the
Gulf of Maine (Fig. 36a). Additional
recaptures in the Gulf of Mexico and
Caribbean Sea show more evidence of
movement between the Atlantic and
Gulf of Mexico (both directions), the
Atlantic and Caribbean Sea, and between U.S. and Mexican waters of the
Gulf.
Long distance shortfin mako recoveries (> 1,000 nmi) at liberty for less
than one year show interesting results
(Fig. 36c). Primarily tagged off the
U.S. Northeast Coast, these fish were
recaptured in the Gulf of Mexico, Caribbean Sea, mid-Atlantic Ocean, and
off Portugal, Morocco, and Western
Sahara. The vast majority of shortfin
14

makos were recovered in an area west
of the Azores (Fig. 36b); fish tagged in
this area show movement to both the
U.S. (from Cape Hatteras, NC, to Cape
Cod, MA) and west toward Europe.
Two shortfin makos tagged off southern Portugal were also recaptured in
the area west of the Azores after less
than one year at liberty. Updated data
show more evidence of shortfin makos crossing the Mid-Atlantic Ridge
demonstrating exchange between the
western and eastern Atlantic. The maximum distance traveled was 3,043 nmi
for a fish that went from Long Island,
NY, to Morocco after 2 years at liberty (Table 1). Maximum time at liberty
was 12.8 years with five shortfin makos at liberty for over 9 years.
Smooth Dogfish
The smooth dogfish is an abundant
coastal species commonly distributed in bays and inshore waters from
Massachusetts southward to Florida and the Gulf of Mexico. They are
also found offshore in the Gulf of
Mexico (Giresi et al., 2015). Northward dispersal is bounded by Cape
Cod, MA, and Nantucket Shoals with
occasional strays found in Massachusetts Bay, the Gulf of Maine, and Passamaquoddy Bay at the mouth of the
Bay of Fundy (Bigelow and Schroeder, 1948). According to Heemstra
(1997), Mustelus canis is replaced by
Mustelus canis insularis in the Caribbean Islands, the Bahamas, Cuba, and
Bermuda. Recent work in the Gulf
of Mexico has suggested the multiple species inhabit this region (smooth
dogfish, Florida smoothhound, Mustelus norrisi, and gulf smoothhound,
Mustelus sinusmexicanus) (Giresi et
al., 2015). These three species can be
distinguished by morphological and
spatiotemporal factors. It would be impossible to unequivocally state that no
cryptic species were tagged; however,
all M/R locations for smooth dogfish
in this study are within the documented geographic and depth range of
Mustelus canis (Kohler et al., 2014).
Smooth dogfish were tagged from the
Gulf of Maine to the Gulf of Mexico
(Fig. 39b). Only two fish were tagged

in the Gulf of Maine; these fish were
caught less than 5 miles from shore,
off Ogunquit, ME. Maximum distance traveled was 460 nmi (Table 1)
for a smooth dogfish tagged off Martha’s Vineyard, MA, and recaptured off
Hatteras Inlet, NC, after 1.1 years. The
maximum time at liberty was 6.8 years
for a female smooth dogfish tagged off
Martha’s Vineyard, MA, that was recaptured 51 nmi to the west of the tag
location. None of the smooth dogfish
moved between the Atlantic and Gulf
of Mexico (Fig. 39a).
Tiger Shark
Tiger sharks are a wide-ranging
coastal-pelagic species, occurring
close to shore and in the open ocean
(Compagno, 1984). They are typically
considered to be residents of tropical
and warm temperate habitats (Ferreira
et al., 2015) with seasonal excursions
into cool temperate areas (Randall,
1992; Last and Stevens, 1994). The
CSTP M/R data (Fig. 42e) show that
tiger sharks are caught year round in
the Caribbean Sea, Gulf of Mexico, and off the U.S. Atlantic coast as
far north as New Jersey. In the warmer months of summer and fall, tiger
sharks are found as far north as Cape
Cod, MA. Recaptures for this species in the Atlantic using conventional
tags has previously shown movements
north and south along the U.S. Atlantic coast; from the U.S. Atlantic coast
to the Gulf of Mexico, Caribbean Sea,
and as far east as 60°W longitude;
from Bermuda to the Gulf of Mexico; and from the Gulf of Mexico to
the U.S. East Coast and Caribbean Sea
(Kohler et al., 1998). Updated information on recaptures from this study
show more evidence of Gulf of Mexico crossings (north–south, east–west)
(Fig. 42b), more exchange between the
U.S. Atlantic coast and Gulf of Mexico; more movements into and out of
the Caribbean Sea (Fig. 42a), and numerous recapture locations across the
North Atlantic Ocean. These long distance movements extend known tiger
shark migrations with recapture locations as far north as 45°N latitude,
as far south as 2.5°N latitude (northMarine Fisheries Review

ern Brazil), and trans-Atlantic movements as far east as the African Coast
(15°W longitude). Some of these extensive movements (> 500 nmi) are
accomplished over a short period of
time (overall mean time at liberty =
1.0 years) with recapture locations in
the Gulf of Mexico, Caribbean Sea,
mid-Atlantic, and eastern Atlantic
Ocean after less than 1 year at liberty (Fig. 42c). Maximum time at liberty
was 11.2 years for a male tiger shark
tagged off Cape Hatteras, NC, and recaptured near Atlantis Canyon (a submarine canyon approximately 100 nmi
south of Cape Cod, MA). Maximum
distance traveled reported in this study
was 3,643 nmi for a female tiger shark
tagged off St. Augustine, FL, and recaptured west of Guinea-Bissau, Africa after 2.5 years at liberty. This is the
longest recorded movement for any tiger shark in the Atlantic Ocean.
Summary
Twenty years of additional CSTP
M/R data provide further insight into
the distributions and movements of the
35 species of Atlantic sharks examined. Distances traveled have been extended for 20 species, some of which
showed exchange into all parts of the
Atlantic Ocean and associated seas.
Times at liberty have been extended for
22 species with some maximums verifying published longevity estimates.
In addition, seasonal distributions have
been included in specific bodies of water. These accomplishments would be
nearly impossible for an individual, or
an individual institution or agency, to
mark and recapture the number of fish
over the geographic and decadal scale
as has been achieved by the CSTP—a
collective of thousands of knowledgeable volunteer recreational and commercial fishermen. This research is
accomplished for little more than the
cost of the tags making the cost/benefit ratio for this program extremely low. The information collected by
CSTP volunteers is a form of public participation in scientific research
(PPSR) or citizen science (Bonney et
al., 2009). The CSTP creates an enormous body of scientific data for under81(2)

standing distributions and migration
patterns for shark species. A further
benefit to PPSR is that tagging has become a socially acceptable component
to recreational fishing, wherein many
anglers solely practice catch, tag, and
release (van der Elst, 1990). The old
adage “the only good shark is a dead
shark” is changing, due partly to the
education received by shark fishermen
through this volunteer shark tagging
program.
Conventional tags continue to have
a valuable role to play in shark conservation and management. Despite
advances in electronic tagging technology, the geographic distributions
and movements for most shark species remain largely unknown (Sims,
2010). Such information, particularly
over large spatial and temporal scales,
is essential for the development of appropriate management strategies (Ferreira et al., 2015) and in determining
the usefulness of conservation measures (Sims, 2010). The wealth of data
garnered from the CSTP highlights the
importance of continuing these longterm tagging programs (Frazier et al.,
2015). Given the fact that shark species are slow growing, long-lived, and
highly mobile, with relatively low return rates for tagged sharks—continued tagging efforts are essential to
provide this critical life history and
population dynamics information.
It is well established that many
shark species segregate by size and
sex (Springer 1960; Clark and von
Schmidt 1965) and undergo partial migrations (Papastamatiou et al., 2013;
Secor, 2015) throughout their lives
with the existence of discrete locations for key life history events, such
as pupping, and mating (Sims, 2010;
Vandeperre et al., 2014). There is also
a distinct seasonal component to these
movements. Understanding these differential migration patterns is crucial for future conservation efforts
(Lea et al., 2015) since long distance
movements could bring different segments of a population into contact
with multiple habitats, ecosystems,
anthropogenic threats (Ferreira et al.,
2015), and overlap with fishing activ-

ity (Queiroz et al., 2016). Finer scale
analysis of the CSTP M/R database
will help bring to light some of these
patterns. Future studies of animal
movements need to be more broadly
based, linking migration research with
ecological and biogeographic studies
(Dingle and Drake, 2007) to begin to
understand the mechanisms and basic driving forces for migration (Alerstam et al., 2003; Ramenofsky and
Wingfield, 2007). In addition, the integration of tagging and population dynamics modelling can greatly enhance
the management and conservation of
highly mobile species such as sharks
(Aires-da-Silva et al., 2009; Braccini
et al., 2016).
Acknowledgments
This report is dedicated to Jack
Casey whose tremendous foresight
and determination created the Cooperative Shark Tagging Program
over 50 years ago. Through his passion, personality, and drive, he created a vast network of enthusiastic and
knowledgeable volunteer taggers who
he called “our windows to the sea.”
We gratefully recognize those thousands of sport and commercial fishermen, fisheries observers, research
vessel captains and crew, tournament officials, and cooperating scientists who have helped make this one of
the largest shark tagging programs in
the world; many thanks to all of you
for your dedication and support of
our program. Special acknowledgements to past and present Apex Predators staff, including Ruth Briggs, Lisa
J. Natanson, Cami McCandless, John
Hoey, Gregg Skomal, Chuck Stillwell,
H. Wes Pratt, Joe Mello, Mike Couturier, Fred Lerch, Larry Lindgren, Nancy Kelley, and Patricia Hadfield for
their assistance and support with data
collection and processing. We are also
grateful to Lisa J. Natanson and Richard McBride for editing the manuscript and helping with the overall
layout, and Kathy Duffy and Jon Hare
for providing valuable technical assistance. We especially thank Lisa J. Natanson for the final push over the finish
line.
15

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17

Atlantic Angel Shark
Sex
Male
Female
Unknown
Total

Tags

Recaptures

Recapture rate
(%)

74
74
22
170

0
0
0
0

0.0
0.0
0.0
0.0

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

Figure 9a.—Distribution of recapture locations for the Atlantic angel shark, Squatina dumeril, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

18

Marine Fisheries Review

Figure 9b.—Distribution of mark/recapture locations for the Atlantic angel shark, Squatina dumeril, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 9c.—Seasonal distribution of mark/recapture locations for the Atlantic angel shark, Squatina dumeril,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2)

19

Atlantic Sharpnose Shark
Sex
Male
Female
Unknown
Total

Tags
2,903
1,804
270
4,977

Recaptures
48
20
11
79

Recapture rate
(%)

Mean distance
traveled (nmi)

1.7
1.1
4.1
1.6

93.6
118.1
99.0
100.6

Maximum distance
traveled (nmi)
570
337
297
570

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.9
2.3
1.8
2.0

7.3
6.1
6.5
7.3

Figure 10a.—Distribution of recapture locations for the Atlantic sharpnose shark, Rhizoprionodon terraenovae, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 10b.—Distribution of mark/recapture locations for the Atlantic sharpnose shark, Rhizoprionodon
terraenovae, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 10c.—Seasonal distribution of mark/recapture locations for the Atlantic sharpnose shark, Rhizoprionodon terraenovae, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2)

21

Basking Shark
Sex
Male
Female
Unknown
Total

Tags

Recaptures

Recapture rate
(%)

46
31
91
168

0
0
0
0

0.0
0.0
0.0
0.0

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

Figure 11a.—Distribution of recapture locations for the basking shark, Cetorhinus maximus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 11b.—Distribution of mark/recapture locations for the basking shark, Cetorhinus maximus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 11c.—Seasonal distribution of mark/recapture locations for the basking shark, Cetorhinus maximus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 23

Bigeye Thresher

Sex
Tags
Recaptures
Male
Female
Unknown
Total

112
120
168
400

2
3
7
12

Recapture rate
(%)

Mean distance
traveled (nmi)

1.8
2.5
4.2
3.0

238.5
900.0
826.1
746.7

Maximum distance
traveled (nmi)
366
1,100
2,067
2,067

Mean time at
liberty (years)

Maximum time
at liberty (years)

5.0
6.3
5.6
5.7

5.7
10.5
9.6
10.5

Figure 12a.—Distribution of recapture locations for the bigeye thresher, Alopias superciliosus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 12b.—Distribution of mark/recapture locations for the bigeye thresher, Alopias superciliosus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 12c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the bigeye
thresher, Alopias superciliosus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 25

Bignose Shark
Sex
Male
Female
Unknown
Total

Tags
61
91
23
175

Recaptures
6
4
1
11

Recapture rate
(%)
9.8
4.4
4.3
6.3

Mean distance
traveled (nmi)
898.5
1,010.8
768.0
927.5

Maximum distance
traveled (nmi)
1,778
1,484
768
1,778

Mean time at
liberty (years)

Maximum time
at liberty (years)

6.5
5.5
2.0
5.7

10.9
11.2
2.0
11.2

Figure 13a.—Distribution of recapture locations for the bignose shark, Carcharhinus altimus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 13b.—Distribution of mark/recapture locations for the bignose shark, Carcharhinus altimus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 13c.—Seasonal distribution of mark/recapture locations for the bignose shark, Carcharhinus altimus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 27

Blacknose Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,340
1,551
67
2,958

17
17
1
35

Recapture rate
(%)
1.3
1.1
1.5
1.2

Mean distance
traveled (nmi)
66.6
35.6
8.0
49.9

Maximum distance
traveled (nmi)
226
196
8
226

Mean time at
liberty (years)

Maximum time
at liberty (years)

2.5
3.1

6.1
9.9

2.8

9.9

Figure 14a.—Distribution of recapture locations for the blacknose shark, Carcharhinus acronotus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 14b.—Distribution of mark/recapture locations for the blacknose shark, Carcharhinus acronotus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 14c.—Seasonal distribution of mark/recapture locations for the blacknose shark, Carcharhinus acronotus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 29

Blacktip Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

3,699
5,681
913
10,293

101
133
35
269

Recapture rate
(%)

Mean distance
traveled (nmi)

2.7
2.3
3.8
2.6

112.8
116.2
81.0
110.9

Maximum distance
traveled (nmi)
1,183
632
607
1,183

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.9
0.8
0.9
0.8

9.3
7.8
5.9
9.3

Figure 15a.—Distribution of recapture locations for the blacktip shark, Carcharhinus limbatus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 15b.—Distribution of mark/recapture locations for the blacktip shark, Carcharhinus limbatus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 15c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the blacktip
shark, Carcharhinus limbatus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 31

Blue Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

48,107
37,055
32,800
117,962

3,914
2,603
1,696
8,213

Recapture rate
(%)

Mean distance
traveled (nmi)

8.1
7.0
5.2
7.0

608.8
491.6
469.2
543.2

Maximum distance
traveled (nmi)
3,997
3,676
3,740
3,997

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.3
0.9
1.0
1.1

15.9
10.3
8.1
15.9

Figure 16a.—Distribution of recapture locations for the blue shark, Prionace glauca, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 16b.—Distribution of recapture locations for the blue shark, Prionace glauca, tagged outside the
U.S. EEZ and distance traveled greater than 1,000 nmi, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 16c.—Long distance recaptures (> 2000 nmi) for the blue shark, Prionace glauca, at liberty for
less than one year from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 33

Figure 16d.—Distribution of mark/recapture locations for the blue shark, Prionace glauca, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 16e.—Monthly seasonal distribution of mark/recapture locations in the western North Atlantic for the
blue shark, Prionace glauca, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 16e.—Continued

81(2) 35

Bonnethead

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,118
3,751
188
5,057

15
199
7
221

Recapture rate
(%)
1.3
5.3
3.7
4.4

Mean distance
traveled (nmi)
3.0
9.8
14.0
9.5

Maximum distance
traveled (nmi)
22
302
40
302

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.4
0.9
0.2
0.9

2.1
7.0
0.3
7.0

Figure 17a.—Distribution of recapture locations for the bonnethead, Sphyrna tiburo, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 17b.—Distribution of mark/recapture locations for the bonnethead, Sphyrna tiburo, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

Figure 17c.—Seasonal distribution of mark/recapture locations for the bonnethead, Sphyrna tiburo, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 37

Bull Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

702
1,167
260
2,129

11
20
5
36

Recapture rate
(%)

Mean distance
traveled (nmi)

1.6
1.7
1.9
1.7

201.6
125.4
122.5
149.0

Maximum distance
traveled (nmi)
428
628
349
628

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.2
0.8
1.0
1.0

6.7
4.8
1.9
6.7

Figure 18a.—Distribution of recapture locations for the bull shark, Carcharhinus leucas, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 18b.—Distribution of mark/recapture locations for the bull shark, Carcharhinus leucas, from the
NMFS Cooperative Shark Tagging Program (1962–2013)

Figure 18c.—Seasonal distribution of mark/recapture locations for the bull shark, Carcharhinus leucas, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 39

Common Thresher Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

63
60
80
203

2
0
2
4

Recapture rate
(%)

Mean distance
traveled (nmi)

3.2
0.0
2.5
2.0

71.0

168.5
119.8

Maximum distance
traveled (nmi)
86

271
271

Mean time at
liberty (years)

Maximum time
at liberty (years)

6.1

4.7
5.4

8.0
6.8
8.0

Figure 19a.—Distribution of recapture locations for the common thresher shark, Alopias vulpinus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 19b.—Distribution of mark/recapture locations for the common thresher shark, Alopias vulpinus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 19c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the common thresher shark, Alopias vulpinus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 41

Crocodile Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

9
10
1
20

0
0
0
0

Recapture rate
(%)
0.0
0.0
0.0
0.0

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

Figure 20a.—Distribution of recapture locations for the crocodile shark, Pseudocarcharias kamoharai,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 20b.—Distribution of mark/recapture locations for the crocodile shark, Pseudocarcharias kamoharai, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 20c.—Seasonal distribution of mark/recapture locations for the crocodile shark, Pseudocarcharias kamoharai, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 43

Dusky Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

2,146
3,723
2,596
8,465

49
72
43
164

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

2.3
1.9
1.7
1.9

607.7
583.5
548.4
581.9

2,015
2,017
2,052
2,052

2.8
3.6
3.0
3.2

11.8
16.1
15.8
16.1

Figure 21a.—Distribution of recapture locations for the dusky shark, Carcharhinus obscurus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 21b.—Distribution of mark/recapture locations for the dusky shark, Carcharhinus obscurus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 21c.—Seasonal distribution of mark/recapture locations for the dusky shark, Carcharhinus obscurus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 45

Finetooth Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,289
1,475
43
2,807

25
35
0
60

Recapture rate
(%)
1.9
2.4
0.0
2.1

Mean distance
traveled (nmi)
55.2
43.5

48.4

Maximum distance
traveled (nmi)
351
365

365

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.5
0.6

0.5

3.9
4.9
4.9

Figure 22a.—Distribution of recapture locations for the finetooth shark, Carcharhinus isodon, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 22b.—Distribution of mark/recapture locations for the finetooth shark, Carcharhinus isodon, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 22c.—Seasonal distribution of mark/recapture locations for the finetooth shark, Carcharhinus isodon,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 47

Galapagos Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

178
195
49
422

7
9
2
18

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

3.9
4.6
4.1
4.3

256.3
71.9
16.0
137.4

1,087
505
32
1,087

Mean time at
liberty (years)

Maximum time
at liberty (years)

3.0
1.7
3.0
2.3

7.1
3.8
4.2
7.1

Figure 23a.—Distribution of recapture locations for the Galapagos shark, Carcharhinus galapagensis,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 23b.—Distribution of mark/recapture locations for the Galapagos shark, Carcharhinus galapagensis, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 23c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the Galapagos shark, Carcharhinus galapagensis, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 49

Great Hammerhead

Sex
Tags
Recaptures
Male
Female
Unknown
Total

102
126
54
282

1
2
2
5

Recapture rate
(%)

Mean distance
traveled (nmi)

1.0
1.6
3.7
1.8

2.0
52.0
575.5
251.4

Maximum distance
traveled (nmi)
2
102
649
649

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.2
1.0
3.1
1.7

0.2
1.2
3.4
3.4

Figure 24a.—Distribution of recapture locations for the great hammerhead, Sphyrna mokarran, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Marine Fisheries Review

Figure 24b.—Distribution of mark/recapture locations for the great hammerhead, Sphyrna mokarran, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 24c.—Seasonal distribution of mark/recapture locations for the great hammerhead, Sphyrna mokarran, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 51

Greenland Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

32
36
0
68

1
0
0
1

Recapture rate
(%)
3.1
0.0
0.0
1.5

Mean distance
traveled (nmi)
0.0

0.0

Maximum distance
traveled (nmi)
0

0

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.0

1.0

1.0
1.0

Figure 25a.—Distribution of recapture locations for the Greenland shark, Somniosus microcephalus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 25b.—Distribution of mark/recapture locations for the Greenland shark, Somniosus microcephalus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 25c.—Seasonal distribution of mark/recapture locations for the Greenland shark, Somniosus microcephalus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 53

Lemon Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,348
1,599
284
3,231

147
113
17
277

Recapture rate
(%)
10.9
7.1
6.0
8.6

Mean distance
traveled (nmi)
45.0
19.7
37.4
34.1

Maximum distance
traveled (nmi)
494
239
286
494

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.2
0.8
1.5
1.1

10.9
5.1
5.2
10.9

Figure 26a.—Distribution of recapture locations for the lemon shark, Negaprion brevirostris, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 26b.—Distribution of mark/recapture locations for the lemon shark, Negaprion brevirostris, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 26c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the lemon
shark, Negaprion brevirostris, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 55

Longfin Mako

Sex
Tags
Recaptures
Male
Female
Unknown
Total

44
19
43
106

5
0
1
6

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

11.4
0.0
2.3
5.7

835.4

434.0
768.5

1852

434
1852

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.7

0.9
1.6

5.5
0.9
5.5

Figure 27a.—Distribution of recapture locations for the longfin mako, Isurus paucus, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 27b.—Distribution of mark/recapture locations for the longfin mako, Isurus paucus, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

Figure 27c.—Seasonal distribution of mark/recapture locations for the longfin mako, Isurus paucus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 57

Night Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

96
120
73
289

4
10
5
19

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

4.2
8.3
6.8
6.6

71.0
582.6
352.8
414.4

225
1,456
789
1,456

Mean time at
liberty (years)

Maximum time
at liberty (years)

8.6
4.0
3.9
4.8

13.8
12.9
8.8
13.8

Figure 28a.—Distribution of recapture locations for the night shark, Carcharhinus signatus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 28b.—Distribution of mark/recapture locations for the night shark, Carcharhinus signatus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 28c.—Seasonal distribution of mark/recapture locations for the night shark, Carcharhinus signatus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 59

Nurse Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,003
944
239
2,186

78
84
13
175

Recapture rate
(%)
7.8
8.9
5.4
8.0

Mean distance
traveled (nmi)
19.3
13.6
31.5
17.3

Maximum distance
traveled (nmi)
297
385
225
385

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.2
1.4
1.2
1.3

7.8
11.6
6.9
11.6

Figure 29a.—Distribution of recapture locations for the nurse shark, Ginglymostoma cirratum, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 29b.—Distribution of mark/recapture locations for the nurse shark, Ginglymostoma cirratum, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 29c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the nurse
shark, Ginglymostoma cirratum, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 61

Oceanic Whitetip Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

152
216
275
643

1
5
2
8

Recapture rate
(%)

Mean distance
traveled (nmi)

0.7
2.3
0.7
1.2

555.0
443.4
625.0
502.8

Maximum distance
traveled (nmi)
555
998
1,226
1,226

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.3
0.1
3.2
0.9

0.3
0.3
3.3
3.3

Figure 30a.—Distribution of recapture locations for the oceanic whitetip shark, Carcharhinus longimanus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 30b.—Distribution of mark/recapture locations for the oceanic whitetip shark, Carcharhinus
longimanus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 30c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the oceanic
whitetip shark, Carcharhinus longimanus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 63

Porbeagle

Sex
Tags
Recaptures
Male
Female
Unknown
Total

727
727
300
1,754

73
80
25
178

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

10.0
11.0
8.3
10.1

215.5
245.0
216.5
228.8

778
889
1,216
1,216

Mean time at
liberty (years)

Maximum time
at liberty (years)

3.0
3.1
3.7
3.1

11.0
16.8
8.0
16.8

Figure 31a.—Distribution of recapture locations for the porbeagle, Lamna nasus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 31b.—Distribution of mark/recapture locations for the porbeagle, Lamna nasus, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

Figure 31c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the porbeagle, Lamna nasus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 65

Reef Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

271
414
83
768

11
11
2
24

Recapture rate
(%)
4.1
2.7
2.4
3.1

Mean distance
traveled (nmi)
3.1
2.9
1.0
2.8

Maximum distance
traveled (nmi)
16
26
2
26

Mean time at
liberty (years)

Maximum time
at liberty (years)

3.0
1.0
0.1
1.9

9.2
2.4
0.2
9.2

Figure 32a.—Distribution of recapture locations for the reef shark, Carcharhinus perezii, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 32b.—Distribution of mark/recapture locations for the reef shark, Carcharhinus perezii, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 32c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the reef shark,
Carcharhinus perezii, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 67

Sandbar Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

12,470
19,363
4,096
35,929

545
784
145
1474

Recapture rate
(%)

Mean distance
traveled (nmi)

4.4
4.0
3.5
4.1

354.0
410.5
784.2
423.4

Maximum distance
traveled (nmi)
2,039
2,031
2,038
2,039

Mean time at
liberty (years)

Maximum time
at liberty (years)

2.8
3.0
5.0
3.1

24.9
26.9
27.8
27.8

Figure 33a.—Distribution of recapture locations for the sandbar shark, Carcharhinus plumbeus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 33b.—Distribution of mark/recapture locations for the sandbar shark, Carcharhinus plumbeus,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 33c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the sandbar
shark, Carcharhinus plumbeus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 69

Sand Tiger

Sex
Tags
Recaptures
Male
Female
Unknown
Total

885
817
317
2,019

31
29
13
73

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

3.5
3.5
4.1
3.6

153.3
141.5
98.1
138.5

641
637
267
641

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.3
1.0
1.4
1.2

5.3
2.3
4.1
5.38

Figure 34a.—Distribution of recapture locations for the sand tiger, Carcharias taurus, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 34b.—Distribution of mark/recapture locations for the sand tiger, Carcharias taurus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 34c.—Seasonal distribution of mark/recapture locations for the sand tiger, Carcharias taurus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 71

Scalloped Hammerhead

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,612
1,504
421
3,537

33
19
10
62

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

2.0
1.3
2.4
1.8

161.1
117.5
194.9
152.5

765
695
902
902

2.7
1.7
3.4
2.5

9.6
7.7
6.0
9.6

Figure 35a.—Distribution of recapture locations for the scalloped hammerhead, Sphyrna lewini, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 35b.—Distribution of mark/recapture locations for the scalloped hammerhead, Sphyrna lewini,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 35c.—Seasonal distribution of mark/recapture locations for the scalloped hammerhead, Sphyrna lewini, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 73

Shortfin Mako

Sex
Tags
Recaptures
Male
Female
Unknown
Total

1,917
3,251
3,357
8,525

373
491
284
1148

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

19.5
15.1
8.5
13.5

512.8
421.4
494.7
469.1

2,867
2,306
3,043
3,043

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.3
1.1
1.2
1.2

12.8
12.4
9.5
12.8

Figure 36a.—Distribution of recapture locations for the shortfin mako, Isurus oxyrinchus, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 36b.—Distribution of recapture locations for the shortfin mako, Isurus oxyrinchus, tagged outside
the U.S. EEZ from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 36c.—Long distance recaptures (> 1000 nmi) for the shortfin mako, Isurus oxyrinchus, at liberty for
less than one year from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 75

Figure 36d.—Distribution of mark/recapture locations for the shortfin mako, Isurus oxyrinchus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 36e.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the shortfin
mako, Isurus oxyrinchus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 36f.—Distribution of mark/recapture locations in the northeastern U.S. for the shortfin mako, Isurus oxyrinchus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 36g.—Distribution of mark/recapture locations in the southeastern U.S. for the shortfin mako, Isurus oxyrinchus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 77

Silky Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

340
464
434
1,238

23
21
21
65

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

6.8
4.5
4.8
5.3

204.0
255.6
132.4
197.6

718
1,288
539
1,288

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.1
0.9
0.7
0.9

6.0
8.6
7.1
8.6

Figure 37a.—Distribution of recapture locations for the silky shark, Carcharhinus falciformis, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 37b.—Distribution of mark/recapture locations for the silky shark, Carcharhinus falciformis, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 37c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for silky shark,
Carcharhinus falciformis, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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79

Smalltail Shark
Sex
Male
Female
Unknown
Total

Tags
8
14
2
24

Recaptures

Recapture rate
(%)

0
0
0
0

0.0
0.0
0.0
0.0

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

Figure 38a.—Distribution of recapture locations for the smalltail shark, Carcharhinus porosus, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 38b.—Distribution of mark/recapture locations for the smalltail shark, Carcharhinus porosus, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 38c.—Seasonal distribution of mark/recapture locations for the smalltail shark, Carcharhinus porosus, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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81

Smooth Dogfish

Sex
Tags
Recaptures
Male
Female
Unknown
Total

263
842
81
1,186

5
29
3
37

Recapture rate
(%)

Mean distance
traveled (nmi)

1.9
3.4
3.7
3.1

147.8
114.2
140.7
120.9

Maximum distance
traveled (nmi)
379
460
152
460

Mean time at
liberty (years)

Maximum time
at liberty (years)

2.1
2.0
0.4
1.9

4.1
6.8
0.8
6.8

Figure 39a.—Distribution of recapture locations for the smooth dogfish, Mustelus canis, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 39b.—Distribution of mark/recapture locations for the smooth dogfish, Mustelus canis, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 39c.—Seasonal distribution of mark/recapture locations for the smooth dogfish, Mustelus canis, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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83

Smooth Hammerhead

Sex
Tags
Recaptures
Male
Female
Unknown
Total

112
121
36
269

2
3
2
7

Recapture rate
(%)

Mean distance
traveled (nmi)

1.8
2.5
5.6
2.6

289.0
61.3
119.5
143.0

Maximum distance
traveled (nmi)
496
178
184
496

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.3
0.8
0.9
1.0

2.1
1.3
1.7
2.1

Figure 40a.—Distribution of recapture locations for the smooth hammerhead, Sphyrna zygaena, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 40b.—Distribution of mark/recapture locations for the smooth hammerhead, Sphyrna zygaena,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 40c.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the smooth
hammerhead, Sphyrna zygaena, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 85

Spinner Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

670
936
117
1,723

9
15
3
27

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.3
1.6
2.6
1.6

170.2
84.1
23.3
106.0

861
312
36
861

1.1
0.8
0.7
0.9

6.8
4.5
1.9
6.8

Figure 41a.—Distribution of recapture locations for the spinner shark, Carcharhinus brevipinna, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 41b.—Distribution of mark/recapture locations for the spinner shark, Carcharhinus brevipinna,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 41c.—Seasonal distribution of mark/recapture locations for the spinner shark, Carcharhinus brevipinna, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 87

Tiger Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

3,800
4,919
1,053
9,772

302
363
44
709

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

7.9
7.4
4.2
7.3

219.1
251.7
512.6
252.6

3,018
3,643
3,089
3,643

Mean time at
liberty (years)

Maximum time
at liberty (years)

0.9
1.0
1.3
1.0

11.2
9.7
7.5
11.2

Figure 42a.—Distribution of recapture locations for the tiger shark, Galeocerdo cuvier, from the NMFS
Cooperative Shark Tagging Program (1962–2013).

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Figure 42b.—Distribution of recapture locations in the western North Atlantic for the tiger shark, Galeocerdo cuvier, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 42c.—Long distance recaptures (> 500 nmi) for the tiger shark, Galeocerdo cuvier, at liberty for less
than one year from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 89

Figure 42d.—Distribution of mark/recapture locations for the tiger shark, Galeocerdo cuvier, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 42e.—Seasonal distribution of mark/recapture locations in the western North Atlantic for the tiger shark, Galeocerdo cuvier, from the NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 42f.—Distribution of mark/recapture locations in the northeastern U.S. for the tiger shark, Galeocerdo cuvier, from the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 42g.—Distribution of mark/recapture locations in the southeastern U.S. for the tiger shark, Galeocerdo cuvier, from the NMFS Cooperative Shark Tagging Program (1962–2013).

81(2) 91

White Shark

Sex
Tags
Recaptures
Male
Female
Unknown
Total

12
27
16
55

0
2
0
2

Recapture rate
(%)

Mean distance
traveled (nmi)

Maximum distance
traveled (nmi)

0.0
7.4
0.0
3.6

465.0

465.0

546

546

Mean time at
liberty (years)

Maximum time
at liberty (years)

1.9

1.9

2.5
2.5

Figure 43a.—Distribution of recapture locations for the white shark, Carcharodon carcharias, from the
NMFS Cooperative Shark Tagging Program (1962–2013).

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Figure 43b.—Distribution of mark/recapture locations for the white shark, Carcharodon carcharias, from
the NMFS Cooperative Shark Tagging Program (1962–2013).

Figure 43c.—Seasonal distribution of mark/recapture locations for the white shark, Carcharodon carcharias,
from the NMFS Cooperative Shark Tagging Program (1962–2013).

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93