ELSEVIER Fisheries Research 29(1997) 101-I 17

Mini review

Status of shark resources in the Southern Gulf of Mexico and

Caribbean: implications for management
Ramh Bonfil *
Fisheries Centre. Uniurrsity oj’Brttish Columhiu, 2204 Main Mull, Vuncouuer. B.C. V6T 124. Gnu&~
Institute Nuciond de lo Prscu, Apdo. Postul 73, Pro~rrso. Yuc, Mexico 97320

Accepted 2 August 1996

Abstract

This paper reviews current knowledge on shark biology, ecology, and fisheries in Mexican waters of the Gulf of Mexico,
focusing on management implications. Original and literature data are summarized covering distribution, migrations, nursery
areas, structure of the landings and catch trends. There are at least 34 species of shark in the area, most of them from the
genera Cat&whims and Sphyma. Fourteen of these species are important in the fisheries. The biology and ecology of
shark stocks in the southern Gulf of Mexico are poorly known. Tagging studies have shown movements of shark from the
eastern US coast to the southern Gulf of Mexico, but migrations have not been properly characterized due to limited tagging
in Gulf waters. There are important shark nurseries for at least five species in the protected waters of Chetumal Bay (Q.
Roo), Yalahau Lagoon (Q. Roe), and Terminos Lagoon (Camp.). Sharks contribute approximately 3.5% of the total fishery
catches in this region, averaging about 13 000 t during I983- 1992. Sharks are harvested by a diverse fleet of small-scale and
large-scale vessels utilizing longlines or gillnets. There seems to be an excessive fishing of juveniles from many of the most
important species. Areas of priority for future research that are pre-requisite for international management are: the life cycle
of the main species in the southern Gulf of Mexico; in-depth study of coastal nursery areas; large scale tagging programmes
coupled with population genetics studies; gathering of fisheries data and fisheries-independent abundance information; and
socio-economic studies of the fisheries. Some precautionary management measures are proposed.

Ktywrds: Sharks; Biology; Ecology; Fisheries management; Overview; Southern Gulf of Mexico

1. Introduction imply that in every case these fisheries should be

The limiting biological characteristics of elasmo-
branchs and the historical failure of shark fisheries
(see Holden, 1977 and Anderson, 1990 for reviews)
= Tel: (604) 822-0618; fax: (604) 822-8934;
ology.ubc.ca.
e-mail: boniil@zo-
carefully managed. This is certainly true for the Gulf
of Mexico, where in the span of 10 years Mexican
and US shark catches rapidly increased more than
threefold, from ca 5500 t in 1980 to nearly 18 000 t
in 1989. These unprecedented levels of exploitation
in the Gulf of Mexico have raised concerns over
possible depletion of shark stocks in the near future
unless adequate and careful management is imple-
mented soon. A collapse of shark stocks in the Gulf

0165-7836/97/S17.00 Copyright 0 1997 Elsevier Science B.V. All rights reserved.

PI1 SO16S-7836(96)00536-X
102 R. Bonjl/Fisheries
of Mexico would jeopardise the fisheries and entail a
significant economic and social cost. There would be
lost catches and lost livelihoods for hundreds of
families in the region. In addition, a collapse of these
apex predators could trigger ecological changes of
unknown dimension (e.g. see van der Elst, 1979).
Shark resources in US waters have received a
reasonable amount of attention. Many biological
studies, including several species of importance in
the fisheries, were made in the early 1980s (see
Prince and Pulos, 1983; Pratt et al., 1990). In the
eastern US, sharks are currently under a management
plan that has brought under control the rapidly in-
creasing exploitation observed there in the late 1980s
(NOAA, 1995).
In contrast, there are very few studies in the
Mexican part of the Gulf of Mexico and no manage-
ment of sharks has been implemented there. Reasons
Fig. 1. Southern Gulf of Mexico, showing the main features mentioned
Reseurch 29 (1997) 101-117
for this are, among others, the perceived healthy
state of the Mexican fishery given the relatively
stable landings during the last ten years, and the lack
of adequate knowledge of the shark stocks and the
characteristics and problems of the fisheries.
Early Mexican research efforts on sharks and their
fisheries were mainly limited to gathering baseline
information about the resources. These efforts yielded
much needed faunistic lists and species catalogues,
sometimes including additional information on
species distribution or their incidence in fisheries
(Castro-Aguirre, 1967; Castro-Aguirre, 1978;
Castro-Aguirre, 1983; Applegate et al., 19791, as
well as details about their utilization (Hemandez-
Carvallo, 197 1). Early research also produced mono-
graphs on fishing technology (Marin, 1964; Castro-
Aguirre, 1965; Hemlndez et al., 1977). However,
none of the above studies provided an adequate
Gulf of Mexico
in the text.
 R. Bonjl/ Fisheries
description of the fisheries nor did they give any
insight into the assessment or management of shark
stocks, This was a consequence of the low level of
shark catches in Mexico before the 1970% which
drove most of the research towards developing the
fisheries as there were no evident needs for regula-
tion (Hemandez-Carvallo, 1971). In the early 1980s
the focus of Mexican research on sharks switched to
their fisheries biology and population dynamics, with
a clear aim on future management. The main thrust
of fisheries biology research came from the works of
scientists at laboratories of the Mexican National
Fisheries Institute (INP) scattered around the south
em Gulf of Mexico (Bonfil et al., 19881, and was
later supplemented by various theses from Universi-
ties in coastal States.
There are presently no coordinated efforts for
shark fisheries management between Mexico and the
USA. However, it is likely that joint management
will probably not proceed while the shark stocks and
the management needs in the southern Gulf of Mex-
ico remain poorly understood.
This paper provides a comprehensive review of
the status of Mexican shark fisheries in the Gulf of
Mexico and the Caribbean. It summarizes present
knowledge on fisheries biology of the species under
exploitation and the fisheries they sustain, stressing
the more evident problems and providing guidance
for immediate research and management actions. It is
hoped that this work will serve as a basis for man-
agers and fisheries scientists to prioritize future re-
search and foster the sound and equitable manage-
ment of sharks in the whole Gulf of Mexico. For the
remaining of the paper, the term ‘southern Gulf of
Mexico’ will be used to refer to the Mexican waters
of the Gulf of Mexico (Fig. I).
2. The fisheries
Mexican shark fisheries are very heterogeneous.
The type of vessels and gears used for shark fishing
vary regionally, as well as the seasonality of harvest
and degree of utilization of the different species.
However, an integrated in-depth study of these as-
pects of the fishery is still lacking for the southern
Gulf of Mexico. Unfortunately there is no formal and
complete record of the types and number of vessels
Rrseurch 29 (1997) 101-I I7 103
and gears involved in this fishery and this severely
hampers the monitoring of effort. There are no offi-
cial effort records, making it impossible to discrimi-
nate whether fluctuations in the catches are due to
changes in shark abundance or availability, or due to
variations in the effort regime.
Mexican shark fisheries include full time and part
time shark fishermen, and directed fisheries for sharks
as well as incidental catches of sharks in other
fisheries. The latter kind includes Mayan octopus
(Octopus maya) and red grouper (Epinephelus
murio) fisheries in Yucatan/Campeche (Bonfil et
al., 1990); king mackerel (Scomberomorus caualla)
in Tamaulipas and Veracruz; and tuna fisheries in all
the southern Gulf of Mexico. However, there is very
poor understanding of the degree of dependence of
the different fishing communities on sharks because
no socio-economic studies exist for these fisheries.
2.1. Vessels and gears
The directed shark fisheries are scattered through-
out the coastline and are mainly composed of small-
scale fleets using outboard-motored boats. These
vessels are generally made of fibreglass with the
exception of a few old wooden boats, and they range
from 7.5 to 12 m in length. Most of them are capable
of only l-3 day trips, with travelling distances deter-
mined by the width of the continental shelf. There
are, however, several medium or large scale vessels
with inboard diesel motors that also direct effort
towards sharks in several States. Vessels of lo-20 m
Fig. 2. Shark catches in the southern Gulf of Mexico 1976- 1992.
104 R. Bonjl/ Fisheries Research 29 (1997) 101-I I7

in length are built of fibreglass, wood, steel or tonnes/year

1976-1992
0 500 lcm 15co 2000 2500 3000 3500
ferro-cement, and can stay for up to 15 days at sea.
The fishing gears used to catch sharks depend Tamaullpas

largely on regional preferences. Various types of

longlines are used mainly in Veracruz and Tamauli-
pas each with 50-150 hooks for small vessels and Tabasco

200-400 hooks for larger vessels. Only a few ves- Campeche

sels use gillnets, which usually have meshes of 13
cm (Montiel, 1988; Marin, 1992). Campeche and
Yucatan fishermen prefer gillnets of mesh sizes rang-
ing from 11 to 40 cm (Uribe, 1984, Seca and Murillo,
1985; Bonfil et al., 1990). Fishermen in Quintana
Roo use both longlines and gillnets (Faustch, 1986).
The fishing of sharks with hand-held harpoons from
sail-powered vessels was abandoned since the early
1970s in the area of Holbox, Quintana Roo.
2.2. Catch patterns and trends’
Mexican catches of sharks in the Gulf of Mexico
have remained relatively stable after the rapid growth
of the early 1980s. Data collected and published by
the Federal Government indicates catches oscillating
around 13 000 t year-’ during 1983-1992 (Fig. 2).
Based on the last 10 years in record, the shark
fishery seems to be in a state of stability with no
increasing trends or problems of declines in the
catches. Although no fishery statistics in the world
Fig. 3. Catches of large and small sharks (see text) by State in the
southern Gulf of Mexico, 1976- 1992.
are 100% accurate, Mexican statistics are thought to
reflect well the trends in the fishery.
Shark fishing is a widespread activity along the
Mexican coast of the Gulf of Mexico (Table 1).
According to mean catches during 1976-1992, Ver-
acruz takes the largest share of the fishery (29%)
followed by Campeche (26%), Tamaulipas (18%)
and Yucatan (16%), with the States of Tabasco and
Quintana Roo contributing only minor catches (8%
and 3% respectively). Shark fisheries are locally of
considerable importance in the southern Gulf of
Mexico. They contributed an average of 3.5% to the
total catches during 1977-1992, significantly more

Table 1
Mexican shark catches by State (tonnes)

Year Tamaulipas Veracruz Tabasco Campeche Yucatan Q. Roo Total

1976 341 708 261 1095 385 239 3029

1977 730 844 547 1361 405 95 3982
1978 572 1025 513 1414 463 107 4094
1979 936 1365 421 1069 182 79 4052
1980 1260 2057 456 882 437 124 5216
1981 3189 4712 594 1199 770 295 10759
1982 1330 4770 540 1553 2093 331 10617
1983 1442 4212 685 3754 2351 385 12829
1984 1679 5623 870 3847 205 1 433 14503
1985 1923 4761 1943 4388 2662 559 16236
1986 1754 2926 1250 2928 225 1 358 11467
1987 2077 2202 1136 4061 1542 256 11274
1988 2375 3121 849 3936 1626 277 12184
1989 2734 3642 742 3397 1912 330 12757
1990 3150 3082 1047 3211 2925 437 13852
1991 2243 2807 1203 2805 2383 288 11729
1992 2529 2958 1160 4577 2417 335 13976
 R. Bonjil/ Fisheries Research 29 (1997) 101-117 105

than the world average of 0.7% reported by Bonfil 19941, shark catches are not broken down into species
(1994). In Yucatan, sharks are the fourth most im- largely due to a lack of species-specific markets.
portant fishery resource (Bonfil et al., 1990). This is a strong obstacle for the management of
In Mexico, as in many other countries (Bonfil, shark fisheries. Official statistics recognize only two

Table 2
Distribution and relative abundance of sharks recorded on Mexican commercial fisheries of the Southern Gulf of Mexico and Carribean
Family Species Tamaulipas Veracnrz Tabasco Campeche Yucatan Q Roo
(1780 t (2989 t (836 t (2675 t (1580 t (290 t
year- ’ 1 year- ‘1 year- ‘) year- ‘) year- ‘1 year- ‘1

Hexanchidae 1 Heptranchius perlo X

2 Hexunchus griseus X X
3 Hexanchus vitulus X 7
X X X
Squalidae 4 Centrophorus grunulosus X XC
5 Centrophorus uyuto X
6 Squalus cuhensis X X
I Squalus mitsukurii XC
Squatinidae 8 Squutinu dumeril X
Ginglymostomatidae 9 Ginglymostoma cirrrrtum a X X ? X xx xxx
Rbincodontidae 10 Rhincodon typus X X
Odontaspididae 1 1 Odontaspis ferox X
Alopiidae 12 Alopicrs superciliosu., X
13 Alopius vulpinus X
Lamnidae 14 I.turus oxyrinchus X X 3
X X X
Triakidae 15 Mustelus cunis b xx xx ? X xxx X
16 Mustelus sp. ? ‘>

Carcharhinidae 17 Carcharhinus crcronotus a xx 3

xx xx X
18 Curchurhinus altimus X X X X
‘7
I9 Curchurhinus brachyurus ?
20 Curchurhinus hrevipinnu X X ? X xx X
2 I Carchurhinus isodon xx
22 Curcharhinus jdciformis b xx xxx ? X xxx xx
23 Curchnrhinus leucus b xxx xxx ? xxx xxx xxx
24 Curchurhinus limhutus b X xxx X xxx xx X
25 Curchurhinus longimanus X X ? ? XC
26 Carchurhinus obscurus b xxx xx ? X xxx X
27 Carchurhinus perezi xx X
28 Curchurhinus plumheus b xxx xxx X X xx X
29 Curchurhinus porosu~ X X
30 Curchurhinus signutus X xx ? ? X
3 1 Goleocerdo cuvier a xx X ? X xx xx
32 Negoprion breuirostris X ? X X xx
33 Rhizoprionodon X xxx 3
xxx xxx xx
terruenovae b
Sphymidae 34 Sphymu lewini b xx xx ? xx xx X
35 Sphyrnu mokarrcm a X X 3
xx xx xx
36 Sphyrna tihuro b X ? xxx xxx X

Average shark catches in parenthesis. (XXX: very common; XX: common; X: rare; ?: unconfirmed report). No superscript letter: no
importance.
a Secondary importance.
b Prime importance.
’ Bonfil, unpublished data.
Source: Backus et al., 1956; Casey et al., 1977- 1993; Applegate et al., 1979; Applegate et al., 1984; Uribe, 1984; Uribe, 1986; Faustch,
1986; Seca and Murillo, 1985; Hemandez, 1987; Montiel, 1988; Marin, 1992; Bonfil, 1995; Bonfil et al., 1990.
106 R. Bonjil/Fisheries Research 29 (1997) 101-117

Campcha

Lu i

L-d L- LadL-
!_-&iuu
1,.
L- (

Lhut ldd l.J.u

luu t bmu
Ld Ll,r R
L- I

1111 i

1n , LEU
i
BMB

uu B.u
i

t!-IL k
 R. Bonjil/ Fisheries Rrteurch
categories: large sharks measuring more than 150 cm
TL,, and ‘cazones’ or small sharks under the 150 cm
TL mark. The latter category comprises small shark
species and the juveniles of larger sharks.
Overall, small sharks averaged about 44% of the
total shark catches of the southern Gulf since 1976,
while the remaining 56% were large sharks. How-
ever, there is some variation in the type of sharks
taken in each State. The fisheries in Tamaulipas
seem to harvest mainly small sharks, whilst in Yu-
catan, Quintana Roo and to a lesser extent in Ver-
acruz, the fisheries seem to concentrate on large
sharks. Campeche and Tabasco catch roughly equal
tonnages of large and small sharks. However, the
absolute catches of small sharks taken in some States
are a cause of concern. Tamaulipas, Veracruz and
Campeche all have catches of small sharks of more
than 1140 t year-’ for the recorded period (Fig. 3).
At least in Campeche, these catches include large
amounts of juveniles of larger species (Seca and
Murillo, 198.5).
2.3. Biodiuersity of sharks in the southern Gulf of
Me.rico
The sharks of the southern Gulf of Mexico in-
clude at least 34 species from 11 families (Table 21,
representing about 10% of the described world shark
fauna. Fourteen shark species are identified here as
the most common in the commercial catches of the
southern Gulf of Mexico, based on the reports of
Uribe (1984); Seca and Murillo (1985); Faustch
(1986); Hemandez (1987); Bonfil et al. (1988); Bon-
fil et al. (1990); Montiel (19881, and Marin (1992).
Sharks of the Families Carcharhinidae and
Sphyrnidae are dominant, as expected in tropical
waters.
A rapid qualitative analysis of the information on
occurrence and relative frequency of the species in
the catches of each State (Table 2) shows that nine
of the 14 main species are of prime importance
Fig. 4. Structure of shark catches in the coastal States of the southern Gulf of
pre-adult; A: adult) are plotted in the x axis for each case according to their
present in the catches; large bars, common in the catches. The total absence
always imply that the species is not present, but that information might not
according to their latitudinal gradient along the coast, from north to south.
29 (19971 101-I 17 107
(these species are common in the four States with
highest shark catches, or very common in at least
one of these States). An additional group of five
species are of secondary importance (these species
are present in the catches of several States and are
common in at least one of the four States with
highest shark catches). Research and management
priorities in the southern Gulf of Mexico should
focus first on the nine species of prime importance.
2.4. Structure of the catches
Data collected during the mid and late 1980s
(Uribe, 1984; Uribe, 1986; Seca and Murillo, 1985;
Faustch. 1986; Hemandez, 1987; Bonfil et al., 1990;
Marin, 1992) provide snap-shot information that al-
lows a rough qualitative description of the demo-
graphic structure of the catches for the main species
in the shark fisheries. For this purpose, four life
stages were identified for each species (newborns,
juveniles, pre-adults and adults) and their importance
in the different fisheries was classified as absent,
present or common (Fig. 4). It is important to note
that these are partial data taken for a period of l-3
years in each site, and that the lack of information on
a species for a certain location does not necessarily
indicate its absence there, but possibly that it has not
yet been reported, or that detailed information on the
catches for that species is not readily available.
Despite these shortcomings, it is possible to iden-
tify a few important features of shark exploitation in
the southern Gulf of Mexico. The newborn and/or
juveniles of at least nine species (blacknose shark,
Carcharhinus acronotus; silky shark, Carcharhinus
falciformis: bull shark, Carcharhinus leucas; black-
tip shark, Carcharhinus limbatus; tiger shark, Galeo-
cerdo cuvier; Atlantic sharpnose shark, Rhizoprion-
odon terraenouae; and the three hammerhead sharks
of the genus Sphyrna) are prominent in the catches
of one or more coastal States. In the case of silky
sharks, while the fisheries take all life stages
Mexico. Four different life stages (N: newborn: J: juvenile; P:
relative abundance: no bars, absent in the catches; short bars,
of information for a species on a particular location does not
be yet available for that area. States are arranged in columns
108 R. Bonjl/Fisheries
throughout its range, the juveniles are dominant in
the catches. In contrast to this, catches of dusky
smooth-hounds (Mustelus canis), dusky sharks (C.
obscurus), and sandbar sharks (C. plum&us) are
almost exclusively comprised of adults and sub-
adults. The catches of bull sharks in the southern
Gulf of Mexico, while including all life stages, are
concentrated on adults and sub-adults except in
Quintana Roo where newborns are also heavily ex-
ploited in nursery areas. There are worrying reports
of large catches of newborn and juveniles of Atlantic
sharpnose shark, silky shark, blacknose shark, and
bonnethead shark (Sphyma riburo) in the Campeche
Bank (Seca and Murillo, 1985; Bonfil, 1990). Simi-
lar concerns have been raised for the heavy exploita-
tion of blacktip, blacknose, bull, coral reef
(Carcharhinus perezi), and lemon sharks (Negup-
rion breuirostris) in Yalahau Lagoon (R. Cruz, Na-
tional Fisheries Institute, pers. comm. August 1985).
3. Ecology
3.1. Migrations and stock identification
Large sharks are known to be strong swimmers
and some species can cross entire oceans several
times during their life span (e.g. blue sharks, Nakano,
1994). While some of them might appear simply to
roam the oceans as they grow, others are known to
display well defined seasonal patterns of migration
(e.g. mako sharks, Zsurus oxyrinchus, Casey and
Kohler, 1992). Defining the number of shark stocks,
their size and geographical boundaries is a key as-
pect for fisheries management. Moreover, the deter-
mination of the timing and routes of their migrations,
and most importantly the rates of movement between
stocks, must also be addressed before management
can be properly implemented. Unfortunately, the state
of knowledge regarding these key aspects for shark
management in the Gulf of Mexico is not encourag-
ing.
To date, there are no published genetic studies of
stock identification for sharks in the Gulf of Mexico.
Given the low genetic variation found in many shark
species (Smith, 1986) this type of approach will
probably need to rely on carefully selected tech-
niques and very well planned surveys that include
Research 29 (1997) 101-117
representative samples covering the entire range of
each species in the NW Atlantic. A lot of work and
cooperation among nations in this area will be essen-
tial for success.
Regarding shark migrations, the available infor-
mation presents a biased picture with much more
data from tagging programmes in the USA than from
Mexican programmes. In the southern Gulf of Mex-
ico, very limited tagging studies have been carried
out in Yucatan and Quintana Roo waters by the
National Fisheries Institute, with support from the
Cooperative Shark Tagging Programme of
NOAA/NMFS (Bonfil et al., 1988). So far, four out
of 15 sharks of six species tagged in four field
campaigns, have been recovered in this programme,
suggesting a relatively high exploitation rate in this
fishery (Fig. 5b). These results indicate a slow coastal
westward movement for a juvenile nurse shark
(Ginglymostoma cirrutum), movements of juvenile
silky sharks inside the Campeche Bank, and an
outside movement up to the coast of western Florida.
This last movement is the first report for a shark
travelling from Mexican waters into the northern
Gulf of Mexico, and highlights the need for in-
creased tagging in this region.
In contrast with the above, information concem-
ing sharks tagged in the US and recovered in the
southern Gulf of Mexico is much more abundant.
Most of this information comes from the Coopera-
tive Shark Tagging Program in the US, whose activi-
ties span for over 25 years and include more than
100000 sharks tagged so far (Casey et al., 1993).
Sandbar and dusky sharks are the most common
species recovered in Mexican waters but an addi-
tional eight species have also been identified (Fig.
5c-d).
Although the occurrence of shark tag recoveries
in Mexican waters might appear high from Fig.
5c-d, the data do not suggest a significant and
seasonal migration of sharks into the southern Gulf.
Recaptures in Mexican waters represent only a very
small fraction of the total recoveries in the western
Atlantic, even for the species most frequently recov-
ered in the southern Gulf of Mexico, the sandbar
shark (approximately 37 Mexican recaptures up until
1992, out of a total of 727 recaptures reported by
Casey et al. (1995)). The limited movement of sand-
bar sharks into the southern Gulf is more evident
 yJeqs a@u!s e JO ayle3!pu! aJr!pua JO u!9uo aql iv SJaqtunulnoqly SMOJJB :(SM~JJB~0 peaq JIZT~U)~~J~AO~~J JO (SMOU~
Jo U!%!JO) pa%%
Jaql!aaJaM l~qls~Jt?qsJOJaqlunu aql alexpu! (p)pw sJaqwnu aqJ_.aJnPg aqlJ0 pueq lqZ!Jaql uo paisy sa!zads aqj 01 puodsauoo ‘(p)
(3)u! SawI 01 pale!zossc!
SJaWM sn U! pa%?, SqJlZqS
JO SlUXIJaAOmJO UOl,E,UaSaJdaJ
3!lkXUaq3S
(p) puk? (3) !S~UXU~AO~UXNElS!p 1SalJOqSpaS!lNUaq3Sa
ou!moqS sas!w:,q3JaasaJuE3!xaw NJ!p?%%el sz$N?qs
Jo Sa!JWOCWj (9) :SgJI?qS UJOqMaU JO a3UWl33~ at,) LCq p3,!23!pU! SE SWJE i(J3SJnN (“) ‘03!XaW JO J[nf) UJaqlnOS al,] JO S.yJeqS JOJ UO!lWLIOjU!lW!%O[O32 JO ,&JeU,,UnS
's ‘3~~1
3
110 R. Bonjil/Fisherie.s
when taking into consideration that these are the
accumulated records of over 25 years of tagging, and
that they sometimes involve recoveries after up to 17
years at liberty. More importantly, the estimation of
the rates of movement of sharks between different
parts of the Gulf of Mexico, and between these and
the US Atlantic coast has not been addressed yet.
With the limited evidence available, there is room
only for speculation about the rate of movement of
sharks between the northern Gulf-eastern US and the
southern Gulf of Mexico.
3.2. Habitat
Although many fish species exhibit ontogenetic
changes in life style and some sharks are known to
range over several marine environments, it is possi-
ble to group the main shark species of the southern
Gulf of Mexico into at least four types roughly based
on their adult-life habitat. This strategy allows to
tackle the complexity of the fishery by systematizing
the species into ‘functional’ groups that better reflect
the particular subsets of the fishery and which could
be used as the basis for management. Coastal-demer-
sal species are the bonnethead shark, the Atlantic
sharpnose shark, the nurse shark, the blacknose shark,
and the dusky smooth-hound. The bull shark and the
sandbar shark form the demersal-pelagic coastal
group. Coastal pelagic species are the blacktip shark,
and the spinner shark (C. breuipinna). Finally, the
coastal-oceanic group is comprised of the silky shark,
the dusky shark, the tiger shark, the scalloped ham-
merhead (Sphyrna lewini), and the great hammer-
head shark (S. mokarran).
3.3. Nursery grounds
The protection of nursery areas is in many in-
stances a key aspect of fisheries management. This is
particularly true for tropical sharks because many
species use coastal areas for pupping and as nursery
grounds. Unfortunately, the utilization of coastal ar-
eas as nurseries by tropical and temperate sharks
exposes their most critical life stages (newborns and
early juveniles) to the potential threats of human
interaction. This is certainly the case for many species
of sharks in the southern Gulf of Mexico, where as
mentioned above (see structure of the catches), sev-
Research 29 (1997) 101-117
eral cases of heavy exploitation of newborns and
juveniles have been reported.
Fig. 5a summarizes the know occurrence of new-
born sharks and the major areas so far identified as
nurseries for sharks of commercial importance in the
southern Gulf of Mexico. There are records of at
least five species using coastal lagoons as nurseries
(Castro-Aguirre, 1978; Applegate et al., 1984;
Faustch, 1986). Chetumal Bay, Yalahau Lagoon, and
Terminos Lagoon, located in the southeast, north-
west and southwest of the Yucatan peninsula respec-
tively (Fig. 11, are of prime importance as shark
nurseries. Six additional shark species seem to have
nurseries in the Campeche Bank (Uribe, 1984; Seca
and Murillo, 1985; Bonfil, 1987; Hemandez, 1987;
Bonfil et al., 1990) and the coastal waters of
Tamaulipas and Veracruz (Montiel, 1988; Marin,
1992). It is possible that the sandbar shark and
perhaps also the dusky shark have nurseries in Mexi-
can waters that have not yet been identified due to
limited research on the subject. However, there is no
detailed information about the seasonality, abun-
dance, time of residence, etc., of newborn and juve-
nile sharks in these nursery areas.
4. Biology
There are very few Mexican studies on the biol-
ogy of sharks of the Gulf of Mexico. Table 3 shows
the biological parameters most relevant for the as-
sessment and management of the main species, as
reported in the literature. Noticeably, this compila-
tion is almost totally composed of studies pertaining
to stocks other than those inhabiting the southern
Gulf. In fact most of the information comes from
populations from the northern Gulf of Mexico, the
eastern US coast, or other parts of the world. This
information could be used to implement conservative
policies according to the precautionary principle of
fisheries management. It can also be useful as refer-
ence and comparison for the biological studies that
must be carried out in the southern Gulf of Mexico,
but it cannot substitute parameters derived specifi-
cally for these stocks. This is especially true in the
light of recent evidence that allopatric shark popula-
tions can have different population parameters (Par-
sons, 1993a; Parsons, 1993b).
Table 3
Life hetory paramcrres of the 14 main shatk spectes I” Mex~ca” commercral fisheries I” the Gulf of Mexico. (Mextcan sudes asterIsked>. I” the case of long-rangtng specjes for which no Mexican dam CXISL.
infommtion is pwen from the nearest studted populauons

Species Common name TLmxx K Length at Litter Gestation B,nh MaI”

(cm) malur~ty (cm) six (months) size (cm) referencrs
Spanish (MCx.)(YucatB”) English (FAO)

Rhi:o,,,_,o,rodon ,erroe,,rxae C&n de Ley Atlantx Sharpnose II0 j 0.359 85 f 32 Parsons (1983):
Shark Parsons (1985):
(Tutrun) 80 m Branstetter (1987a)
Caz6n Cabeza de Pala Bonnethead Shark II7 f 0.34-0.37 f SO-95 f 9-12 2s. 40 Parsons (1987):
(Chara) 93 m 0.53~0.58 m 68-80 m Parsons (1993.x):
Parsons (1993b):
Clark and van Schmidt (1965):
Alvarez(1988)*
Car6n Mamdn Dusky Smooth- 152 0.296 H 90 f 20 39 Francis (I 98 I ): Pratt and Casey f 1990):
hound Compagno (I 984)
82 m
Corchrrrhrnu, o<ronr,,ur Caz6n Nariz Negra Blacknosr Shark 164 0.138 f 101-103 f 3 6 45.50 Clark and vo” Schmidt (1965).
(Ganguay) 0.117 m Garr,ck (1982):
Schwartz (1984)
Currhorhinur lrmharur Tiburd” Puntas Negras Blacktip Shark 212 0.197-0.276 158-162 f IO 60 Clark and van Schmtdt (1965):
133-136 m K,llam and Parsons (1989):
Branstetter(l987b)
Corchorhrnur hrwrp,nno Tibur6” Maromero Spinner Shark 225 0.2 I2 180 f 6-8 60-70 Brsnstetter (I 98 I ):
170 m Branstetter (1987b)
Corchorhmus ,,lumhru.r Tibur6” Al&” Sandbar Shark 245 a 0.04&0.046 f 183 f 8-12 56-75 Casey and Natanso” (1992):
0.046-0.05 m 178 m Casey et al. (1985):
Spri”ger(1960)
Nurse Shark 304 0.068 H 230-240 f 21-28 28 Compagno (I 984):
22s m Pm” and Casey (1990)
Sphymo leuini Cornuda ComGn Scalloped 309 0.054-0.249 f 210-250 f 30 49 Branstetter (1987~):
Hammerhead Pratt and Casey (I 990):
0.054-0.222 m 163-210 m Che” et al. (1990):
Kllmley (1987)
TiburBn Toro Bull Shark 324 0.039-0.076 180-230 f I2 60-75 Pratt and Casey (1990):
(Xmoa) 157-226 m Bransretter and Sttles (1987):
Thorson and Lacey (1982):
Garrick (1982)
Carcharhinur fdciformis Tlbur& Sedoso Silky Shark 330 0.101~0.153 232-245 f I2 76 Bonfil et al. (1993)*:
225 ” Branstetter (1987~);
Garrick (1982)
Carrhorhinus ohrcurus Tiburdn Negrillo Dusky Shark 365 0.039 f 257-300 f I4 100 Natanson et al. (I 995):
0.038 m 280 m Sprmger (I 960):
Clark and vo” Schmbdt (1965)
Golencerdo cuuirr Tinforera Tlper Shark 550 0.184 315-320 f 43-82 51-85 Kauffman (1950):
310 m Compagno (1984):
Branstetter et al. (1987)
Sphyrna mokormn Comuda Figante Great 560 0.133 H 250-300 f 40 50-70 Castro (1983):
Hammerhead Compagno (1984):
243-269 m Pratt and Casey (I 990)
’ R. Bonfll. unpublished data. K. coefficient of rhe Vo” Berralanffy growth model, eslimated by direct methods excepr where marked by an H (method of Holden. 1974): f: females: m: males)
112 R. Bonfil/Fisheries
In the southern Gulf of Mexico there is informa-
tion on fisheries biology for only three species.
Alvarez (1988) provides growth parameters for the
Atlantic sharpnose shark and the bonnethead shark
of Yucatan, estimated via length frequency analysis.
His results show values of La = 120 cm TL; K =
0.152; I, = - 1.75 years, for the bonnetbead shark,
and Lx=ll2cmTL; K=0.169; to= -1.90years
for the Atlantic sharpnose shark. However, Alvarez’s
estimates of the growth constant K may be an under-
estimate, as suggested by the results of direct aging
methods in the northern Gulf of Mexico for the
bonnethead shark (Parsons, 1987) and the Atlantic
sharpnose shark (Branstetter, 1987a), the latter vali-
dated with tetracycline marking. In addition, Alvarez
did not detect the well-established differential growth
of the sexes in bonnethead sharks. Although different
growth patterns cannot be discounted for populations
of sharks in the northern and southern Gulf of Mex-
ico, more likely the low power of the length fre-
quency technique used by Alvarez (Basson et al.,
1988) is responsible for these differences.
Bonfil et al. (1993) studying the silky sharks of
the Campeche Bank, report on reproductive and
growth parameters, the latter determined by vertebral
readings and verified via back-calculation and length
frequency analysis. Males attain maturity at about
225 cm TL or 10 years and females at about 232-245
cm TL or after 12 years. Silky sharks in the
Campeche Bank are born during May-July at about
76 cm TL after approximately 12 months of gesta-
tion. Litter sizes are generally of 12 pups. Growth in
both sexes is described with VBGF parameters Lm
=311 cmTL; K=O.lOl;and to= -2.72years.
There are no studies focused on feeding habits of
sharks in the southern Gulf of Mexico. There is only
ancillary information reporting food items for some
shark species (Hemandez, 1987; Marin, 1992) and
the field observations of the author that silky sharks
prey heavily on the Mayan octopus.
5. Assessment and management
There are no assessments of the shark fisheries of
the southern Gulf of Mexico as a single unit, either
for individual species or for the whole system as an
aggregated multi-species fishery. Yucatan is the only
Research 29 (1997) 101-117
area where some preliminary assessments have been
done for three of the most important species. This
shortage of assessment studies is largely due to a
lack of adequate baseline information: effort data are
not routinely compiled by the Department of Fish-
eries and catches are not available for separate
species. Furthermore, there is no fishery-independent
abundance information for sharks in the southern
Gulf. Finally, as pointed out previously, the under-
standing of the biological parameters of local stocks
is appallingly poor and has been tackled only re-
cently and just for a few species.
The assessment studies carried out for three
species in Yucatan, although preliminary in nature,
provide some insight into possible problems facing
shark resources in the area. The surplus-production
assessments (equilibrium Schaefer and Fox models,
and the Walter (1975) non-equilibrium modification)
of bonnethead and Atlantic sharpnose sharks of Al-
varez (1988) suggest MSY values around 22 and 75 t
year - ’ for each species respectively. Alvarez points
out that both species were close to these levels of
exploitation. His Beverton-Holt yield per recruit as-
sessment suggests that the Atlantic sharpnose shark
is already overexploited and the bonnethead shark is
at optimum exploitation. Alvarez further reports VPA
estimates of standing stocks of 246 t for the Atlantic
sharpnose shark and 86 t for the bonnethead shark.
For the silky shark, Bonfil (1990) finds growth
overfishing when applying the yield per recruit model
of Beverton and Holt (19571, and suggests that the
size at first capture should be set at about 140-170
cm TL. These authors coincide in suggesting a cap-
ping of the fishing effort for shark fisheries in the
area.
The two studies above are only preliminary and
need to be superseded by more appropriate ap-
proaches, which unfortunately demand more and bet-
ter information. Both authors resort to some neces-
sary extreme assumptions due to limitations in avail-
ability of information: crude estimates of effort (un-
certain data on total fisheries fleet size) and likely
underestimates of growth in the first study; ‘gues-
stimates’ of natural mortality in the latter. Further-
more, the use of yield per recruit approach assumes
that recruitment is independent of variations in the
parent stock, which is probably not true for viviparous
sharks. However, this preliminary approach can
 R. Bonjil/ Fisheries Research 29 (1997)
sometimes evidence unquestionable facts such as the
growth overfishing of the silky shark reported by
Bonfil ( 1990).
The apparently persistent trend of concentrating
fishing effort in juvenile sharks should be a major
cause of concern. Dealing with the potential problem
of overfishing of juveniles is perhaps one of the most
pressing problems in the Mexican shark fisheries.
Unfortunately, until the time this review was submit-
ted for publication, no specific management mea-
sures had ever been taken for shark fisheries in
Mexico. However, the Under-Ministry of Fisheries is
currently considering this issue and shark manage-
ment might become a reality soon.
6. Discussion
6. I. Deficiencies in information, and priorities
research on sharks of the Gulf of Mexico
Current knowledge prevents a complete and sound
assessment of the situation of shark fisheries in the
southern Gulf of Mexico. Information is incomplete
or non-existent in several areas of particular impor-
tance for fisheries assessment and management. Fur-
thermore, there is a need for the establishment of
efficient management structures that allow for scien-
tific information to be translated into timely and
proficient management and enforcement. Future re-
search on sharks and their fisheries in the southern
Gulf of Mexico should focus on six areas of special
attention discussed below.
The biology of most stocks in the southern Gulf
of Mexico must be unveiled. Altogether, there is
only partial information on age, growth, and repro-
duction for three out of the 14 main species in the
area. This is insufficient if assessments are to be
done via single-species age-structured models. Life
history information from the NW Atlantic or the
northern Gulf of Mexico cannot simply be extrapo-
lated to the southern Gulf. Age-structured models are
often sensitive to life history parameters, and geo-
graphic variation in these has been documented for
distant stocks of shark (e.g. silky sharks from the
western Atlantic vs. those in the Indian Ocean; Bon-
fil et al., 1993) as well as for neighbouring popula-
tions (e.g. bonnethead shark; Parsons, 1987). Life
101-117 113
history studies of the populations residing in the
southern Gulf of Mexico should begin by focusing
on the nine main species in the fishery, then consid-
ering the five species of secondary importance. The
determination and validation of age and growth by
direct methods, as well as the understanding of the
reproductive cycles should be given priority. While
this research is in progress there is room only for
precautionary management measures such as those
suggested below. An alternative line of research with
much wider application, would be the development
of robust methods that allow the estimation of impor-
tant life history parameters based on the knowledge
of just one or two parameters for a specific stock of
fish.
The lack of historical records of catch and effort
has restricted the use of dynamic surplus-production
models to assess the status of the exploitation of
for sharks in the region. Although this type of informa-
tion will not be available for individual species for a
long time (probably until separate markets are devel-
oped for different species), it is possible to rescue
valuable information from log-books or payment slips
from private fishing companies or cooperatives
throughout the southern Gulf of Mexico, to build
time series of aggregated shark catches and corre-
sponding efforts. This can be achieved quickly and
cheaply, and should be an immediate task that would
allow for a first assessment of the situation of the
fisheries. The detail of the information should be
properly preserved, so that patterns could be identi-
fied for different species groups. The advantage of
obtaining different time series of information scat-
tered throughout the region is that this would permit
also a rough assessment of the spatial distribution of
effort. Such information could prove vital for avoid-
ing situations of hyperstability or hyperdepletion that
severely hamper the utilisation of commercial CPUE
data as an index of abundance (Hilbom and Walters,
1992). Alongside this, a start should be made in
collecting fishery independent abundance informa-
tion for sharks. Given the lack of reliable catch and
effort statistics per species, and the caveats of aggre-
gated commercial CPUE data, the availability of
long-term abundance surveys is likely to play a
major role in the sound assessment of shark re-
sources. For this purpose, cooperation between man-
agement and industry could prove to be very benefi-
114 R. Bonfil/ Fisheries
cial. It is likely that ‘chartering’ of commercial
fishing boats to do surveys, will be the most efficient
and economic way to obtain fishery independent
abundance indices for sharks.
The identification of migratory patterns, the proper
delimitation of fish stocks and a good understanding
of the interdependence between stocks/substocks,
are fundamental parts of fisheries management. These
three frequently interrelated aspects are essential in
order to make sense ecologically, but more impor-
tantly, they pafom also as facilitators for the ade-
quate implementation of effective management poli-
cies. This is especially important in the event of
shared stocks between neighbouring States or Na-
tions.
Despite some statements indicating the existence
of single stocks of large sharks between Mexican
and US waters of the Gulf of Mexico (Branstetter,
1990; Hoff and Musick, 1990) there is no hard
evidence to confirm this. Tagging studies are incom-
plete because the large majority of the information
pertains to sharks tagged outside the Gulf of Mexico.
This provides at best a biased and partial picture of
the main patterns of shark movements. Moreover,
the rates of movement of sharks between different
areas and the proportion of the populations involved
in such movements have not been assessed. A wide
scale long-term cooperative tagging programme for
sharks in Mexican waters of the Gulf needs to be
implemented soon. Newborn and early juvenile
sharks can be tagged in the identified nursery areas
while older sharks will have to be tagged through
fishing cruises for this specific purpose given the
lack of a large sport fishery in the area. Without this
information, the timing and size of migrations, the
rates of movement between different parts of the
stocks, and the delimitation of such stocks will re-
main unknown.
Our poor understanding of shark movements in
the Gulf of Mexico and neighbouring areas is further
complicated by the lack of direct and comprehensive
stock identification studies for the main species based
on population genetic techniques. In summary, there
have been not enough studies directed toward resolv-
ing the status and dynamics of the stocks of sharks in
the NW Atlantic including the Gulf of Mexico. This
must be addressed as a requirement for wide scale
fisheries management (i.e. joint management be-
Research 29 (1997) IO1 - I1 7
tween Mexico and the USA) and it will entail the
combination of genetic identification of stocks and
the tagging studies outlined above. The genetic study
of populations must ensure a representative sampling
of the whole distribution range of each species in the
region. Once the main stocks and substocks have
been identified, the degree of mixing throughout
their geographical ranges should be considered for
establishing more appropriate management proce-
dures between the two countries, if this is warranted.
Meanwhile, precautionary management should be
implemented by the Mexican Government as out-
lined below.
A detailed study of the utilization of coastal la-
goons as shark nursery areas and the impacts of
human activities in those lagoons should be con-
ducted promptly. Identifying the most important
shark nursery areas and unveiling their dynamics
would allow for more effective management mea-
sures. Such measures should ensure the viability of
the stocks by protecting newborn and early juveniles,
which are the most fragile stages of sharks’ life
cycles. In the longer term, monitoring of carefully
chosen nursery areas should focus on determining
the relationship between stock and recruitment and
the estimation of natural mortality of newborns and
juveniles that are essential for age-structured models.
In addition, modelling efforts should be oriented
towards determining the utility of protecting nursery
areas and the degree of protection needed in order to
positively impact the stocks.
Last but not least, the socio-economic importance
of shark fisheries in the different subregions of the
southern Gulf of Mexico must be addressed. This
should include studies of the importance of each of
the two main fisheries, directed and incidental. All
this information will be a vital part of any fisheries
management decision because management has to be
consistent not only with biological processes, but
also with the social and economic needs of fishing
communities along States and Nations.
6.2. Precautionary and immediate management
Despite the lack of a complete understanding of
the situation of shark fisheries in the southern Gulf
of Mexico, there are some important features on the
information presented here. Such features can be
 R. Bonjl/Fisheries
used to outline immediate management needs pri-
marily based on the precautionary principle of fish-
eries management.
The incidence of large catches on newborn and
juveniles of several shark species in nursery areas is
of particular concern. Such an exploitation regime,
apart from leading to the wasteful practice of growth
overfishing, will compromise the future recruitment
of species of prime importance to the fishery if it is
not controlled soon. Possible immediate management
measures directed at reducing juvenile fishing mor-
tality include the banning of small meshed gillnets in
all coastal lagoons identified as important nurseries
for sharks. Minimum size restrictions could be im-
posed for the capture of species with nursery areas in
continental shelf waters.
The fragility of shark stocks and the situation of
the US shark fisheries call for a capping of the effort
and catches of sharks in the Gulf of Mexico. Al-
though Mexican shark catches in the area seem to be
fairly stable and without signs of trouble, catch
stability is not enough guarantee of healthy stocks,
specially for such fragile species as sharks. The lack
of any abundance index for these resources should
warn against the adoption of any risk-prone policies
such as expanding the fishery. The large increase in
catches in the last 1.5 years in the Gulf of Mexico
and the known low reproductive potential of most
elasmobranchs can lead to a major collapse in the
fisheries if further growth in the effort is allowed.
Shark fisheries should not expand until we gain a
better understanding of the biology and ecology of
the main stocks in the Gulf of Mexico.
As a complement to any preliminary management
measures, a system for the permanent collection of
catch and effort data should be established in the
main shark fishing centres of the southern Gulf of
Mexico. Preferably such data should be collected on
a species specific basis, or at least discriminating the
most important species groups and trying to preserve
any spatial detail of the information.
Acknowledgements
I am grateful to Daniel Pauly for helpful discus-
sions, useful suggestions to improve the original
manuscript, and for arranging Fig. 5 and getting it
Reseurch 29 (1997) 101-I 17 115
redrawn at ICLARM, Manila. The suggestions of
two anonymous referees for improving the
manuscript are also appreciated.
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