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AQUATIC CONSERVATION: MARINE AND FRESHWATER ECOSYSTEMS

Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)


Published online 14 July 2006 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/aqc.784

Effects of trap fishing on reef fish communities

JULIE P. HAWKINSa,*, CALLUM M. ROBERTSa, FIONA R. GELLa,1


and CALVIN DYTHAMb
a
Environment Department, University of York, Heslington, York, UK
b
Department of Biology, University of York, Heslington, York, UK

ABSTRACT
1. Trap fishing is widespread on coral reefs but the sustainability of this practice is causing concern
because it is efficient and unselective. The effects of trap fishing were investigated by comparing fish
assemblages among six Caribbean islands subject to different trapping pressures. These ranged from
none in Bonaire and Saba increasing through Puerto Rico, St Lucia, Dominica and Jamaica
respectively.
2. Fish were censused at depths of 5 m and 15 m on fore-reef slopes by counting the numbers within
replicate 10 m diameter areas for 15 min. Between 64 and 1375 counts were made in each country.
3. In St Lucia and Jamaica abundance of fish censused on the reef was compared to representation
in traps which were visually sampled underwater in the area of fish counts. Twenty-three traps were
sampled in Jamaica and 75 in St Lucia. For some comparisons between these islands, St Lucian
sampling effort was reduced to that of Jamaica (23 traps and 112 counts) by randomly sub-sampling
10 times.
4. Traps contained 54 different species in St Lucia and 22 in Jamaica, while there were 90 and 57
respectively in counts. After reducing St Lucian sampling effort to Jamaican levels, an average of 35
species were found in traps and 70 seen in counts. Of these, 76% in St Lucia and 73% in Jamaica
were relatively more abundant in traps than they were on the reef.
5. Species were considered to be highly susceptible to trapping if the ratio of their abundance in
traps compared to that on the reef exceeded 3:1. Trapping pressure was approximately three and a
half times greater in Jamaica than St Lucia. After equalizing sampling effort, there was an average of
16 highly trappable species in St Lucia compared to 13 in Jamaica. Species did not always appear
highly trappable in both countries. Eleven of St Lucia’s highly trappable species were absent from
Jamaica (falling to 8.5 on average after equalizing sampling effort), but none vice versa, suggesting
that trapping may have contributed to their absence or rarity on Jamaican reefs.
6. The Tetraodontiformes, which include many non-target species, were particularly susceptible to
trapping in both countries. Their abundance in the six islands censused was inversely related to trap
fishing pressure, as was that of two other non-target families, butterflyfish (Chaetodontidae) and
angelfish (Pomacanthidae).

*Correspondence to: Julie P. Hawkins, Environment Department, University of York, Heslington, York, Y010 5DD, UK. E-mail:
jph7@york.ac.uk
1 Present address: Wildlife and Conservation Division, Department of Agriculture, Fisheries and Forestry, Knockaloe Farm, Peel, Isle

of Man IM5 3AJ, UK.

Copyright # 2006 John Wiley & Sons, Ltd.


112 J.P. HAWKINS ET AL.

7. To determine whether fish that are common in traps in St Lucia are reaching sexual maturity
before capture, size frequency data for 23 species from a sample of trap catches were gathered and
examined for their state of maturity. In seven species, more than a third of 705 trapped fish were
immature, indicating that trap fishing causes growth over-fishing (premature removal of fish), and
calling into question the sustainability of yields for these species.
8. In conclusion, at the intensities seen in this study, trap fisheries cause serious over-fishing,
reduce biodiversity, and alter ecosystem structure. While commonly perceived as low impact, coral
reef trap fisheries in the Caribbean and further afield, need tighter regulation and control.
Copyright # 2006 John Wiley & Sons, Ltd.

Received 2 December 2003; Revised 1 February 2006; Accepted 15 February 2006

KEY WORDS: coral reef; fish pot; overfishing; marine reserves

INTRODUCTION

Trap fishing is one of the oldest fishing methods and is widely practised throughout the world in both
tropical and temperate regions (Recksiek et al., 1991; Slack-Smith, 2002). It is the most common fishing
technique used on Caribbean coral reefs (Gobert, 1998). In this region traps typically consist of a bamboo,
wooden, or steel frame covered with chicken wire with a mesh size ranging from 2–5 cm (Stevenson and
Stuart-Sharkey, 1980). Depending on their design, traps may have one or two funnel-shaped openings
through which fish can enter. Fish can also escape from these openings or by squeezing through the mesh
(Gobert, 1998). Trap retention rates vary between locations and depend on how long a trap has been
soaked for, and the trap’s internal structure (Whitelaw et al., 1991; Sheaves, 1995).
There are many reasons why traps are popular with reef fishers. They are easy to deploy (Ferry and
Kohler, 1987), relatively inexpensive to make (Garrison et al., 1998), and can be used in areas with rugged
substrata (Miller and Hunte, 1987). If the weather is bad, fishers can leave their traps out until conditions
improve. Traps are also effective at catching a wide variety of species (Stevenson and Stuart-Sharkey, 1980).
Robichaud et al. (2000) reported there to be little quantitative information on what factors influence the
catchability of reef species. Their own study concluded that mobility was important because it increased the
rate at which fish encountered traps and hence their likelihood to enter them. While they found that prey
tended to avoid entering traps containing predators, they suggest that prey might also enter traps in an
effort to avoid predation. On balance, they found that the presence of prey in traps increased the
catchability of piscivores by reducing their escape rates. The study also noted that behavioural
characteristics such as schooling species following each other into traps and solitary species avoiding
them may influence catchability, while Wolff et al. (1999) have suggested that some species could be
attracted into traps merely by the structure they provide.
Some of the factors that make trapping popular with fishers can also cause problems. As a fishing
method that is effective but unselective it produces much unwanted bycatch which is commercially wasteful
and could be threatening to biodiversity (Dayton et al., 1995; Alverson and Hughes, 1996; Boehlert, 1996).
Around Florida about 100 different species are caught in traps with levels of bycatch ranging from 38% to
54% (Sutherland and Harper, 1983; Taylor and McMichael, 1983). Taylor and McMichael (1983) found
that unwanted angelfish (Pomacanthidae) constituted 17% of catches, trunkfish (Ostraciidae) 9%, and
butterflyfish (Chaetodontidae) and wrasse (Labridae) 5–6% each. In the Gulf of Mexico levels of bycatch
from trap fishing are reported to have reached 65% (NMFS, 1995).
What counts as bycatch varies between regions depending on local traditions, income levels and intensity
of exploitation. In parts of the Caribbean, such as Jamaica and Haiti, where many people are very poor and
fisheries intense, almost nothing is discarded and fish are used either for food or for bait (Ferry and Kohler,
1987; Sary, 1995). However, even if all the catch is used, the potential threat to biodiversity remains.

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
TRAP FISHING EFFECTS ON REEF FISH 113

As desirable food fish become increasingly rare, even people from wealthy nations resort to eating species
that were once considered ‘trash’ such as squirrelfish (Holocentridae) and angelfish. This has happened in
Bermuda, where, by the 1980s, trap fisheries had decimated the grouper (Serranidae) and snapper
(Lutjanidae) populations (Butler et al., 1993).
Although trapping is a multi-species fishing technique, it has more serious impacts on some species than
others. For example, in the US Virgin Islands trapping, together with hook and line fishing, has virtually
eliminated the Nassau grouper (Epinephelus striatus) (Garrison et al., 1998). Over a 6-year period during
the 1970s, a spawning aggregation which consisted of tens of thousands of fish was wiped out (Olsen and
LaPlace, 1978). Trap fishing also poses a threat to juvenile fish because of the small mesh sizes used (Sary
et al., 1997), particularly to high-bodied species such as surgeonfish (Acanthuridae). In areas with high
levels of trapping, juveniles can represent a significant proportion of the catch (Robichaud et al., 1999) and
trapping can lead to reduced productivity through growth over-fishing, i.e. premature removal of fish
leading to lower catches.
A further problem associated with trap fishing is the damage it causes to corals and other bottom-living
organisms when traps are dropped onto the reef (Yoshikawa and Asoh, 2004). In St Lucia we observed
fishers breaking corals with grappling hooks while retrieving their traps (pers. obs.). If traps are lost (these
are often called ‘ghost traps’), they can continue catching fish for weeks, or even months, until they fall
apart. Biodegradable panels, which give way sooner than traps, help reduce this waste (Stewart and Ferrell,
2002). Despite the problems associated with trap fishing it has received little regulation other than mesh size
restrictions and limits on numbers and not much is known about how such management measures will
improve sustainability. Fishers have been reluctant to install biodegradable panels into their traps, even
when required to by law (Blott, 1978; Garrison et al., 1998; pers. obs.). In Bermuda, the problems
associated with trapping led to the fishery being closed down in 1990 (Butler et al., 1993) and plans are
under way to phase it out in the Gulf of Mexico by 2007 (http://www.reefguardian.org/Campaigns/
FishTrapGOM/MoreInfoFTrapGOM.html).
In this study, we investigate questions relating to the sustainability and biodiversity impacts of trap
fishing in the Caribbean by examining the sensitivity of a broad range of species to trapping and looking at
the incidence of capture of juvenile fish of commercially valuable species.

METHODS

Coral reef fish assemblages were censused during single visits to the islands of Bonaire, Saba, Puerto Rico,
Dominica and Jamaica between 1994 and 1998 (Figure 1), and on five visits to St Lucia, either annually or
biennially, between 1994 and 2000. Fish were counted over sections of coast covering ca 5–20 km,
depending on island size, using an adaptation of the stationary point visual census method developed by
Bohnsack and Bannerot (1986). For each 15-min count a 10-m tape was laid on the reef to indicate the
diameter of a cylinder extending 5 m upwards. The location for the first count was the point of entry for the
dive and subsequent counts were made along the same depth contour. Approximately equal numbers of
counts were performed at depths of 5 m and 15 m and each was separated by at least 10 m. The number of
counts performed in each country was: 72 in Bonaire, 78 in Saba, 98 in Puerto Rico, 64 in Dominica and
112 in Jamaica. In St Lucia, we used a sample of 275 counts in 1995 for an inter-island comparison of fish
assemblages. For comparisons between fish caught in traps and counted on the reef we used a sample of
1375 counts taken between 1995 and 2000.
Comparable measures of the absolute fishing intensity for these islands were not possible, because data
have been reported at different levels of detail, and the types of fishing gear used vary between islands.
However, using estimates of number of fishers per kilometre of reef, it is possible to rank fishing pressure
among the islands. Data from published studies, information from fisheries departments and marine parks,

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
114 J.P. HAWKINS ET AL.

Figure 1. Location of the islands studied.

and personal observations on numbers of active fishers and gear types used were combined for the purpose
(Table 1). On this basis fishing pressure was low in Bonaire (0.06 fishers km1 of reef), light in Saba
(0.12 fishers km1) and higher in Puerto Rico (0.83 fishers km1). Fishing pressure was intense in St Lucia
(3.23 fishers km1) and Dominica (3.03 fishers km1) and extremely intense in Jamaica (7.14 fishers km1).
These measures include all methods of fishing. If trapping is considered independently, then Bonaire and
Saba have none, levels were moderate in Puerto Rico, high in St Lucia and Dominica, and extremely high in
Jamaica. One other way of comparing fishing pressure is from trap counts. We counted traps during fish
counts in Jamaica and St Lucia only. Twenty-three traps were observed during 112 fish counts in Jamaica
while in St Lucia 80 traps were observed during 1375 counts, implying that trapping pressure is
approximately three and a half times greater in Jamaica.

Effects of trap fishing pressure on non-target species throughout the Caribbean


The vulnerability of families containing species that are not generally popular food fish, but which do get
caught in traps, was investigated by comparing fish abundance on the reef among the six Caribbean islands
of a superfamily, the Tetraodontiformes (pufferfish (Tetraodontidae), boxfish and trunkfish (Ostraciidae),
triggerfish and filefish (Balistidae)), butterflyfish (Chaetodontidae) and angelfish (Pomacanthidae). The
abundances of soldierfish (Holocentridae) which have no commercial value in Saba and Bonaire, are low-
value food in Puerto Rico, and are commonly eaten in the other three countries where they make up a large
proportion of trap catches were also compared. Species from these families are predominantly taken by
trap fishing but some are also caught by spearfishing. Mean abundance per count of each family was
analysed with one-way ANOVAs and post hoc LSD tests. To avoid any possibility of pseudoreplication, we
reduced the degrees of freedom in the test to one for each island, i.e. treating islands as samples rather than
replicates within islands. Data for St Lucia were taken from the 1995 survey prior to implementation of
marine reserves there (Roberts et al., 2001).

Fish abundance on the reef versus that in traps


In St Lucia and Jamaica species’ abundances on the reef were compared to that inside traps. Trap contents
were recorded during dives to census fish after the fish counts were complete. On a single dive
approximately 100–120 m linear distance of reef was searched for traps within the area of the fish counts.
The number of traps counted per dive ranged from 0 to 5 in both countries. The numbers of each species
present were recorded for each trap. The contents of 23 traps in Jamaica and 75 traps in St Lucia were

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
TRAP FISHING EFFECTS ON REEF FISH 115

sampled. In order to compare results between the two countries it was necessary to reduce the level of
sampling intensity in St Lucia to that in Jamaica. This was done by selecting 10 random samples of 112 fish
counts from the 1375 performed in St Lucia, and ten random samples of the contents of 23 traps from the
75 traps sampled in St Lucia. Average values obtained from these two sets of 10 random samples are used in
the results to make direct comparisons possible with Jamaica.

Analysis of fish trappability


The trappability of a species is expressed as the ratio of its proportional abundance in traps (i.e. the fraction
that each species represented of the total sample of trapped fish) compared to its proportional abundance in
all fish counts. This provides a measure of vulnerability to trapping. Data were pooled across depth and
observer and, from counts in St Lucia, for all the years that were sampled. Species with a trappability value
of one were equally represented in traps and fish counts, those scoring higher were more represented in
traps and those scoring lower were more represented in counts. Species were considered highly trappable if
their trappability ratio was greater than 3 and highly untrappable if it was less than 0.33 (i.e. if a species was
three times less represented in traps than on the reef). Trappability at the family level was also assessed.
Prior to analyses all species shown in the Appendix were removed from the dataset. These were generally
either too small (maximum cross-sectional body dimension less than 3 cm) or too big (maximum cross-
sectional body dimension greater than 10 cm) to be caught by traps, or could not be counted effectively
using visual census because they were cryptic. Cryptic species would tend to be under-represented in counts,
thereby giving misleading trappability ratios. Small juveniles of some species included in the analysis could
pass freely through the traps and large numbers of such fish on the reef could lead to underestimates of
trappability for such species.

Size at maturity of fish caught within traps in St Lucia


Between September 2000 and February 2001, the contents of 530 traps landed from 134 fishing trips that
had taken place within the study area in St Lucia were sampled. Size frequency distributions for 23
commercially important species were compiled and the gonads of 705 fish bought from trap fishers over a
two-day period in September 2002 were examined and the minimum size at maturity calculated, and where
possible the size at which 50% and 100% were mature.

RESULTS

Effects of trapping on non-target species throughout the Caribbean


Abundances of Tetraodontiformes, butterflyfish and angelfish on the reef, were all inversely related to
fishing pressure (Figure 2) (one-way ANOVAs: F5;6 ¼ 33:5; p50:001 (Tetraodontiformes); F5;6 ¼ 48:0;
p50:001 (butterflyfish); F5;6 ¼ 38:6; p50:001 (angelfish)). Tetraodontiformes abundance was much greater
in Bonaire and Saba, where there was no trap fishing, compared to those islands where trapping did occur.
Butterflyfish, abundance declined steadily over the fishing pressure gradient. However, the difference in
abundance between the most- and the least-fished countries was great, with only 6% as many fish present
in Jamaica as Bonaire. Angelfish were the least abundant of the families sampled. They were most common
in Bonaire and Saba, less so in Puerto Rico and were uncommon to absent in the most heavily fished islands
of St Lucia, Dominica and Jamaica. Soldierfish abundance initially increased with increasing fishing
pressure but fell abruptly in intensively fished Jamaica (one-way ANOVA: F5;6 ¼ 42:7; p50:001).

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
116

Table 1. Characteristics of fisheries and relative fishing intensities at each of the six study locations

Location No. of active Length of Gears used Species targeted Fishing Reference

Copyright # 2006 John Wiley & Sons, Ltd.


fishers reef fished intensity
(km) (fishers/
km reef)
Bonaire } 4 full-time 61.8 hook and near reef pelagics (e.g. 0.06 K. De Meyer (Bonaire
leeward coast line jacks, rainbow runners) Marine Park Manager),
from Klein personal communication,
Bonaire north and personal observations
to Slagbaai

Saba } 10 part-time ¼ 6.9 spear mainly predatory fish such 0.12 David Kooistra (Saba
J.P. HAWKINS ET AL.

leeward coast 0:83 f.t.e.a as large snappers and Marine Park Manager),
from SE to N groupers personal communication,
of island and personal observations

Puerto Rico } 64 full-time 2 78.3 lines, trap, predominantly snappers, 0.83 Matos-Caraballo, 1997
reefs around part-time ¼ 65 nets, spear but grunts and near-reef Morelock et al., 1994
La Parguera in f.t.e.a pelagics important; a wide
SW of island variety of reef associated
species also targeted

St Lucia } 25 full-time 4 8.4 traps, hook wide range of species 3.23b Gell et al., 2001
reefs of the SW part-time ¼ 27 and line, including many small- Goodridge, 1996
coast around f.t.e.a limited bodied planktivores,
Soufrière spearing, invertivores, omnivores
few gill nets and herbivores; predators
are highly desirable but
only limited numbers
caught; inedible species

Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)


DOI: 10.1002/aqc
are used for bait
Dominica } 32 full-time 18.5 traps, hook wide range of species 3.03b H. Guiste, (Chief Fisheries
reefs of the 47 part-time and line, including many small- Officer, Dominica),
NW coast =56 f.t.e.a extensive bodied planktivores, personal communication
around spear and invertivores, omnivores Roberts and Hawkins, 1996
Portsmouth nets and herbivores; predators
are highly desirable but
only limited numbers
caught; inedible species
are used for bait

Jamaica } 58 full-time 24 9.9 traps, hook virtually everything larger 7.14 Sary, 1995
reefs of the part-time ¼ 70 and line, than 10 cm is targeted. Sandeman and Woodley,
North coast f.t.e.a extensive Majority of catch is small- 1994
around spear and bodied; large predators Allison, 1992
Discovery Bay nets desirable but extremely

Copyright # 2006 John Wiley & Sons, Ltd.


uncommon
a
The f.t.e. is full-time equivalents. In Saba we assumed that each fisher fished for 1 day every 2 weeks (assuming a 6-day working week) because all fishing on this island is
recreational. Hence, each fisher is considered one-twelfth a full-time equivalent. In all other countries, where fishing is done on a commercial or subsistence basis, we
assumed that part-time fishers fished for half of their time.
b
The amount of reef available to fishers in St Lucia and Dominica is virtually identical. However, because the range of gears used in St Lucia was more limited (i.e. less
spear and net fishing), we considered this island to be slightly less heavily fished than Dominica.
TRAP FISHING EFFECTS ON REEF FISH
117

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DOI: 10.1002/aqc
118 J.P. HAWKINS ET AL.

Bonaire
Saba
Puerto Rico
St Lucia
Dominica
Jamaica

4 2
Mean number per reef census

3
Tetradontiformes Angelfish
Spearman's rho = - 0.45 Spearman's rho = - 0.36
2 1

0 0
0 2 4 6 8 0 2 4 6 8

4 Butterflyfish 25 Soldierfish
Mean number per reef census

Spearman's rho = - 0.57 20 Spearman's rho = 0.17


3

15
2
10

1
5

0 0
0 2 4 6 8 0 2 4 6 8
Fishers / km reef Fishers / km reef

Figure 2. Mean abundance ðs:e:Þ of Tetraodontiformes, butterflyfish, angelfish and soldierfish versus fishing pressure.

Trappability of species
Figures 3 and 4 compare species abundance in traps to that on the reef in Jamaica and St Lucia respectively.
Species are listed in rank order of their abundance in counts. In Jamaica, Haemulon plumieri, Holacanthus
ciliaris, Lutjanus apodus, Lutjanus synagris, Lactophrys polygonia and Chilomycterus antillarum were absent
from counts but present in traps, while the same was true for C. antillarum and Haemulon striatum in
St Lucia. The most abundant species in both countries was the wrasse, Clepticus parrae, which has been
omitted from all graphs and tables since its dominance obscures results for other species. When C. parrae
was included in the analysis it comprised 67.6% of the fish population in Jamaica and 66.5% in St Lucia,
but only 0.6% and 3.3% respectively in traps, showing that this schooling, mid-water, planktivorous
species, not preferable for eating, is not readily trappable.
Catches were generally diverse. In both countries Stegastes planifrons, Acanthurus bahianus, Sparisoma
aurofrenatum and Scarus taeniopterus were among the six most numerous in counts. In Jamaica, 22 species
were recorded in traps and 57 in counts (Figure 3), while in St Lucia there were 54 in traps and 90 in counts
(Figure 4). Reducing the sampling intensity of St Lucia to that of Jamaica gave average figures of 35 species
in traps and 70 in counts, for 10 randomly picked sub-samples of 23 traps and 112 counts.

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
TRAP FISHING EFFECTS ON REEF FISH 119

20

Traps
10
Counts
% in counts

10

20
% in traps

30
Cephalopholis fulva

Lutjanus mahogoni
Cephalopholis cruentata
Hypoplectrus puella

Lutjanus jocu

Lactophrys polygonia
Scarus iserti

Sparisoma viride

Holocentrus vexi

Stegastes dorsopunicus

Sparisoma rubripinne
Stegastes diencaeus
Acanthurus bahianus
Stegastes planifrons
Scarus taeniopterus

Holocentrus marianus

Pseudopeneus maculatus
Haemulon flavolineatum

Hypoplectrus indigo

Lutjanus analis
Sparisoma aurofrenatum

Myripristis jacobus
Holocentrus rufus
Acanthurus coeruleus

Epinephelus guttatus
Chaetodon ocellatus

Scorpaena plumieri
Xanthichthys ringens
Haemulon carbonarium
Haemulon aurolineatum

Lutjanus synagris
Holacanthus ciliaris

Chilomycterus antillarium
Stegastes leucostictus

Microspathodon chrysurus
Haemulon chrysargyreum
Bodianus rufus

Hypoplectrus guttavarius
Cantherhines pullus
Halichoeres radiatus
Chaetodon capistratus
Sparisoma chrysurus
Haemulon sciurus
Hypoplectrus nigricans
Hypoplectrus chlorurus
Holacanthus tricolor

Chaetodon striatus

Bothus lunatus
Acanthurus chirugus
Melichthys niger
Diodon holacanthus
Rypticus saponaceus
Pricanthus cruentatus

Holocentrus adscensionis
Plectrypops retrospinis
Diodon hystrix

Lutjanus apodus
Stegastes variabilis

Hypoplectrus unicolor

Sphoeroides spengleri

Lacophrys triqueter

Haemulon plumieri
Mulloidichthys martinicus

Species
Figure 3. Proportional representation of species in traps compared to on the reef in Jamaica.

Table 2 shows the trappability ratios of species in Jamaica and St Lucia. These represent the ratio
of abundance of species in traps compared to that on the reef in counts. All data for trappability ratios
were taken from underwater observations since data from trap landings may have come from different
habitats or depths to our fish counts. In Jamaica 73% of species caught in traps were proportionally
more common in traps than they were on the reef, and 35% of those potentially trappable had been
caught. In St Lucia these figures were 76% and 59% respectively for the full sample of 75 traps. Reducing
the sampling intensity to that of Jamaica gave average figures of 76% and 39% respectively, for 10
randomly picked sub-samples of 23 traps and 112 counts. These figures are virtually the same as those for
Jamaica.
Including species caught in traps but not seen in counts there were 13 highly trappable and 43 highly
untrappable species in Jamaica. St Lucia contained over twice as many highly trappable species (29) as
Jamaica, and virtually the same number of highly untrappable (44), suggesting that the higher fishing
pressure in Jamaica could have eliminated species that are susceptible to trapping. However, reducing the
St Lucia sample size to that of Jamaica (23 traps and 112 counts) produces means of 16 highly trappable
species and 43 highly untrappable species, showing that the difference was simply a sampling effect. Only six
species were considered highly trappable in both countries; three Tetraodontiformes, C. antillarum,

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
120 J.P. HAWKINS ET AL.

20

Traps
% in counts

10
Counts

10
% in traps

Myripristis jacobus
Stegastes planifrons
Scarus taeniopterus
Sparisoma aurofrenatum
Acanthurus bahianus

Sparisoma viride
Abudefduf saxatilis
Haemulon chrysargyreum
Mulloidichthys martinicus
Acanthurus coeruleus
Cephalopholis cruentata

Hypoplectrus guttavarius
Holocentrus rufus

Microspathodon chrysurus
Scarus vetula
Holocentrus marianus
Haemulon aurolineatum
Stegastes diencaeus
Stegastes dorsopunnicus
Haemulon flavolineatum
Lutjanus mahogoni
Chaetodon capistratus
Paranthius furcifer
Hypoplectrus puella

Acanthurus chirugus
Scarus iserti
Chaetodon striatus
Cephalopholis fulva
Bodianus rufus
Kyphosus sectatrix

Cantherhines pullus
Lactophrys triqueter
Holacanthus tricolor

Pseudopeneus maculatus
Priacanthus cruentatus
Sparisoma rubripinne
Chaetodon aculeatus
Hypoplectrus chlorurus
Lutjanis griseus
Melichthys niger
Halichoeres radiatus
Haemulon carbonarium
Holocentrus vexillarius
Bothus lunatus
Hypoplectrus sp.
Hypoplectrus nigricans
Xanthichthys ringens
Stegastes leucostictus
Haemulon plumieri
Lactophrys bicaudalis
Lutjanus apodus
Scorpaena plumieri
Pomacanthus paru
Sparisom chrysopterum

Lactophrys polygonia
Diodon holacanthus
Plectrypops retrospinis
Balistes vetula

Epinephelus adscensionis
Lutjanus buccanella
Cantherhines macrocerus
Haemulon parra
Rypticus saponaceus
Holocentrus adscensionis
Lutjanus jocu
Holacanthus ciliaris
Chaetodon ocellatus
Lutjanus analis
Haemulon melanurum

Epinephelus guttatus
Aluterus scriptus
Anisotremus surinamensis
Haemulon macrostomum

Mycteroperca interstitialis
Haemullon sciurus

Stegastes variabilis
Gerres cinereus

Diodon hystrix
Bairdiella sanctaeluciae
Rypticus subbifrenatus
Rhomboplites aurorubens
Scarus coeruleus
Mycteroperca venenosa
Sphoeroides spengleri
Calamus bajonado
Lutjanus synagris

Haemulon striatum
Ocyurus chrysurus

Holocentrus coruscum
Calamus calamus
Bothus ocellatus

Chilomycterus antillarum
Species
Figure 4. Proportional representation of species in traps compared to on the reef in St Lucia.

L. polygonia, Diodon holocanthus; one snapper, L. synagris and two grunts, H. plumieri and Haemulon
flavolineatum. Of species that were highly trappable in St Lucia, 38% (11) were absent from both traps and
counts in Jamaica: H. striatum, Calamus calamus, Balistes vetula, Mycteroperca interstitialis, Aluterus
scriptus, Haemulon melanurum, Kyphosus sectatrix, Lactophrys bicaudalis, Cantherhines macrocerus,
Pomacanthus paru, and Ocyurus chrysurus. All occur throughout the Caribbean and so were expected to
occur in Jamaica (Humann, 1996). A third (4) of these highly trappable species were Tetraodontiformes
with most other families represented by just one species. No species trapped in Jamaica were absent from
St Lucia. To explore the possibility that this difference is an artefact of the greater sampling intensity in
St Lucia compared to Jamaica, we randomly reduced the St Lucia sample size to that of Jamaica, repeated
10 times, and counted the number of these vulnerable species that remained in the sample. On average, the
reduced samples retained 8.5 of the 11 highly trappable species.
Only the Tetraodontiformes and snappers appear highly trappable in both St Lucia and Jamaica
(Table 3). Of the four other families that were highly trappable in St Lucia, no butterflyfish or groupers
(Serranidae not including Epinephelinae) were trapped in Jamaica while chub (Kyphosidae) and porgies
(Sparidae) were not seen there. No family appeared to be highly untrappable in both countries.

Catch composition and size at maturity


Several commercially important species in St Lucia are commonly caught before they reach maturity
(Figure 5, Table 4). For seven of the 23 species sampled, over one-third of the catch taken was immature,

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TRAP FISHING EFFECTS ON REEF FISH 121

Table 2. Trappability of species in Jamaica and St Lucia calculated as the ratio of their proportional abundance in traps compared to
that in fish counts. Species with an asterisk (*) were found in traps but not seen in counts. A trappability of zero indicates that the
species was seen in counts but not in traps

Species Family Jamaica St Lucia


trappability trappability
Bothus ocellatus Bothidae not present 0
Chaetodon aculeatus Chaetodontidae not present 0.8
Epinephelus adscensionis Epinephelinae not present 0
Mycteroperca interstitialis Epinephelinae not present 11.2
Mycteroperca venenosa Epinephelinae not present 0
Gerres cinereus Gerreidae not present 0
Anisotremus surianamensis Haemulidae not present 0
Haemulon melanurum Haemulidae not present 7.7
Haemulon parrai Haemulidae not present 0
Haemulon striatum Haemulidae not present *
Holocentrus coruscum Holocentridae not present 0
Kyphosus sectatrix Kyphosidae not present 6.2
Lutjanus buccanella Lutjanidae not present 0
Lutjanus griseus Lutjanidae not present 1.1
Ocyurus chrysurus Lutjanidae not present 3.3
Rhomboplites aurorubens Lutjanidae not present 0
Pomacanthus paru Pomacanthidae not present 3.6
Abudefduf saxatilis Pomacentridae not present 1.6
Scarus coelestinus Scaridae not present 0
Scarus vetula Scaridae not present 0.1
Bairdiella sanctaeluciae Sciaenidae not present 0
Hypoplectrus sp. Serranidae not present 0
Paranthius furcifer Serranidae not present 1.2
Rypticus subbifrenatus Serranidae not present 0
Calamus bajonado Sparidae not present 0
Calamus calamus Sparidae not present 14.7
Aluterus scriptus Tetraodontiformes not present 9.2
Balistes vetula Tetraodontiformes not present 14
Cantherhines macrocerus Tetraodontiformes not present 4.1
Lactophrys bicaudalis Tetraodontiformes not present 4.9
Haemulon plumieri Haemulidae * 13.3
Lutjanus apodus Lutjanidae * 0
Lutjanus synagris Lutjanidae * 438.6
Holacanthus ciliaris Pomacanthidae * 0
Chilomycterus antillarum Tetraodontiformes * *
Lactophrys polygonia Tetraodontiformes * 18.3
Haemulon aurolineatum Haemulidae 210.3 0.2
Acanthurus chirurgus Acanthuridae 112.2 2.3
Diodon holocanthus Tetraodontiformes 42.1 18.3
Sparisoma chrysopterum Scaridae 18 0
Acanthurus coeruleus Acanthuridae 12 2.6
Acanthurus bahianus Acanthuridae 3.6 2
Haemulon flavolineatum Haemulidae 3.5 7.7
Holacanthus tricolour Pomacanthidae 1.3 0.3
Scarus taeniopterus Scaridae 1.2 0.2
Sparisoma viride Scaridae 1.1 1.6
Bodianus rufus Labridae 0.5 0
Microspathodon chrysurus Pomacentridae 0.5 0.8
Sparisoma aurofrenatum Scaridae 0.4 1.3
Cephalopholis cruentata Epinephelinae 0.3 0.3
continued over

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DOI: 10.1002/aqc
122 J.P. HAWKINS ET AL.

Table 2 continued
Species Family Jamaica St Lucia
trappability trappability
Stegastes leucostictus Pomacentridae 0.2 1.4
Scarus iserti Scaridae 0.04 0
Bothus lunatus Bothidae 0 0
Chaetodon capistratus Chaetodontidae 0 2.3
Chaetodon ocellatus Chaetodontidae 0 7.3
Chaetodon striatus Chaetodontidae 0 9.8
Cephalopholis fulva Epinephelinae 0 0
Epinephelus guttatus Epinephelinae 0 9.1
Haemulon carbonarium Haemulidae 0 7.3
Haemulon chrysargyreum Haemulidae 0 0.3
Haemulon sciurus Haemulidae 0 0
Holocentrus adscensionis Holocentridae 0 14.7
Holocentrus marianus Holocentridae 0 0
Holocentrus rufus Holocentridae 0 0.6
Holocentrus vexillarius Holocentridae 0 0
Myripristis jacobus Holocentridae 0 0.3
Plectrypops retrospinis Holocentridae 0 0
Halichoeres radiatus Labridae 0 1.1
Lutjanus analis Lutjanidae 0 0
Lutjanus jocu Lutjanidae 0 0
Lutjanus mahogoni Lutjanidae 0 3.3
Mulloidichthys martinicus Mullidae 0 0.7
Pseudupeneus maculatus Mullidae 0 7
Stegastes diencaeus Pomacentridae 0 0
Stegastes dorsopunnicus Pomacentridae 0 0
Stegastes planifrons Pomacentridae 0 0.02
Stegastes variabilis Pomacentridae 0 0
Priacanthus cruentatus Priacanthidae 0 0.6
Sparisoma rubripinne Scaridae 0 3.6
Scorpaena plumieri Scorpaenidae 0 0
Hypoplectrus chlorurus Serranidae 0 0
Hypoplectrus guttavarius Serranidae 0 0
Hypoplectrus indigo Serranidae 0 not present
Hypoplectrus nigricans Serranidae 0 0
Hypoplectrus puella Serranidae 0 0
Hypoplectrus unicolor Serranidae 0 not present
Rypticus saponaceus Serranidae 0 4.6
Cantherhines pullus Tetraodontiformes 0 8.6
Diodon hystrix Tetraodontiformes 0 0
Lactophrys triqueter Tetraodontiformes 0 2
Melichthys niger Tetraodontiformes 0 7
Sphoeroides spengleri Tetraodontiformes 0 0
Xanthichthys ringens Tetraodontiformes 0 4.8

indicating that serious growth over-fishing is occurring. These include the snappers O. chrysurus (71%
immature) and Lutjanus mahogoni (36% immature), surgeonfish Acanthurus coeruleus (45% immature),
parrotfish Sparisoma viride (85% immature), grunts Haemulon flavolineatum (38% immature), Haemulon
chrysargyreum (43% immature) and the goatfish Mulloidichthys martinicus (50% immature). In 12 species,
for example Pseudopeneus maculatus and C. parrae, most fish (>85%) were sexually mature when they
were caught.

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TRAP FISHING EFFECTS ON REEF FISH 123

Table 3. Trappability at the family level in St Lucia and Jamaica calculated as a ratio of a family’s abundance in traps compared to
that in fish counts. The Serranidae (*) have been split into four groups representing species of more similar life history characteristic.
Species denoted as ** were found in traps but not seen in counts

Family Trappability ratios


St Lucia Jamaica
Sparidae (porgies) 12.19 **
Kyphosidae (chub) 6.18 **
Tetraodontiformes (includes triggerfish, pufferfish, boxfish, filefish) 5.95 7.75
Chaetodontidae (butterflyfish) 4.67 0
Serranidae* } Grammistinae (soapfish) 4.07 0
Lutjanidae (snappers) 3.14 12.94
Haemulidae (grunts) 2.28 3.5
Acanthuridae (surgeonfish) 2.18 6.12
Mullidae (goatfish) 1.5 0
Serranidae* } Paranthias furcifer (creole fish) 1.19 **
Scaridae (parrotfish) 0.84 0.46
Pomacanthidae (angelfish) 0.74 2.55
Priacanthidae (cardinalfish) 0.65 0
Holocentridae (soldierfish) 0.36 0
Serranidae* } Epinephilinae (groupers) 0.36 0.18
Pomacentridae (damselfish) 0.35 0.05
Labridae (wrasse) 0.34 0.45
Bothidae (flounders) 0 0
Serranidae* } Hypoplectrus (hamlets) 0 0
Gerreidae (mojarras) 0 **
Sciaenidae (croakers) 0 **
Scorpaenidae (scorpionfish) 0 0

DISCUSSION

This study confirms that fish traps are highly unselective fishing gears, catching a wide variety of species
from many different families. Some families and species are particularly vulnerable to being caught,
including many that have traditionally been of little or no commercial value. Measures of abundance of
highly trappable families across the Caribbean showed steep declines as fishing pressure increased.
A comparison of trap catches between St Lucia and Jamaica showed that 11 species that were highly
vulnerable to trapping in St Lucia appeared to be absent from Jamaica. Since current levels of trap fishing
in Jamaica were approximately three and a half times higher than in St Lucia, and fishing there has been
intense for over 100 years (Jackson, 1997), it seems that the absence of these species is due to long-term
intensive trap fishing.
Examination of 705 fish gonads in St Lucia revealed that in 23 species constituting an important
component of fish catches, there were seven where over a third of the fish caught were immature.
Severe growth over-fishing is occurring and suggests that St Lucia’s trap fishery is wasteful and may
be unsustainable. Studies of trap fisheries in other parts of the world have reached similar conclusions.
For example, work in Kenya showed that on average 39% of the fish caught in small traps and 49% of those
in big traps were juveniles (Mangi, 2006). For one of these species (Lethrinus xanthochilus) juveniles comprised
100% of the catch while in five others, the figure was 75% or more. From St John in the US Virgin Islands,
Garrison et al. (2004) reported that most of the commercially important Balistes vetula found in traps are
immature and concluded that the island’s trap fishery had been severely over-exploited. Similarly in the

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124 J.P. HAWKINS ET AL.

Acanthurus bahianus Acanthurus coeruleus


180 35

160
30
140 30.2% 45.1%

Number of fish
Number of fish
25
120
n = 514 n = 193
100 20

80 15
60
10
40
5
20

0 0
8 10 12 14 16 18 20 22 24 26 28 6 8 10 12 14 16 18 20 22 24 26 28

Sparisoma viride Sparisoma aurofrenatum


14 60

12 50
Number of fish

10

Number of fish
85.2% 40 3.8%
8
n = 89 30
n = 235
6
20
4

2 10

0 0
10 12 14 16 18 20 22 24 26 28 30 5 10 15 20 25 30 35

35
Scarus taeniopterus 8
Sparisoma rubripinne
Min. size = 17
30
0.8% 7.1%
6
Number of fish

Number of fish

25

20 n = 124 n = 14
4
15

10
2

0 0
12 14 16 18 20 22 24 26 28 30 14 16 18 20 22 24 26

Mulloidichthys martinicus Pseudopeneus maculatus


100 14
Min. size = 14
12
80 50% 0%
Number of fish

10
Number of fish

60 n = 510 8 n = 47
6
40

4
20
2

0 0
5 10 15 20 25 30 35 14 16 18 20 22 24 26 28 30
Size (cm) Size (cm)
Figure 5. Size frequency distributions of species commonly caught in traps in St Lucia, from a sample of 530 fish catches. Arrows mark
fish size at first maturity. Data for maturity were obtained from a sample of 705 fish catches that were not included in the size frequency
distributions. ‘Min. size’ also describes fish size at first maturity and is indicated on graphs where individuals of this length were
absent from catches analysed for size frequency distributions. Percentages shown indicate the fraction of fish landed below the size
at first maturity.

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TRAP FISHING EFFECTS ON REEF FISH 125

Haemulon carbonarium Haemulon chrysargyreum


4 35

30

3 6.6% 43.4%

Number of fish
25
Number of fish

n = 15 20
n = 83
2
15

10
1
5

0 0
14 16 18 20 22 24 10 12 14 16 18 20

50
Haemulon flavolineatum 4
Haemulon parrai
Min. size = 16
40
3
Number of fish

Number of fish
38.2% 0%
30
2
20
n = 136 n = 11

1
10

0 0
8 10 12 14 16 18 20 22 18 19 20 21 22 23 24 25 26

Haemulon plumieri Haemulon aurolineatum


8 100

80
6
10.3% 25%
Number of fish

Number of fish

60
4 n = 29 n = 436
40

2
20

0 0
12 14 16 18 20 22 24 26 28 30 32 34 6 8 10 12 14 16 18 20 22 24

Ocyurus chrysurus Lutjanus mahogoni


25 16

14
20
70.9% 12
36%
Number of fish

Number of fish

15 10

n = 179 8 n = 114
10 6

4
5
2

0 0
15 20 25 30 35 40 45 50 55 10 15 20 25 30 35

Size (cm) Size (cm)

Figure 5. continued

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DOI: 10.1002/aqc
126 J.P. HAWKINS ET AL.

Cantherhines pullus 20
Clepticus parrae
14
4.7% 0%
12

n = 43 15 n = 49

Number of fish
Number of fish 10

8
10
6

4
5

0 0
10 15 20 25 30 35 16 18 20 22 24 26

8
Cephalopholis cruentata 12 Cephalopholis fulva

10
6 12.2% 20%

Number of fish
Number of fish

n = 82 n = 25
4 6

4
2
2

0 0
10 15 20 25 30 35 40 12 14 16 18 20 22 24 26

180
Holocentrus rufus 7
Holocentrus adscensionis
160
6
140
Number of fish
Number of fish

5
120 4.9% 20%
100 4

80 n = 597 3 n = 30
60
2
40
1
20

0 0
8 10 12 14 16 18 20 22 24 10 15 20 25 30 35

Size (cm)

Myripristis jacobus
250

200
4.7%
Number of fish

150
n = 701
100

50

0
8 10 12 14 16 18
Size (cm)

Figure 5. continued

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TRAP FISHING EFFECTS ON REEF FISH 127

Table 4. Catch composition from fish traps in St Lucia showing fish size at first maturity and where possible the size at 50% and 100%
maturity

Species Sample size Minimum size Range Size at which Size at which
at maturity 50% mature 100% mature
Acanthurus bahianus 38 14 12–19 16
Acanthurus coeruleus 20 15 11–21 15 15
Scarus taeniopterus 38 15 14–24
Sparisoma aurofrenatum 67 14 14–23
Sparisoma rubripinne 43 17 17–31 22
Sparisoma viride 10 25 15–25
Cephalopholis cruentata 50 16 12–29 26 26
Cephalopholis fulva 23 19 12–25
Lutjanus mahogoni 14 20 18–34
Ocyurus chrysurus 19 32 17–43 32 32
Haemulon aurolineatum 24 14 14–17 16 17
Haemulon carbonarium 22 16 15–22
Haemulon chrysargyreum 21 15 15–18
Haemulon flavolineatum 42 15 13–19 19
Haemulon parrai 55 16 13–20
Haemulon plumieri 23 17 14–28 25 25
Holocentrus adscensionis 46 17 13–24 19 20
Holocentrus rufus 34 13 13–17 14 15
Myripristis jacobus 29 11 10–15 11 14
Pseudupeneus maculatus 19 14 13–21
Mulloidichthys martinicus 23 20 13–25
Cantherhines pullus 17 13 13–20
Clepticus parrae 21 17 17–24 17 17

Arabian Gulf the high incidence of juvenile catch in traps has led to a call for fishery reform (Grandcourt
et al., 2004).
Comparing the species that were most susceptible to trapping in St Lucia and Jamaica revealed
differences between these islands. Trappability ratios indicated that the Tetraodontiformes were highly
susceptible to trapping in both Jamaica and St Lucia, and butterflyfish were highly susceptible in St Lucia,
but not in Jamaica. Long-term intensive exploitation could eliminate species susceptible to trapping, and
this has probably happened to butterflyfish and several of the Tetraodontiformes that were highly trappable
in St Lucia but rare or absent from Jamaica. Although, the results from Jamaica are based on a relatively
small sample size of 23 traps and 112 counts, a study by Sary (1995) in the same area a year earlier involved
sampling 886 traps. Comparing our list of highly trappable species in St Lucia, with the species that Sary
found in traps in Jamaica reveals that two-thirds of St Lucia’s highly trappable species were missing from
traps in Sary’s study. This suggests that these species: H. striatum, C. calamus, M. interstitialis, A. scriptus,
H. melanurum and K. sectatrix, really could have been fished out from this part of northern Jamaica and
that our findings are not simply an artefact of under-sampling. It is already acknowledged that once-
abundant species of snappers, groupers and large parrotfish are virtually absent on Jamaica’s reefs (Koslow
et al., 1994). Furthermore, reducing St Lucia samples to Jamaican levels left 8.5 of 11 highly trappable
species still represented in St Lucia.
Soldierfish had low trappability ratios in both Jamaica and St Lucia, but were much more abundant in St
Lucia than in Jamaica. Since this study showed trapping pressure in Jamaica to be around three and a half
times greater than in St Lucia, it is possible that even families which can usually withstand intensive fishing
will become over-exploited if trapping reaches extremely high levels.

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128 J.P. HAWKINS ET AL.

It is important to be cautious in attributing among-island differences in fish assemblages to differences in


fishing pressure. There are other factors among islands besides fishing that could have played a role. For
example, islands differ in size, relief, human population densities and level of development, and these
may influence processes on adjacent reefs. They also differ in terms of scale and history of human and
other impacts, including coral disease outbreaks, hurricanes, watershed clearance and nutrient input, to
name a few (e.g. see Gardner et al., 2003). However, findings from the study uphold many predicted
effects of fishing (Polunin and Roberts, 1996). Other factors, such as degree of watershed development,
although not tested statistically, do not appear to relate in obvious ways to inter-island differences
in fish abundance or assemblage composition, nor are mechanisms by which they might exert these
effects clear. One possible exception is that coral cover and habitat structural complexity decrease on the
study reefs as fishing pressure increases (Hawkins and Roberts, 2004), which may affect reef carrying
capacity and favour some species over others. Hawkins and Roberts (2003) argue that this habitat shift is in
itself partly a consequence of overfishing of herbivores. Biomass of herbivores decreases 10-fold over this
fishing pressure gradient, from Bonaire to Jamaica, with a consequent reduction in grazing pressure on
algae. Without undertaking experimental manipulations of fishing pressure it is not possible to rule out
influences other than exploitation, but our findings clearly show that trap fishing has the capacity to alter
fish assemblage composition, and historical fishery data suggest that it already has in Jamaica (Koslow
et al., 1994).
Some species may be highly trappable because of behavioural traits. For example, most butterflyfish
form strong pair bonds (Roberts and Ormond, 1992), so if one member of a pair goes into a trap its
partner is likely to follow. Such behaviour would double the likelihood of a species getting trapped
and indeed Chaetodon striatus (trappability ratio 9.8) and Chaetodon ocellatus (trappability ratio 7.3)
were highly trappable in St Lucia. Similarly, species which move around in groups or schools might
follow each other into traps. Highly trappable species which this might apply to in Jamaica include
Acanthurus chirugus (trappability ratio 112), Acanthurus coeruleus (trappability ratio 12) A. bahianus
(trappability ratio 3.6) and Sparisoma chrysopterum (trappability ratio 18), and, in St Lucia, Melichthys
niger (trappability ratio 7), K. sectatrix (trappability ratio 6.2) and Sparisoma rubripinne (trappability ratio
3.6). Other species may be highly trappable simply because they are curious and readily enter enclosed
spaces.
Differences in fish behaviour between St Lucia and Jamaica could also help explain why trappability
ratios differed between these countries. In Jamaica, species such as Balistes vetula and Calamus calamus,
tended to avoid our counting area but were less timid in St Lucia. Wary behaviour towards people is often
encountered in places that are intensively spear-fished and, from personal observations, spear-fishers were
much more common in Jamaica than St Lucia.
Species that grow slowly and reproduce late may be extirpated by trapping even when trapping pressure
is relatively light. Groupers and snappers contain species that are prime examples of animals whose life
history characteristics (e.g. late maturity) render them particularly vulnerable to over-exploitation
(Reynolds, 2003). In St Lucia such species with high trappability ratios were the groupers M. interstitialis
(trappability ratio: 11.2) and Epinephelus guttatus (trappability ratio: 9.1) and the snappers L. synagris
(trappability ratio: 439), O. chrysurus (trappability ratio: 3.3) and L. mahogoni (trappability ratio: 3.3).
L. synagris had the highest trappability ratio of any species in St Lucia. This species was rare in counts and
may have moved into traps at night after coming up from deeper water (www.fishbase.org). The
trappability ratio of L. synagris thus probably represents an overestimate of the difference in abundance
between reef and traps for this species.
Trap fishing is widely perceived as a relatively benign method whereby small-scale fishers can
obtain a living from coral reefs. However, it is clear from the steep gradients in fish abundance across
our inter-island fishing pressure gradient that fishing, particularly with traps, has greatly affected Caribbean
fish assemblages. Historical data from the region place this inter-island comparison in a different

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TRAP FISHING EFFECTS ON REEF FISH 129

perspective, showing steep declines in high-value, vulnerable species over time. In Puerto Rico, for
example, there were 1403 fishers in 1931, setting 4239 traps and catching 1400 tonnes of reef fish
(Appeldoorn et al., 1992). By 1990, trap numbers climbed to 13 555, while catches fell to 854 tonnes.
Most of the catch in 1931 was composed of Nassau groupers, Epinephelus striatus and other high-value
snappers and groupers. By 1990, more than half of the catch was composed of low-value species such as
grunts, surgeonfish and parrotfish. Similar trends of increasing fishing effort, falling catches and
disappearance of high-value species have been seen over much of the region, including the US Virgin
Islands (Rogers and Beets, 2001), Florida (Ault et al., 1998) and Jamaica (Jackson, 1997). The perception
that trap fishing is a benign fishing method has perhaps permitted these fisheries to operate with
little regulation or control. There is an urgent need for reappraisal, such as that under way in Florida
where traps are due to be phased out by 2007 (http://www.reefguardian.org/Campaigns/FishTrapGOM/
MoreInfoFTrapGOM.html).
Species with high trappability ratios may not necessarily be highly threatened by fishing, even in places
where trap fishing is intense. Species which are small at maturity, have short life-cycles and high turnover
rates might be able to withstand being highly trappable because their life history characteristics enable them
to persist despite high rates of fishing mortality. In countries where levels of trap fishing are high these
species may constitute the majority of catches. However, intensive fishing may threaten sustainability of
even these species eventually. Mesh size limits help reduce the impact of trapping on fish populations by
allowing juvenile fish to escape capture until after they reach maturity (Hunte and Mahon, 1994). They can
also increase catches by reducing growth over-fishing. Work by Sary et al. (1997), done at the same place in
Jamaica as this study and at a similar time, showed that fishery yields increased when traps with 2.5 cm
and 3.2 cm meshes were replaced with ones of a larger size of 3.8 cm mesh. However, meshes will need
to be bigger than this if large species with late maturity and a susceptibility to trapping are to gain any
protection. One way round this problem could be to introduce slots, known as ‘escape gaps’, into traps that
are larger than the mesh size. These help provide an opportunity for juveniles to escape (Stewart and
Ferrell, 2002).
In summary, certain taxa such as the Tetraodontiformes appear to be particularly vulnerable to
trapping. Whatever reasons lure fish into traps and regardless of the fact that some species appear more
trappable than others, trapping is a highly non-selective form of fishing. If populations of vulnerable
species are to be maintained on reefs, and more resilient species are to support productive and sustainable
fisheries, then trapping needs to be regulated. This can include measures such as limiting numbers of trap
fishers and how many traps each can use, increasing mesh sizes and introducing escape gaps to traps.
Education could also help reduce waste from trap fisheries. For example, fishers could be encouraged to
throw back the small fish they catch from shallow water. Given that fisheries regulations are often difficult
to enforce in areas with trap fisheries, marine reserves that are off-limits to fishing probably provide the
easiest and most effective way to tackle the problems associated with trap fishing, particularly if used in
association with the other forms of management discussed (Murray et al., 1999; Roberts and Hawkins,
2000; Russ, 2002; Gell and Roberts 2003). Even in areas where trapping pressure is fairly low, marine
reserves will provide essential refuges for species that are particularly vulnerable to over-exploitation
(Roberts, 2005).

ACKNOWLEDGEMENTS

We would like to thank the UK Natural Environmental Research Council, UK Department for International
Development, University of Puerto Rico Seagrant and USAID for funding this work. Special thanks to the Bonaire
Marine Park, Saba Marine Park and Scuba St Lucia for generous use of their facilities during field trips. Erkki Siirila,
Harold Guiste, Daniel Matos, Graciela Garcia-Moliner, Michelet Fontaine and John Clark helped us track down
information on fishing pressure. Thanks to John Baxter and two reviewers whose comments greatly improved the
manuscript.

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
130 J.P. HAWKINS ET AL.

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Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc
132 J.P. HAWKINS ET AL.

APPENDIX: SPECIES REMOVED FROM TRAPPABILITY ANALYSIS (ARRANGED IN


ALPHABETICAL ORDER)
Species Size (cm) Cryptic
Amblycirrhitus pinos 9 Yes
Apogon maculates 9 Yes
Aulostomus maculatus 75 No
Bairdiella sanctaeluciae 15 Yes
Canthigaster rostrata 9 No
Centropyge argi 6 Yes
Centropyge aurantonotus 6 Yes
Channomuraena vittata 120 Yes
Chromis cyanea 10 No
Chromis insolata 9 No
Chromis multilineata 14 No
Cryptotomus roseus 9 Yes
Echidna catenata 45 Yes
Enchelycore carychroa 25 Yes
Enchelycore nigricans 60 Yes
Equetus acuminatus 20 Yes
Equetus lanceolatus 20 Yes
Equetus punctatus 23 Yes
Fistularia tabacaria 120 No
Ginglymostoma cirratum 270 No
Gramma loreto 6 Yes
Gymnothorax funebris 150 Yes
Gymnothorax miliaris 45 Yes
Gymnothorax moringa 90 Yes
Halichoeres bivittatus 18 No
Halichoeres cyanocephalus 11 No
Halichoeres garnoti 15 No
Halichoeres maculipinna 13 No
Halichoeres pictus 14 No
Halichoeres poeyi 14 No
Hemiemblemaria simulus 9 Yes
Hemipteronotus martinicensis 9 Yes
Heteroconger halis 38 Yes
Hippocampus erectus 10 Yes
Inermia vittata 18 No
Liopropoma carmabi 4 Yes
Liopropoma rubre 8 Yes
Lucayablennius zingaro 4 Yes
Malacanthus plumieri 45 No
Megalops atlanticus 120 No
Monacanthus tuckeri 9 Yes
Myrichthys breviceps 75 Yes
Myrichthys ocellatus 75 Yes
Narcine brasiliensis 38 No
Odontoscion dentax 19 Yes
Ophichthus ophis 75 Yes
Opistognathus aurifrons 8 Yes
Pempheris schomburgki 13 Yes
Scorpaenodes caribbaeus 10 Yes
Serranus baldwini 5 Yes
Serranus tabacarius 11 No
Serranus tigrinus 9 No
Sparisoma atomarium 10 Yes
Sparisoma radians 13 Yes
Stegastes partitus 9 No
Synodus intermedius 35 No
Synodus saurus 18 No
Thalisoma bifasciatum 13 No
Urolophus jamaicensis 30 No

Copyright # 2006 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. 17: 111–132 (2007)
DOI: 10.1002/aqc

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