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7
Bull. N.J. Acad. Sci., 53(2), pp. 7–11.
© 2008, by the New Jersey Academy of Science
Assessment of Fish and Decapod Distributions Between Mangrove and
Seagrass Habitats in St. John, U.S.V.I.
Charles C. Kontos1 and Paul A. X. Bologna2
1Department of Biology and Molecular Biology, Montclair State University, Montclair, NJ 07043
2Aquatic and Coastal Sciences Program, Department of Biology and Molecular Biology,
Montclair State University, Montclair, NJ 07043, email: bolognap@mail.montclair.edu
Abstract: Fish and decapod community structure and catch per unit effort
(CPUE) were assessed within a mangrove previously impacted by a hurricane and
an adjacent seagrass bed in St. John, United States Virgin Islands. Diel sampling
utilizing minnow traps showed signicantly greater total sh CPUE in the seagrass
habitat with few individuals using the mangrove regardless of the time of day.
Decapods showed no difference in CPUE and were equally distributed between
mangrove and seagrass habitats. When individual species were analyzed, Haemulon
avolineatum, H. sciurus, Calcinus elegans and Panulirus argus had signi-
cantly higher CPUEs in seagrass beds compared to the mangrove, while Panopeus
occidentalis showed the opposite distribution. Panulirus argus and C. elegans had
signicantly higher CPUEs during the night, while Eurypanopeus abbreviatus was
signicantly more abundant during the day. A Similarity of Percentages analysis
showed low species similarity among individual habitat-diel samples (<25%), but
strong dissimilarities among all sampling combinations (>90%) and an Analysis
of Similarity showed signicant differences in the fauna among each habitat-diel
conguration. These analyses reect the minimal use of the mangrove by sh and the
strong diel habitat differences for individual decapod species including an example
of potential niche partitioning between two mud crabs species in the mangrove.
Keywords: mangrove, Rhizophora mangle, seagrass, Thalassia
testudinum, niche partitioning
INTRODUCTION
Coastal tropical systems are often dominated by a mix of mangrove,
seagrass and coral reef habitats. Each of these habitats is important
as refuge and feeding regions for a variety of sh (Nagelkerken et al.
2000, Dorenbosch et al. 2004) and invertebrates and frequently they
are linked through trophic transfer (Baelde 1990). As coastal systems
are impacted by natural and anthropogenic disturbances, the value
and linkages among these habitats may often be altered or degraded.
Human disturbances, such as over-shing, habitat destruction, and
eutrophication, generally lead to signicant or permanent ecosystem
damage (Fondo and Martens 1998), while natural disturbances, such
as hurricanes, are generally followed by natural recovery.
The value of seagrasses and mangroves as nursery habitat has been
well documented (see Heck et al. 1997, Faunce and Serafy 2006, and
references within). Primary productivity in these systems is directly
consumed (Kirsch et al. 2002, Feller and Chamberlain 2007), but a
substantial portion of this production enters detrital pathways (Ce-
brian et al. 1997, Lee 1999). High levels of secondary production
lead to substantial trophic transfer to higher level consumers (Wolff
et al. 2000). Often, movement among these habitats leads to energy
export to coral reef communities; as juveniles take up residence on
the reefs and also through daily foraging of adults into mangroves
and seagrass beds (Nagelkerken et al. 2000). Consequently, commu-
nity disturbance may reduce overall productivity of a coastal system
(Wolff et al. 2000). This is especially evident when mangroves are
destroyed for aquaculture, because the resultant loss of habitat leads
to increased erosion and turbidity, loss of essential habitat for all sh
and invertebrates previously utilizing this habitat and subsequent
declines in adjacent seagrass and coral reef communities (Primavera
2006). Less understood is how the faunal communities change and
recover following a natural disturbance, such as a hurricane.
This research focused on a mangrove-seagrass habitat complex in
the northern part of Great Lameshur Bay, U.S. Virgin Islands that had
been severely impacted by two hurricanes (Hugo 1989, Marilyn 1995).
While the seagrass beds have recovered (Kendall et al. 2001), the man-
grove has seen very limited natural recovery (Nemeth et al. 2004). Our
objective was to investigate the diel usage patterns of the mangrove and
adjacent seagrass habitat by sh and decapods to determine whether
the limited recovery of the mangrove was providing usable habitat for
these organisms. Little was known regarding the use and value of this
particular mangrove community prior to the hurricane, but currently
this system remains substantially degraded with diminutive re-growth
of the mangrove interior (Kendall et al. 2001).
MATERIALS AND METHODS
Great Lameshur Bay is inside the boundaries of Virgin Islands National
Park, a United Nations Biosphere Reserve. The region is characterized
by several communities including seagrass beds, coral reefs, mangrove
forests, and unvegetated zones covering a total area of roughly 50
acres (Kendall et al. 2001). Our experiment was conducted among
the prop roots of a red mangrove (Rhizophora mangle) forest and the
adjacent seagrass (Thalassia testudinum) habitats within the bay. The
mangrove forest was damaged during Hurricanes Hugo (1995) and
Marilyn (1995) causing severe internal damage resulting in piles of
dead mangrove trees and large regions of unvegetated mudat (Nem-
eth et al. 2004, Bologna pers. obs.).
8 Charles C. Kontos and Paul A.X. Bologna
Minnow traps were used to sample juvenile sh and decapods in
each habitat. The traps were constructed of 3.2 mm seine net and
a steel wire frame with two 5.1 cm diameter door openings at each
end. The traps measured 25.4 cm × 25.4 cm × 43.2 cm and had two
zippered pockets for bait and emptying catch along with a 3.7 m
drop cord. Six traps were placed within the red mangrove prop roots
by kayaking along the mangrove channel and six traps were placed
into the adjacent Thalassia testudinum bed. Squid was used to bait
traps during both day and night sampling events. Traps were set in
the morning (sunrise) and retrieved approximately 11–11.5 hours
later (sunset) and then reset to assess nighttime utilization of the two
habitats. Traps were deployed for 3 daily cycles in January 2006. Fish
and decapods from retrieved traps were enumerated, measured, and
identied to their lowest taxonomic level and then released. Data were
standardized for trap shing time to generate a catch per unit effort
(CPUE) by dividing the abundance of organisms by trap collection
time (# sh, decapods/ # hours sampled). To assess habitat use and
diel differences among the common sh and decapods, CPUE was
analyzed using a 2-way ANOVA with habitat (i.e., seagrass, mangrove)
and diel stage (day, night) as independent variables with an α = 0.05.
The four habitat-diel stage combinations were further assessed by
conducting an Analysis of Similarities (ANOSIM, Bray-Curtis matrix
on non-transformed data) and Similarity of Percentages (SIMPER,
non-transformed data) analysis using the PRIMER software package
(Clarke and Warwick 1994) on the species specic CPUE to determine
how related the sampled faunal communities were.
RESULTS
Fish
Traps collected eight sh species: French grunt (Haemulon avolinea-
tum), schoolmaster (Lutjanus apodus), bluestriped grunt (Haemulon
sciurus), yellowtail snapper (Ocyurus chrysurus), slippery dick (Hal-
ichoeres bivittatus), marked goby (Ctenogobius stigmaticus), squirrelsh
(Holocentrus adscensionis), and damselsh (Stegastes adustus) (Table 1).
Signicantly more sh used seagrass habitat than mangrove (F1,64 =
13.04, P <0.0006, Fig. 1a), but utilization was similar between day and
night. Haemulon avolineatum were signicantly more abundant in
seagrass beds (F1,64 = 15.8, P <0.002) as were H. sciurus (F1,64 = 5.92, P
<0.02). No other sh showed signicant afliations with either habitat.
Average sh length indicated that primarily juvenile H. avolineatum,
H. sciurus, and O. chrysurus were using these habitats (Table 1).
Decapods
Decapods were represented by ten species including common mud
crab (Panopeus occidentalis), lobate mud crab (Eurypanopeus abbre-
viatus), blue crab (Callinectes sapidus), spiny lobster (Panulirus argus),
ocellate swimming crab (Portunus sebae), peppermint shrimp (Lysmata
wurdemanni), grass shrimp (Palaemonetes pugio), snapping shrimp (Al-
pheus heterochaelis), sponge decorator crab (Stenocionops furcata), and
blue-legged hermit crabs (Calcinus elegans) (Table 1). Total decapod
CPUE was not different between habitats (F1,64 = 0.08, P <0.77) or
time of collection (F1,64 = 1.32, P <0.26; Fig. 1b). Despite this lack of
overall habitat or time effect at the decapod community level, several
Table 1. Species Specic Catch Per Unit Effort (CPUE). Values represent the mean CPUE ± 1 SE for all species identied from this study. Mean size values provided
for sh are standard length values ± 1 SE, crabs carapace width ± 1 SE, and shrimp/lobsters carapace length ± 1 SE. NR represent values not recorded.
Mangrove Seagrass
FISH Species Night Day Night Day Mean Size (cm)
Ctenogobius stigmaticus 0.007 ± 0.005 0 0 0 3.60 ± 1.27
Haemulon avolineatum 0.014 ± 0.014 0 0.265 ± 0.105 0.212 ± 0.098 8.78 ± 1.95
Haemulon sciurus 0 0 0.015 ± 0.01 0.019 ± 0.014 10.18 ± 0.95
Halichoeres bivittatus 0 0 0.023 ± 0.16 0 9.97 ± 0.40
Holocentrus adscensionis 0 0 0 0.006 ± 0.006 14.60
Lutjanus apodus 0.027 ± 0.015 0.006 ± 0.006 0 0 13.59 ± 2.39
Ocyurus chrysurus 0.003 ± 0.003 0 0 0.012 ± 0.008 6.77 ± 4.22
Stegastes adustus 0.003 ± 0.003 0 0 0 9.40
DECAPOD Species
Alpheus heterochaelis 0.003 ± 0.003 0 0 0 3.90
Calcinus elegans 0.003 ± 0.003 0 0.182 ± 0.062 0 NR
Callinectes sapidus 0.007 ± 0.005 0.003 ± 0.157 0 0 6.6 ± 2.26
Eurypanopeus abbreviatus 0.014 ± 0.014 0.076 ± 0.021 0 0.033 ± 0.027 2.23 ± 0.79
Lysmata wurdemanni 0 0 0.008 ± 0.008 0 3.30
Palaemonetes pugio 0.151 ± 0.082 0.02 ± 0.015 0.008 ± 0.008 0.076 ± 0.069 NR
Panopeus occidentalis 0.051 ± 0.019 0.03 ± 0.018 0.015 ± 0.01 0 2.60 ± 0.55
Panulirus argus 0 0 0.03 ± 0.013 0 8.92 ± 1.10
Portunus sebae 0 0 0 0.013 ± 0.009 4.47 ± 0.93
Stenocionops furcata 0 0 0.008 ± 0.008 0 0.70
Fish and Decapod use of Tropical Habitats 9
individual species exhibited habitat or diel preferences. Specically,
signicantly more P. occidentalis (F1,64 = 4.3, P <0.04) were collected
in mangroves compared to seagrass beds, while C. elegans and P. argus
were signicantly more abundant in seagrass habitat (F1,64 = 13.28,
P <0.0005, F=9.56, P <0.0061, respectively). Eurypanopeus abbrevia-
tus were signicantly more abundant during the day than at night
(F1,64=6.86, P <0.01), while C. elegans and P. argus were signicantly
Table 3. SIMPER Community Comparisons among the habitat-diurnal treatments. Values represent the individual percent contribution and the cumulative
contribution to dening the fauna responsible for the relationship.
Seagrass Day (SD) Seagrass Night (SN) Mangrove Day (MD) Mangrove Night (MN)
SD
Average SD Similarity 9.99
Ind.% Cum.%
H. avolineatum 84.9 84.9
P. sebae 9.4 94.3
SN
Average Dissimilarity 90.4%
Ind.% Cum.%
H. avolineatum 38.6 38.6
C. elegans 25.9 64.5
P. pugio 7.1 71.6
P. argus 6.8 78.4
E. abbreviatus 4.4 82.7
P. sebae 4.4 87.1
H. sciurus 4.3 91.4
Average SN Similarity 23.9
Ind.% Cum.%
C. elegans 51.5 51.5
H. avolineatum 39.6 91.1
MD
Average Dissimilarity 97.5
Ind.% Cum.%
H. avolineatum 29.1 29.1
E. abbreviatus 25.2 54.2
P. pugio 13.1 67.3
C. sapidus 9.4 76.6
P. occidentalis 7.9 84.6
P. sebae 6.4 91.0
Average Dissimilarity 99.8%
Ind.% Cum.%
C. elegans 26.3 26.3
H. avolineatum 25.9 52.3
E. abbreviatus 15.8 68.1
P. argus 6.8 74.9
C. sapidus 6.8 81.7
P. occidentalis 6.1 87.8
P. pugio 3.7 91.5
Average MD Similarity 18.1
Ind.% Cum.%
E. abbreviatus 79.3 79.3
C. sapidus 12.9 92.3
MN
Average Dissimilarity 97.3%
Ind.% Cum.%
H. avolineatum 31.6 31.6
P. pugio 22.9 54.6
P. occidentalis 12.3 66.8
P. sebae 7.7 74.5
E. abbreviatus 6.2 80.8
O. chrysurus 4.3 85.1
L. apodus 4.2 89.3
H. sciurus 3.5 92.9
Average Dissimilarity 98.2
Ind.% Cum.%
C. elegans 27.4 27.4
H. avolineatum 26.7 54.1
P. pugio 12.4 66.6
P. occidentalis 9.5 76.0
P. argus 8.0 84.0
L. apodus 3.3 87.3
S. furcata 3.2 90.5
Average Dissimilarity 94.2
Ind.% Cum.%
E. abbreviatus 28.5 28.5
P. pugio 22.2 50.7
P. occidentalis 21.4 72.1
C. sapidus 13.1 85.1
L. apodus 6.6 91.7
Average MN Similarity 8.5%
Ind.% Cum.%
P. occidentalis 48.4 48.4
P. pugio 41.3 89.7
L. apodus 6.8 96.5
Table 2. ANOSIM comparisons among the habitat-diurnal treatments.
Comparison Pair Groups R Statistic Signicance
Level
Seagrass Day vs. Seagrass Night 0.16 0.028
Seagrass Day vs. Mangrove Day 0.29 0.001
Seagrass Day vs. Mangrove Night 0.296 0.001
Seagrass Night vs. Mangrove Day 0.507 0.001
Seagrass Night vs. Mangrove Night 0.449 0.001
Mangrove Day vs. Mangrove Night 0.238 0.002
more abundant at night (F1,64=12.53, P <0.0008, F=8.05, P <0.0061,
respectively).
Community Response
Fish and decapod communities varied substantially in the use of each
of the habitat-diel category (Table 2). Two important features were
apparent. First, the low similarity percentage for each habitat-diel
category (8–24% SIMPER) suggests that there is substantial variability
among the replicates present in the data set, but also that only a few
species contributed greatly to the overall response. Second, the very
high dissimilarity percentages (90–99.8%) demonstrate that the faunal
communities using each habitat-diel conguration are substantially
different (Table 3).
DISCUSSION
It is well known that diel changes in species composition occur in
reef-associated sh populations such as Haemulidae, Lutjanidae, and
Apogonidae as a result of inter-habitat movement and foraging (Wein-
stein and Heck 1979, Nagelkerken et al. 2000, Unsworth et al. 2007).
10 Charles C. Kontos and Paul A.X. Bologna
Additionally, sh species assemblages within seagrasses contain similar
families throughout the world (Pollard 1984). While higher abundance
and diversity of Lutjanids (snappers) have been found in Thalassia
testudinum habitats near coral reefs than near mangroves (Baelde 1990),
we found that Lutjanus apodus was found only in mangroves, but
Ocyurus chrysurus was found in both habitats (Table 1). More juvenile
Haemulon avolineatum and H. sciurus utilized seagrass habitats and
their presence dominated the sh fauna captured. Juveniles of these
two grunts species are also dominant in barrier reef lagoons in Be-
lize (Sedberry and Carter 1993), while newly settled Haemulidae are
among the most abundant species found in the backreef embayments
of nearby St. Croix, U.S.V.I. (Mateo 2002). The migration of juvenile
grunts is photomechanically attuned to light intensities suggesting that
they are crepuscular (McFarland et al. 1979). In the backreef areas of
Belize, H. avolineatum were found mostly in sand ats at night, while
H. sciurus were found in seagrass beds at night indicating that these
sh may be leaving the reef at dusk to forage in these adjacent habitats
(Burke 1995). It is interesting to note that H. avolineatum and H.
sciurus were found almost exclusively in the seagrass, but distributed
equally in the day and night (Table 1). This suggests that without the
mangrove habitat as an alternate, these species utilized the seagrass
bed to a much higher extent than expected.
While the value of mangroves as sh habitat has been demonstrated
(Faunce and Serafy 2006), our results showed signicantly more sh
were collected within seagrass compared to mangrove habitat during
both day and night sampling periods (Fig. 1a). This nding may largely
be attributed to the reduced structural habitat and the highly turbid,
warm, and hypoxic conditions present within the hurricane impacted
mangrove. In fact, temperature exceeding 34°C and dissolved oxygen
concentrations below 5 mg/l during the day were observed (Bologna
unpubl. data). These environmental conditions could certainly curtail
the use of the mangrove, especially for vertebrates with higher respira-
tion demands. Furthermore, unlike decapods, many reef-associated
sh species may be unable to tolerate the higher temperatures and
hypoxic conditions of the hurricane-impacted mangrove, especially
during the day, and our results support the near absence of juvenile
sh use of the mangrove during that time period (Fig.1a). Schools of
large tarpon were also observed feeding along the edge of the turbid-
ity plume exiting the mangroves during daytime ebbing tide (Bologna
pers. obs.) potentially inuencing the presence of sh in this area
indicating that anti-predator behavior may be affecting the use of the
mangrove (sensu Abrahams and Katteneld 1997).
Decapod species composition, however, varied extensively based
on habitat type and diel sampling period with more decapods being
collected at night, but with similar distributions between mangrove
and seagrass habitats (Fig. 1b). The most abundant decapod spe-
cies, Palaemonetes pugio, utilized predominantly mangrove habitat at
night, but it was present in seagrass habitat during the day (Table 1).
This suggests a diel migration into the mangrove to feed and active
refuge seeking during the day within the seagrasses. The second most
abundant species, Calcinus elegans, preferred nocturnal foraging within
seagrass habitat with 96% of the individuals being collected during this
time frame. Juvenile Panulirus argus showed a similar pattern of being
only collected at night, within the seagrass beds (Table 1). Panulirus
argus is strongly nocturnal (Cox et al. 1997) especially in regards to
juvenile movement and feeding.
One unique nding relates to the potential habitat segregation or
niche partitioning between two species of mud crabs in the mangrove.
Specically, Panopeus occidentalis was signicantly more abundant in
mangroves and dominated the mud crab abundance at night there
(80% CPUE, mud crabs), while Eurypanopeus abbreviatus dominated the
mangrove during the day (70% CPUE). This apparent diel segregation
of the mangrove may limit interspecic competition between them.
Other species of these genera exhibit a similar relationship. Speci-
cally, P. herbstii and E. depressus appear to have different life histories
with respect to size of individual and diet, which allows them to co-
occur in intertidal oyster reefs (McDonald 1982) and Meyer (1994)
demonstrated that they partitioned the intertidal oyster reef habitat
in North Carolina. Our results indicate a similar niche partitioning
potential between P. occidentalis and E. abbreviatus, but further research
is needed to ascertain this.
Species assemblages of tropical marine ecosystems can be indica-
tive of changing or altered environmental conditions. In this study,
several sh and decapod species varied considerably based on time
of day and habitat type. Considering the degraded condition of the
mangrove habitat, the reduced abundance and species composition
detected during our sampling period compared to other studies (see
Pinto and Punchihewa 1996, Dorenbosch et al. 2004) was not unex-
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
0.45
Seagrass Day Seagrass Night Mangrove Day Mangrove Night
Habitat-Diurnal Category
CPUE (+ SE)
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
0.45
Seagrass Day Seagrass Night Mangrove Day Mangrove Night
Habitat-Diurnal Category
CPUE (+ SE)
Figure 1. Total Catch per Unit Effort (CPUE) for sh and decapods collected in experimental traps. Values represent mean CPUE ± 1 SE. a. Fish. b. Decapods.
Fish Decapods
Fish and Decapod use of Tropical Habitats 11
pected. Our community assessment indicated that signicant differ-
ences existed in the fauna using the mangrove and seagrass habitat
as well as diel usage (Table 2). While we have been able to document
a divergence in faunal utilization of these habitats, more research
into how these differences impact communities and trophic transfer
is necessary to assess the recovery and functioning of the mangrove.
For each habitat-time conguration, we documented low similarity
among replicates and these relationships were dominated by two or
three species (Table 3). Not surprisingly, each habitat conguration
showed extremely high dissimilarities (90–99.8%), with the greatest
dissimilarity being seen between mangrove day and seagrass night,
where nocturnal foraging of C. elegans in seagrass and the lack of H.
avolineatum in mangroves dominated the relationship (Table 3). The
mangrove is functioning as a habitat for decapods, but it is providing
little value to the juvenile sh in this bay. Given the original impact
year of 1985 and the lack of interior recovery, it may be many more
years before this mangrove is contributing to the sh abundance and
productivity of Great Lameshur Bay.
One primary issue in coastal systems is understanding how the loss
of habitats impacts the communities of organisms previously using
them. While it is clear that human impacts on systems like mangroves
detrimentally affect the associated fauna (Wolff et al. 2000), our un-
derstanding of how natural disturbances inuence the fauna and the
potential route toward recovery is still unclear. Long-term monitoring
of these sites is necessary to resolve the return to functionality of these
valuable communities. Unfortunately, many of these communities are
also under threat from anthropogenic pressures and recovery may
never occur naturally. Due to the protected nature of this Biosphere
Reserve, long-term monitoring and assessment on this site are pos-
sible and we can evaluate the potential recovery trajectory for use of
the mangroves by sh and decapods.
ACKNOWLEDGMENTS
We would like to acknowledge Daniel Ward for his instrumental role
throughout the eld portion of this study; without his perseverance,
effort, and expertise this project could not have been completed. We
would also like to thank Cathleen Dale, Rita Papagian, Suzann Regetz,
and Taryn Townsend for their assistance and recommendations which
have proved especially valuable in the completion of this manuscript.
We would like to thank Rafe Boulon with the Virgin Islands National
Park for administrative support of this project. Last but not least,
we would like to thank the volunteers and staff at the Virgin Islands
Environmental Resource Station for their jovial support during this
eld study.
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Notes
The Bulletin.
Dr. Michael J. Kennish, Institute of
Marine and Coastal Sciences, Rutgers University, New Brunswick,
New Jersey 08901 (email: kennish@imcs.rutgers.edu).
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