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Porifera Research: Biodiversity, Innovation and Sustainability - 2007
131
Spongivory by juvenile angelfish (Pomacanthidae)
in Salvador, Bahia State, Brazil
Bárbara R. Andréa(1), Daniela Batista(1,2), Cláudio L.S. Sampaio(3), Guilherme Muricy(1*)
Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro. Quinta da Boa Vista, s/no.,
São Cristóvão. 20940-040 Rio de Janeiro, RJ, Brasil. [email protected]
(2)
Departamento de Biologia Marinha, Universidade Federal Fluminense, Niterói, Rio de Janeiro, Brasil
(3)
Departamento de Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, Brasil.
(1)
Abstract: Adult angelfish of the genera Pomacanthus and Holacanthus (Family Pomacanthidae) are considered the most
important spongivorous fishes of the Caribbean, with sponges comprising more than 70% of their gut contents. However,
despite their commercial importance as ornamental fish, little is known about the diet of juvenile angelfish, which are
generally considered to be cleaners. The goal of this study was to identify through gut content analysis the sponge species
eaten by juveniles of the angelfish Pomacanthus paru, Holacanthus ciliaris and Holacanthus tricolor in Salvador, Bahia state,
Brazil. We also estimated the frequency of occurrence of each sponge species in the diet of juvenile angelfish, and tested the
correlation between fish size and number of sponge species preyed. In Salvador, 34 species of sponges were found in the gut
contents of 14 out of 16 specimens of juvenile angelfish. Twenty-two species of sponges were eaten by Holacanthus tricolor,
15 by H. ciliaris, and 14 by Pomacanthus paru. There was a significant positive correlation between fish size and number
of preyed sponge species, but the coefficient of determination was low and even the smallest fishes had sponges in their gut
contents. These findings indicate that juveniles of all three species of angelfish are generalists in the consumption of sponges
in Brazil, a diet similar to that of the adults. The most frequent sponges in juvenile angelfish gut contents were Tedania ignis,
Mycale sp., and Spirastrella sp., all with 37.5% of frequency in the 16 stomachs analyzed. Tedania ignis was not consumed by
H. ciliaris in Salvador, but it was consumed by the other fish species, being the most frequent prey of H. tricolor and P. paru.
Juvenile angelfish probably adapt to a cleaning behavior or to benthic feeding according to local environmental conditions.
The development of sponge-based artificial foods may allow longer maintenance and reproduction of angelfish in aquaria,
thus helping to protect their natural populations.
Keywords: Holacanthus, Pomacanthus, Porifera, Predation, Southwestern Atlantic
Introduction
Sponges are important structural and functional components
of coral reefs, where they participate in numerous ecological
relationships such as competition, commensalism, symbiosis,
bioerosion and predation (reviewed by Diaz and Rützler 2001,
Wulff 2001, 2006a). Although reef sponges are abundant
and many species live exposed to predators, few animals
are known to feed on them, probably due to their physical
(spicules) and chemical (secondary metabolites) defenses
(Pawlik et al. 1995, Hill et al. 2005, Wulff 2006a, 2006b).
Among invertebrates, spongivory is shown by opisthobranchs
(e.g., Glossodoris pallida, Archidoris montereyensis,
Peltodoris atromaculata, Tylodina perversa), sea stars (e.g.,
Oreaster reticulatus, Echinaster echinophorus), sea urchins
(e.g., Eucidaris tribuloides and Lytechinus variegatus) and
hermit crabs (Reiswig 1973, Wulff 1995, 2006b, Becerro
and Paul 1998, Wadell and Pawlik 2000, Santos et al. 2002,
Gemballa and Schermutzki 2004). Some associated copepods,
amphipods, isopods and alpheid shrimps also consume their
host sponges (Pawlik 1983, Ríos and Duffy 1999, Mariani
and Uriz 2001). The main vertebrate sponge-feeders are the
sea turtle Erethmochelys imbricata (Meyland 1988) and reef
fishes belonging to the families Pomacanthidae, Ostraciidae,
Tetraodontidae, Ephippidae and Monacanthidae. Among
these, the family Pomacanthidae contains the most important
sponge-feeding fishes in the Caribbean, particularly in the
genera Pomacanthus and Holacanthus (Randall and Hartman
1968, Wulff 1994).
The family Pomacanthidae includes 88 species distributed
in all tropical seas (Allen et al. 1998, Debelius et al. 2003).
Within the family there is a diverse range of feeding
specializations, including herbivory, planktivory, cleaning
activity, and omnivory (Böhlke and Chaplin 1968, Hourigan
et al. 1989, Allen et al. 1998, Deloach 1999, Bellwood et
al. 2004). Ontogenetic variation in diet contents has been
observed in many pomacanthids (Thresher 1980, Deloach
1999) as well as in other reef fish families such as Blenniidae,
Kyphosidae and Scaridae (Bellwood 1988, Sturm and Horn
1998, Muñoz and Ojeda 2000). In the Caribbean, sponges
comprise over 70% of the diet of adults of the common
pomacanthid species Pomacanthus paru (French angelfish),
Pomacanthus arcuatus (Gray angelfish), Holacanthus ciliaris
(Queen angelfish) and Holacanthus tricolor (Rock beauty)
132
(Randall and Hartman 1968, Dunlap and Pawlik 1996). In
Brazil, adults of Pomacanthus paru also feed mostly on
sponges and algae, in variable proportions according to
the locality studied (Batista 2006). The feeding behavior
of juveniles, however, appears to be different from that of
adults in these species. Juveniles of H. ciliaris, P. paru and P.
arcuatus may act as cleaners until they have approximately
10-15 cm in total length (Feder 1966, Thresher 1980, Deloach
1999, Sazima et al. 1999). The primary food of P. paru and
P. arcuatus juveniles was reported to be filamentous algae,
with copepods picked from client fishes and few free-living
copepods making up to 25% of their diet (Deloach 1999).
In Abrolhos Archipelago, Brazil, juveniles of the French
angelfish P. paru removed ectoparasites of 31 species of reef
fishes (Sazima et al. 1999). Food items in their stomachs
included caligid (15-30%) and harpaticoid copepods (5-10%),
together with both red and green algae (30-70%). Juveniles of
H. ciliaris also feed on algae until they reach sexual maturity
(Deloach 1999). The diet of H. tricolor juveniles is largely
unknown, but it is assumed that it is a combination of drifting
plankton, small benthic invertebrates, and possibly the mucus
and parasites of larger fishes (Thresher 1979, 1980, Gasparini
and Floeter 2001).
Brazil is one of the five leading exporting countries of
tropical aquarium fishes in the world, and the interest in
marine ornamental organisms has increased substantially from
mid- to late 1990’s (Gasparini et al. 2005, Floeter et al. 2006).
Juvenile angelfish are preferred over the adults for ornamental
purposes due to their smaller size and beautiful color patterns.
In Ceará State, NE Brazil, angelfish juveniles have been
captured and exported in large numbers: 43,730 specimens
of H. ciliaris, 22,969 of P. paru, and 8,757 specimens of H.
tricolor between 1995 and 2000 (Monteiro-Neto et al. 2003),
although these values may be overestimated (Gasparini et al.
2005). Cleaner species play an important ecological role in
reef habitats, and their removal may negatively affect other
fish species, including commercially important ones (Sazima
et al. 1999, Monteiro-Neto et al. 2003). Therefore, knowledge
about the feeding behavior of juvenile angelfish is important
for the conservation of angelfish species and of coral reef
communities.
In this study we tested the hypothesis that sponges are an
important part of the diet of juvenile angelfish, and not only
of the adults. We identified the sponge species found in the
gut contents of juveniles of Pomacanthus paru, Holacanthus
ciliaris and H. tricolor in Salvador, Bahia state, Brazil. We
also estimated the frequency of each sponge species in the
gut contents of juvenile angelfish, and tested the correlation
between fish size and number of prey species. Our results
may aid in the development of a better diet for the growth
and reproduction of angelfish in captivity, thus helping to
protect angelfish species which are currently threatened by
the increase of marine ornamental fish trade in Brazil.
Materials and methods
Sixteen specimens belonging to three species of angelfish
were studied: Pomacanthus paru (n=5); Holacanthus ciliaris
(n=6) and Holacanthus tricolor (n=5). Fish were collected in
four locations in Salvador, Bahia state, Brazil (Fig. 1): Praia
Fig. 1: Location of the study area.
da Ribeira, in Todos os Santos Bay (12º54’S–38º29’W),
Barra Grande, in Itaparica Island (13º03’S–38º38’W), Rio
Vermelho (13º00’S–38º30’W), and Praia da Pituba (13º00’S–
38º30’W). The specimens were collected from 18/II/2003 to
10/III/2004 by commercial ornamental fisheries (Axé Online
Ltd.) in shallow reefs and rocky bottoms. Only juveniles
were caught, easily recognized by their special color patterns,
which change after sexual maturity is reached (Böhlke and
Chaplin 1968, Thresher 1980). Specimen size varied between
5.0-14.5 cm in H. ciliaris, 5.0-14.0 cm in H. tricolor, and
3.0-14.5 cm in P. paru. The stomachs were removed and
their contents were separated based on color, texture and
consistency. Only the sponge fragments were identified to
lower rank taxons, and no attempts were made to quantify the
abundance of each species consumed in the gut contents of
the 16 angelfish specimens analyzed. The sponge fragments
were dehydrated in an alcohol series (50-100%) with a final
xylene step and included in paraffin. Transverse sections were
mounted on microscope slides for identification. Free spicules
were not considered as evidence of predation, only fragments
large enough to be sectioned and in which the skeleton could
be observed. Sponges were deposited in the sponge collection
of Museu Nacional, Universidade Federal do Rio de Janeiro,
Brazil. A linear regression between fish size (standard length SL) and number of prey species was calculated online (http://
faculty.vassar.edu/lowry/VassarStats.htm).
133
Results
Sponges comprised more than 90% of the gut contents
of juvenile angelfish in Salvador, together with a few
unidentified filamentous algae (< 10%). The guts of two
juveniles of H. ciliaris were empty. A total of 34 sponge
species were found in the diet of angelfish in Salvador (Table
1). The greatest species richness was observed in gut contents
of Holacanthus tricolor (22 sponge species), followed by H.
ciliaris and Pomacanthus paru (15 and 14 sponge species,
respectively). Twelve sponge species were only predated by
H. tricolor: Acanthanchora sp., Coelosphaeridae unidentified,
Desmapsamma anchorata, Lissodendoryx sp., Microcionidae
unidentified 1, Microcionidae unidentified 2, Monanchora
sp., Myxillina unidentified, Pachastrellidae unidentified,
Phellodermidae unidentified, Plakinastrella sp., and Timea sp.
Only three species were predated exclusively by Holacanthus
ciliaris: Artemisina sp., Chalinidae unidentified, and Tethya
sp. Six species were eaten exclusively by P. paru: Chondrilla
nucula complex, Cyamon sp., Mycale laxissima, Niphatidae
unidentified, Raspaciona sp., and Tetillidae unidentified (Table
1). As a whole, the sponge species most frequently predated
by pomacanthids in Salvador were Tedania ignis, Mycale sp.,
and Spirastrella sp., all present in 37.5% of the 16 specimens
analyzed. Although Tedania ignis was not consumed by H.
ciliaris, it was very frequent in the gut contents of P. paru
(80% of the specimens examined) and common in H. tricolor
(40%; Table 1). The number of sponge species per stomach
varied from 4-9 in H. tricolor, from 1-7 in P. paru, and from
0-8 in H. ciliaris.
The curves of accumulated richness of sponge species in
angelfish diet did not reach stabilization with the small sample
sizes studied here, neither with each fish species considered
separately nor when they were considered together (Fig. 2).
This indicates that the real number of sponge species in the
diet of juvenile angelfish in Salvador is probably much higher
than that shown by our results.
There was a significant positive correlation (p<0.05)
between fish size and number of sponge species in gut
contents, although with a low coefficient of determination
(r2 = 0.465; Fig. 3). The single smallest fish (3 cm SL) had
only one sponge species in its stomach, and three of the five
smallest fishes (3.0-5.5 cm SL) fed on few sponge species (15 species per stomach).
Table 1: Frequency of occurrence of sponge species in gut contents of juvenile angelfish in Salvador, Brazil (in %). Combined = frequency
in all species pooled together (n=16).
Sponge species
Acanthanchora sp.
Acervochalina sp.
Artemisina sp.
Chalinidae unidentified
Chondrilla nucula complex
Clathria sp. 1
Clathria sp. 2
Coelosphaeridae unidentified
Cyamon sp.
Desmapsamma anchorata
Halichondriidae unidentified
Haplosclerida unidentified
Lissodendoryx sp.
Microcionidae unidentified 1
Microcionidae unidentified 2
Mycale laxissima
Mycale sp.
Mycalidae unidentified
Myxillina unidentified
Monanchora sp.
Niphatidae unidentified
Pachastrellidae unidentified
Phellodermidae unidentified
Plakinastrella sp.
Raspaciona sp.
Spirastrella sp.
Stelleta sp. 1
Stelleta sp. 2
Strongylacidon sp.
Suberitidae unidentified
Tedania ignis
Tethya sp.
Tetillidae unidentified
Timea sp.
Species richness
P. paru
20
20
40
20
20
40
20
40
20
20
60
20
80
20
14
H. tricolor
20
40
20
20
40
20
20
20
20
20
60
40
40
H. ciliaris
17
17
17
33
17
50
17
50
17
20
20
20
20
20
20
20
40
33
17
17
17
17
17
20
22
15
Combined
6.25
12.50
6.25
6.25
6.25
25.00
25.00
6.25
6.25
12.50
12.50
31.25
6.25
12.50
6.25
6.25
37.50
25.00
12.50
12.50
6.25
6.25
6.25
6.25
6.25
37.50
12.50
12.50
12.50
12.50
37.50
6.25
6.25
6.25
34
134
Fig. 2: Curves of accumulated richness of sponge prey species in juvenile angelfish gut contents. A, Holacanthus tricolor; B, Holacanthus
ciliaris; C, Pomacanthus paru; D, all species pooled together.
Discussion
Juvenile angelfish fed largely on sponges in Salvador (>
90%), and only a few filamentous algae were also found in
their gut contents. These figures are similar to those of adult
angelfish diet in the Caribbean, where sponges comprised
74.8% to 97.1% of the diet of Pomacanthus paru and
Holacanthus ciliaris, respectively (Randall and Hartman
1968). Algae were also the second most common item in
the diet of adult pomacanthids in the Caribbean, varying
from 0.8% to 13.4% of the diet of H. tricolor and P. paru,
respectively (Randall and Hartman 1968). Other food items
common in Caribbean adults such as tunicates, hydroids,
zoantharians, and bryozoans were not found in juveniles from
Salvador.
Fourteen out of 16 fishes (87.5%) collected in one year had
sponge remains in their stomachs, indicating that spongivory
is not occasional, but frequent and widespread in the juvenile
angelfish population in Salvador. Adult angelfish appear to
have a more varied diet, at least in the Caribbean, where
sponges were found in only 12 of 23 stomachs of P. paru
(52.1%) and 18 out of 34 specimens of P. arcuatus (52.2%).
Adults of H. tricolor are more specialized in sponges, with
sponge remains in 22 of 24 specimens (91.7%; Randall and
Hartman 1968).
The diet of juvenile angelfish in Salvador includes at least
34 sponge species. All three angelfish species studied here,
Pomacanthus paru, Holacanthus ciliaris, and H. tricolor
were generalists in the consumption of sponges, eating 14 to
22 different species each. The real number of sponge species
consumed by these angelfish is probably higher than that, as
indicated by the shape of the accumulated richness curves
(Fig. 2). Although there was a significant positive correlation
between fish size and number of preyed sponge species
(Fig. 3), the coefficient of determination was low and even
the smallest fishes had sponges in their gut contents. Part of
this correlation may reflect greater gut volume in larger fish
individuals, allowing them to have more bites (and therefore
more species) in their guts at the time they were collected.
The diet of pomacanthid juveniles in Salvador is very
similar to that of the adults, both in the Caribbean and in
Brazil: they are “smorgasbord-feeding sponge specialists”,
consuming a variety of benthic invertebrates and algae,
apparently chosen according partly to prey palatability and
partly to prey availability in the environment (Randall and
Hartman 1968, Hourigan et al. 1989, Wulff 2006a, Batista
2006). This is at odds with some studies which suggest that
juveniles of H. ciliaris and P. paru are cleaners until they have
approximately 10-15 cm in total length, when they change to
benthic feeding of a mainly sponge and algae diet (Feder 1966,
Böhlke and Chaplin 1968, Hourigan et al. 1989, Sazima et al.
1999). Other studies however found little cleaning activity in
juveniles of P. paru (Wicksten 1995, 1998). Among Tropical
Western Atlantic angelfish, juveniles of P. paru seem to be the
most specialized as cleaners, with P. arcuatus and H. ciliaris
having a mixed diet composed mostly of algae and detritus
135
Fig. 3: Linear regression between fish size (standard length in
cm) and the number of sponge species in gut contents of juvenile
angelfish. Slope = 1.1173, intercept = 5.0347, Standard error of the
estimate = 3.169.
and being only occasional cleaners (Thresher 1980, Deloach
1999). The diet of H. tricolor juveniles was hitherto poorly
known, although cleaning activity has been reported in a few
other studies (Thresher 1979, 1980, Gasparini and Floeter
2001).
The method of study employed by each author is important
to explain such disagreements. When only direct underwater
observation is considered (e.g., counting the bites given in each
prey species), there is a trend to increase the relative importance
of algae and of cleaning activity in the diet of pomacanthids
(Böhlke and Chaplin 1968, Thresher 1980, Hourigan et
al. 1989, Sazima et al. 1999, Batista 2006). Juveniles of
angelfish such as H. tricolor dwell mostly in cryptic habitats
inaccessible for divers (Thresher 1980). These habitats are
often dominated by sponges (Sarà and Vacelet 1973). Other
species of angelfish, including adult specimens, also use
small caves and reef crevices for sheltering and foraging,
biasing the direct observation of preyed items towards more
exposed organisms such as algae and gorgonians (Thresher
1980, Batista 2006). When only gut contents are analyzed, the
relative importance of spongivory seems to increase, both in
proportion to algae and benthic invertebrates and in number
of species (e.g., Randall and Hartman 1968, Deloach 1999). It
is possible that sponge fragments remain recognizable in the
guts of angelfish longer than copepods and filamentous algae,
due to their high collagen and spicule content (Chanas and
Pawlik 1995). This would artificially increase the proportion
of spongivory in studies based only in gut contents and
without direct observations, such as the present one (see also
Randall and Hartman 1968). The complementary use of both
field observations and gut content analysis allows a better
understanding of pomacanthid feeding habits (Hourigan et al.
1989, Sazima et al. 1999, Batista 2006, Wulff 1994, 2006a).
It is possible therefore that juvenile angelfish in Salvador are
not almost exclusively sponge-eaters as shown by our results,
but might also consume other benthic organisms and act as
cleaners occasionally. This issue can only be solved through
systematic direct observations on juvenile angelfish, which
could not be carried out in this study.
Another explanation for the variation in the diet of
pomacanthid juveniles in different studies is a high dietary
plasticity of pomacanthid species. Allopatric angelfish
populations may specialize to use different resources,
depending on their availability in each locality. For example,
the abundance of reef fish in general is greater in the Abrolhos
National Marine Park, which is a protected area, than in
Salvador, which has been subject to heavy fishing (Gasparini
et al. 2005, Floeter et al. 2006). It may thus be easier for
juveniles of pomacanthids such as P. paru to establish a
cleaning station in Abrolhos than in Salvador, where they
have to adapt their diet to the available resources, mainly
sponges and algae. The presence of a suitable habitat for
juveniles close to a reef fish community also helps to explain
the intense cleaning activity of P. paru juveniles in Abrolhos
reefs (Sazima et al. 1999).
Although always composed mostly of sponges, the specific
composition of the diet of adult angelfish varied strongly
in different localities such as the Bahamas, Virgin Islands,
and Panama in the Caribbean (Randall and Hartman 1968,
Feddern 1968, Hourigan et al. 1989, Wulff 1994), and Atol
das Rocas, Abrolhos and Ilha Grande in Brazil (Batista 2006),
irrespective of the study method. In all these locations and
in Salvador, each species of angelfish fed on a diverse array
of sponge species, ranging from 12 in P. paru to 40 in H.
ciliaris, both in the U.S. Virgin Islands (Randall and Hartman
1968, Hourigan et al. 1989). Together, Pomacanthus paru
and P. arcuatus ate 64 sponge species in Panama (Wulff
1994). The only exception was H. tricolor, which was
observed feeding on a single sponge species in Panama and
was therefore considered a possible specialist (Wulff 1994).
However, adults of H. tricolor are known to fed on 14 to 28
different sponge species in the U.S. Virgin Islands and Puerto
Rico (Randall and Hartman 1968, Hourigan et al. 1989), and
juveniles in Salvador fed on 22 sponge species. This indicates
that H. tricolor is also a generalist in the consumption of
sponges, like most other pomacanthids. The number of sponge
species eaten by each individual fish in Salvador varied from
0-9 (average 4.5). This tolerance to predate upon diverse
sponge species may be a strategy of angelfish to overcome
the effects of sponge toxins by eating small amounts of each
toxic substance from many different sponge species (Wulff
1994). It has also be suggested as an adaptation of benthic
fish species with pelagic larvae such as angelfish to increase
their chances of survival in areas with different availability
of specific food items (Hourigan et al. 1989). The similarity
between gut contents of different but co-specific specimens
was low in Salvador (maximum 0.273 between two specimens
of H. tricolor; data not shown), indicating that individual fish
choose their food independently. This may be related to the
very small home range of angelfish juveniles (around 1-2 m2),
much smaller than that of the adults (up to 2,300 m2 in P.
paru; Thresher 1980, Hourigan et al. 1989).
All specimens studied here came from the same general
region in Salvador, but dietary similarity between the three
species was low (maximum 0.296 between H. ciliaris and H.
136
tricolor; data not shown) and in the same range of intraspecific
similarity. Most of the interspecific variation found in
angelfish diet in Salvador thus appears to be random, and due
to the sum of independent choices of individual fishes of the
three different species studied. As most fishes, spongivores
have great plasticity in their feeding behavior. This variability
could have influence from food availability, fish size, species
and other temporal and spatial factors. However, part of
the differences in dietary composition between sympatric
species is probably due to an active choice of preys, possibly
restrained by the physiological tolerance of each fish species to
the prey’s toxins. A certain degree of choice in pomacanthids
is supported by observations of predation upon relatively
rare sponge species (Hourigan et al. 1989, Wulff 1994).
Holacanthus passer often feeds on plankton and fish faeces
in the water column in the eastern Pacific (Aburto-Oropeza et
al. 2000); there are few exposed sponges on coral reefs there,
but when cryptic sponges were exposed the angelfish quickly
swam down out of the water column to feed on them (Wulff
1997). Caribbean angelfish also prefer to eat mangrove or
cryptic sponge species whenever they are made available in
the reefs (Dunlap and Pawlik 1996, Wulff 2005). In general,
spongivorous fish tend to choose the most palatable or
undefended prey species available in a given locality (Dunlap
and Pawlik 1996), although which prey is more palatable
varies according to the predator species (Wulff 2006a). For
instance, the fire-sponge Tedania ignis is commonly eaten by
H. tricolor and P. paru, but apparently not by H. ciliaris, both
in Brazil and in the Caribbean (Table 1; see also Randall and
Hartman 1968). The starfish Oreaster reticulatus also feeds
on Tedania ignis in Belize, but not on the sibling species T.
klausi (Wulff 2006b). Also, the dietary similarity between
congeneric species of both Pomacanthus and Holacanthus is
greater than between species from different genera (Table 1;
see also Hourigan et al. 1989). Whether this truly represents
a species-specific choice of prey by pomacanthids remains to
be experimentally demonstrated.
Both adult and juvenile angelfish present high plasticity,
tolerance, and a certain degree of choice in their smorgasbord
sponge-feeding specialized strategy. This appears to be
a highly derived feeding strategy, which is coupled with
morpho-functional modifications in the teeth, jaws, and gill
rakers that make pomacanthids (especially Holacanthus and
Pomacanthus) more apt to feed on structurally resilient and
firmly attached benthic prey such as sponges, gorgonians and
tunicates (Hourigan et al. 1989, Bellwood et al. 2004; Konow
and Bellwood 2005). Sponges have been evolving during at
least 580 MY (Li et al. 1998), and they developed chemical
and physical defenses so effective that only very specialized
predators such as angelfish and opisthobranchs are able to
feed on them. This may help to explain why there are so few
predators of sponges, despite their great abundance in coral
reefs.
Angelfish are threatened by marine ornamental fish trade
in Brazil, and juveniles are the targets preferred by aquarists
(Gasparini et al. 2005, Floeter et al. 2006). The maintenance
of angelfish in aquaria is difficult due to their aggressiveness
and their diet made up mostly of sponges (Thresher 1980). The
development of artificial food based on the sponge species
found in this study to be eaten by angelfish juveniles in the
field might help their reproduction and survival in captivity,
allowing aquarists to enjoy angelfish in their saltwater tanks
without eliminating them from coral reefs, where they play
important ecological roles.
Acknowledgements
We thank José de Anchieta Nunes, Camilo Ferreira and Ericka Coni
for laboratory assistance. We are also grateful to Samuele Clerici
of Axé Online Fishes for the kind donation of specimens for study.
We thank the two anonymous reviewers for their comments, which
greatly improved the manuscript. This study was supported by grants
and fellowships from Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) and Fundação Carlos Chagas Filho
de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ).
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