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Green2009-herbivore monitoring

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Herbivorous Coral Reef Fishes<br />

HERBIVOROUS CORAL REEF FISHES<br />

KEY FAMILIES AND THEIR FEEDING MODES<br />

Herbivorous reef fishes are taxonomically diverse, comprising at least nine families (Choat 1991). The<br />

most important families are the Acanthuridae (surgeonfishes and unicornfishes), Pomacentridae<br />

(damselfishes), Labridae (Tribe Scarini 1 : parrotfishes), Siganidae (rabbitfishes) and Kyphosidae<br />

(rudderfishes: Choat 1991). Some of these families are quite speciose, particularly surgeonfishes and<br />

parrotfishes (Choat 1991).<br />

Another potentially important family is the Blenniidae (blennies), which is usually overlooked because<br />

of their cryptic habits. Other families comprising some herbivorous representatives, include the<br />

Pomacanthidae (angelfishes), Gobiidae (gobies) and Ephippidae (batfishes: Choat 1991, Bellwood et<br />

al 2006). Some Monacanthidae (filefishes), Balistidae (triggerfishes) and other Labridae (wrasses)<br />

may also be <strong>herbivore</strong>s, but their trophic status has not been confirmed (Myers 1999, Randall et al<br />

1996, G.P Jones unpubl. data).<br />

Herbivorous reef fishes exhibit a range of feeding modes and ingest a variety of plant material,<br />

including macroalgae, epilithic algal turf, detrital material, algal mats and associated organisms<br />

(including bacteria: reviewed by Choat 1991). There is considerable variation among and within<br />

families, and they do not all perform the same role nor do they have similar impacts on coral reef<br />

ecosystems (Choat 1991).<br />

There are two groups of parrotfishes, which show major differences in their jaw morphology and<br />

feeding behaviour: excavators and scrapers (Bellwood and Choat 1990). The majority of parrotfishes<br />

(scarines 1 ) are scrapers (most Scarus and Hipposcarus species) that feed on epilithic algal turf<br />

growing on hard substrata (Bellwood and Choat 1990), although some Indo-Pacific species graze<br />

mainly over sand (Choat 1991). They take non-excavating bites and scrape material from the surface<br />

of the substratum. Excavators differ from scrapers by taking deeper, excavating bites and removing<br />

more of the substratum as they feed (Bellwood and Choat 1990). They include Bolbometapon<br />

muricatum, Cetoscarus bicolor and species of the genus Chlorurus. Members of the genus<br />

Calotomus are browsers, while some members of the Caribbean subfamily Sparisomatinae feed on<br />

marine angiosperms (Choat 1991). B. muricatum also grazes substantial amounts of live coral (Choat<br />

1991, Bellwood et al 2003). Four other Indo-Pacific parrotfish species have been observed grazing on<br />

live corals (Chlorurus microrhinos, Cetoscarus bicolor, S. frenatus and S. rivulatus: Bellwood and<br />

Choat 1990, Bellwood 1995), although corals only account for a small proportion of their diet (Hoey<br />

and Bellwood 2008).<br />

There is a greater diversity of feeding habits within the Acanthuridae (Choat 1991). Most species of<br />

the genera Acanthurus and Zebrasoma graze on epilithic algal turf over reef substrata (Choat 1991,<br />

Choat et al 2002), although some Acanthurus species graze over sand and feed on a combination of<br />

algae and detritus (ringtail surgeonfishes including A. blochii, A. dussumieri, A. leucocheilus, A.<br />

maculiceps, A. nigricauda, A. olivaceus, A. pyroferus, A. tristis and A. xanthopterus) while others are<br />

planktivores (A. albipectoralis, A. mata, A. nubilus and A. thompsoni: Choat 1991, Choat et al 2002).<br />

All members of the genus Ctenochaetus are primarily sediment and detritus feeders (Choat 1991),<br />

while the single species of the monotypic genus Paracanthurus is a planktivore (Choat 1991, Choat et<br />

al 2002). The majority of unicornfishes are also planktivores (Choat 1991, Choat and Clements 1998,<br />

Choat et al 2002), with the largest individuals (N. annulatus, N. hexacanthus and Naso lopezi) feeding<br />

on macroplankton in open<br />

water (Choat 1991). A second group of unicornfishes, including N. unicornis and N. tuberosus, feed<br />

on large macroscopic algal species (Choat and Clements 1998).<br />

1 Recent phylogenetic analyses have confirmed that parrotfishes, previously Family Scaridae, belong to the Tribe<br />

Scarini within the Family Labridae (Westneat and Alfaro 2005, Cowman et al in press). Therefore parrotfishes,<br />

previously known as scarids, are now known as scarines.<br />

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