Mar Biol (2016) 163:6 DOI 10.1007/s00227-015-2788-4 ORIGINAL PAPER Abundance, diet, foraging and nutritional condition of the banded butterflyfish (Chaetodon striatus) along the western Atlantic Ana M. R. Liedke1,2 · Diego R. Barneche3 · Carlos E. L. Ferreira4,5 · Barbara Segal2,5 · Lucas T. Nunes2 · Ana P. Burigo2 · José A. Carvalho6 · Sonia Buck2,7 · Roberta M. Bonaldo8 · Sergio R. Floeter2 Received: 5 April 2015 / Accepted: 13 November 2015 © Springer-Verlag Berlin Heidelberg 2016 Abstract The feeding behaviour and diet plasticity of the most important items in seven study sites. Therefore, C. a given species are usually shaped by the relationship striatus may be considered as a non-coral generalist feeder, between species physiology and the quality and availabil- as it feeds on a wide variety of items and substrata along ity of resources in the environment. As such, some species the studied range, with no consistent selectivity pattern for may achieve wide geographical distributions by utilizing foraging substratum across sites. Individuals from all sites multiple resources at different sites within their ranges. We but Salvador (NE Brazil) had similar RNA/DNA ratios, studied the distribution and feeding of Chaetodon striatus, suggesting that C. striatus nutritional condition is similar the most widespread butterflyfish in the Atlantic, by assess- along its extensive distribution. Our findings highlight the ing its density and foraging rates in eight sites enclosing importance of assessing different sites within the distribu- 44° of latitude. We also evaluated the relationship between tion range of generalist butterflyfishes, and different vari- fish density and foraging rates with nutritional condition ables, to a better comprehension of the feeding ecology of and diet across study sites, and the gut length relative to these species. body size. Density and foraging rates did not differ among studied sites. In 169 stomachs analysed, we found 52 dif- ferent items (12–23 per site). Polychaeta and Cnidaria were Introduction Diet and feeding behaviour are primarily important to the ecology, biology and evolution of organisms, as they are Responsible Editor: K.D. Clements. essential traits that affect species distribution and popu- Reviewed by R. Francini-Filho, M.S. Pratchett lation structure (Brown 1984; Cole and Pratchett 2014). In this sense, the feeding behaviour and diet plasticity of Electronic supplementary material The online version of this a given species are usually shaped by the relationship article (doi:10.1007/s00227-015-2788-4) contains supplementary material, which is available to authorized users. * Sergio R. Floeter 5 Instituto Coral Vivo, Porto Seguro, BA 45816‑000, Brazil sergio.floeter@ufsc.br 6 Faculdade de Ciências da Terra, do Mar e do Ambiente, 1 Programa de Pós Graduação em Ecologia e Conservação, Universidade do Algarve, Campus de Gambelas, 8000 Faro, Universidade Federal do Paraná, Curitiba, PR 81531‑990, Portugal Brazil 7 Departamento de Ciências Ambientais, Universidade Federal 2 Departamento de Ecologia e Zoologia, Universidade Federal de São Carlos, Rod. Washington Luis, Km 235, São Carlos, de Santa Catarina, Florianópolis, SC 88010‑970, Brazil SP 13565‑905, Brazil 8 3 Centre for Geometric Biology, School of Biological Departamento de Ecologia, Instituto de Biociências, Sciences, Monash University, Clayton, VIC 3800, Australia Universidade de São Paulo, São Paulo, SP 05508‑900, Brazil 4 Departamento de Biologia Marinha, Universidade Federal Fluminense, Niterói, RJ 24001‑970, Brazil 1 3 6 Page 2 of 13 Mar Biol (2016) 163:6 between species morphology, physiology and the quality with only a small fraction of its diet, if any, represented by and availability of resources in the environment (Brown scleractinian corals (e.g. Randall 1967; Pitts 1991; Sazima 1984; Lawton et al. 2012; Cole and Pratchett 2014). This and Sazima 2001; Bonaldo et al. 2005). Chaetodon striatus relationship ultimately influences species distribution and is the butterflyfish with the widest distribution range in the density in a given location (Gerking 1994; Lawton et al. western Atlantic, which encompasses two biogeographic 2012). As such, some species may achieve high local abun- provinces (Caribbean and Brazil) and a diversity of envi- dance and wide geographical distributions by utilizing ronmental conditions, such as temperature (17–30 °C), hab- multiple resources at different sites throughout their ranges itat type (e.g. coral and rocky reefs), and food items/avail- (Lawton et al. 2012). ability (Floeter et al. 2001). This diversity of environmental The butterflyfishes (Chaetodontidae) include about 130 and biotic conditions may ultimately affect the abundance reef fish species that live closely associated to the substra- (e.g. via recruitment rates; Pratchett and Berumen 2008), tum, and most of them consume a variety of benthic prey diet, foraging selectivity (e.g. by changes in resource avail- items, such as anthozoans, polychaetes and crustaceans ability) and nutritional condition of C. striatus along its (Randall 1967; Birkeland and Neudecker 1981; Sazima and distribution range. However, given the apparent high feed- Sazima 2001; Pratchett 2005). Butterflyfishes are gener- ing versatility and generalist habits of this species, it is ally classified into three major guilds according to their diet possible that differences in prey availability do not exert a and feeding selectivity: obligate corallivores, facultative strong influence on local abundance of C. striatus along its corallivores and non-coral generalist feeders. Most studies distribution, especially in comparison with highly special- on butterflyfish feeding have been focused on Indo-Pacific ized corallivorous butterflyfishes. corallivores, which usually have wide longitudinal and, in We compared the abundance, diet and foraging rates and some cases, latitudinal distributions on tropical coral reefs selectivity of C. striatus among eight distinct sites, span- (e.g. Hobson 1974; Irons 1989; Tricas 1989; Pratchett ning 44° of latitude, testing whether local variation in the 2005; Gregson et al. 2008). Generalist corallivorous but- availability of prey exerted an important influence on this terflyfishes with wide distributions in the Indo-Pacific may seemingly generalist species. We particularly investigated change their diet and foraging preferences over large spa- whether fish density, foraging rates, foraging selectivity tial scales, in response to food availability (Berumen et al. and nutritional condition varied along the studied gradient. 2005; Cole and Pratchett 2014) and intraspecific/interspe- We verified key dietary resources of C. striatus by validat- cific competition (Cox 1994; Pratchett 2005). On the other ing in situ feeding observations using dietary analyses. We hand, this variation is less noticeable in highly specialized also assessed C. striatus gut length relative to body size, to corallivorous butterflyfishes in the Indo-Pacific, which are verify whether this metric supported the food habit found more sensitive to differences in the availability of their pre- for this species. As we show below, we found no differ- ferred prey species (Devictor et al. 2010; Cole and Pratch- ences in C. striatus density and foraging rates among sites. ett 2014). Also, our results suggest that C. striatus has high feeding In contrast to corallivorous butterflyfishes, very few plasticity, as it forages on a wide variety of food items studies assessed feeding habits and dietary composition of and substrata along the studied sites, without clear differ- non-coral feeder species (e.g. Randall 1967; Birkeland and ences in nutritional condition. Therefore, the present study Neudecker 1981; Bonaldo et al. 2005). Non-coral feeders reveals how wide-ranging species can be better understood differ from other butterflyfishes in their geographical dis- using a plethora of complementary approaches. tribution (Lawton and Pratchett 2012; Cole and Pratchett 2014), as their range is not only associated with coral reef habitats and may include temperate environments (Lawton Materials and methods and Pratchett 2012; Cole and Pratchett 2014). Non-coral feeder butterflyfishes also differ from other butterflyfishes Study sites in their morphology, as their gut length/body size ratio is usually intermediate to that of corallivores and planktivores The present study was conducted in eight sites distrib- (Berumen et al. 2011). However, little is known on the uted along the western Atlantic. These sites included three feeding ecology of non-coral feeder butterflyfishes and how types of reefs: tropical biogenic reefs (one in Puerto Rico their abundance and feeding habits respond to environmen- and three in Brazil: Tamandaré, Salvador and Abrolhos), tal differences within their distribution. volcanic rocky reefs (Trindade Island) and subtropical In the western Atlantic, the banded butterflyfish, Chae- rocky reefs (Guarapari, Arraial do Cabo and Florianópolis; todon striatus, is classified as a non-coral generalist feeder all situated in Brazil) (Fig. 1; Table 1). These eight sites (also known as benthic invertebrate feeder sensu Cole and were chosen because of their different reef formations and Pratchett 2014), as it feeds on a wide variety of prey items, because they encompass a wide latitudinal range along 1 3 Mar Biol (2016) 163:6 Page 3 of 13 6 −96 −62.5 −29 80 Puerto Rico 80 Tamandaré 60 60 30 40 40 20 20 0 0 15 80 Salvador 80 Abrolhos 60 60 40 40 ) 20 20 0 0 0 80 Trindade
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