Local Variations in Microhabitat Use by Stegastes Fuscus (Cuvier, 1830
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Brazilian Journal of Biological Sciences, 2016, v. 3, no. 6, p. 375-384. ISSN 2358-2731 http://dx.doi.org/10.21472/bjbs.030613 Local variations in microhabitat use by Stegastes fuscus (Cuvier, 1830) (Teleostei: Pomacentridae) in a tropical reef of Brazil Paulo Roberto de Medeiros¹, Ana Luisa Pires Moreira² and Ana Maria Alves de Medeiros³ ¹Universidade Federal de Campina Grande. Centro de Formação de Professores. Unidade Acadêmica de Ciências Exatas e da Natureza. Rua Sérgio Moreira de Figueiredo, S/Nº. Cajazeiras-PB. Brazil (CEP 58900-000). Email: [email protected]. ²Instituto de Desenvolvimento Sustentável e Meio Ambiente do Rio Grande do Norte (IDEMA). Núcleo de Unidades de Conservação (NUC/IDEMA). Área de Proteção Ambiental dos Recifes de Corais (APARC). Av. Alm. Alexandrino de Alencar, S/Nº. Tirol. Natal/RN (CEP: 59015-350). ³Laboratório de Ecologia. Departamento de Sistemática e Ecologia. Centro de Ciências Exatas e da Natureza. Universidade Federal da Paraíba. João Pessoa-PB. Brazil. (CEP 58050-900). Abstract. Due to their widespread abundance and territorial habits, damselfishes play central roles in reefs worldwide, but the extent to Received which they modify the reef’s substrate varies both amongst and November 17, 2016 within species. The present study evaluated microhabitat preferences and the role of the common damselfish Stegastes fuscus (Cuvier, Accepted December 22, 2016 1830) (Teleostei: Pomacentridae) as a benthic community modifier in a tropical algal-dominated reef. We employed video survey Released techniques to assess fish density and microhabitat use and, December 31, 2016 additionally, the biomass of sessile components were compared inside and immediately outside algal farms of S. fuscus. Individuals Open Acess Full Text Article showed pronounced microhabitat preference, but with prominent local (i.e. small-scale) variations in benthic cover. These local variations are reported for the first time and were due to individuals defending one of two contrasting microhabitats (i.e. turf-dominated or Palythoa caribaeorum-dominated). Further, significant differences in the biomass of sessile organisms were observed within algal farms suggesting that S. fuscus plays a keystone role in the benthic community of the reef. The implications of the prominent local variation in microhabitat use are discussed and strongly illustrate the behavioral plasticity of this damselfish. ORCID Keywords: Damselfish; Habitat Complexity; Benthic Biomass; 0000-0002-8534-7331 Southwestern Atlantic. Paulo Roberto de Medeiros 0000-0001-7748-6131 Ana Luisa Pires Moreira 0000-0002-8119-0197 Ana Maria Alves de Medeiros ISSN 2358-2731/BJBS-2016-0070/3/6/13/375 Braz. J. Biol. Sci. http://revista.rebibio.net 376 Medeiros et al. Introduction an intraspecific component, with fish of different size (Foster, 1985) and from Territorial pomacentrids such as the different stages (Medeiros et al., 2010a) damselfishes play central roles as benthic defending territories with different algal modifiers in reefs worldwide (Hixon, 1996; composition. Ceccarelli, 2007). These ecologically Hata and Kato (2002; 2004) significant tasks are largely the result of referred to damselfishes that strongly their ability to specific remove undesired defend small, monospecific farms and and surplus components, therefore selectively remove unpalatable algae as maintaining selected species (Ceccarelli et intensive farmers. On the other hand, al., 2001; Ceccarelli et al., 2005) and, often, territories of less aggressive damselfishes monoculture algal farms (Hata and Kato, tend to be larger and include indigestible 2002; Hata and Kato 2003). Additionally, algae, characterizing an extensive farmer. effects on the benthic community come Intensive farmers gain an increased food about from the territorial exclusion of supply per unit area at the expense of a herbivores (e.g. fishes and sea urchins) via higher energy prompted during territory aggressive behaviors (Kamura and defense and weeding, whereas extensive Choonhabandit, 1986). Therefore, farmers dispend less energy in territory damselfishes’ territories are often regarded maintenance, but territories are larger due as zones of high productivity (Klumpp et to the low crop yield per unit area. al., 1987). Hixon and Brostoff (1983) were The overall aim of the present study amongst the first to demonstrate how was to determine microhabitat preference damselfishes maintain benthic diversity via by the dusky damselfish in a tropical intermediate-disturbance, but several other shallow reef, and to compare benthic studies also investigated the role of composition inside and outside their damselfishes as keystone species in reefs territories as a means of understanding the worldwide (e.g. Brawley and Adey, 1977; role played by these fishes as benthic Branch et al., 1992; Ceccarelli et al., 2001; community modifiers. Barneche et al., 2009). Nonetheless, although territoriality Materials and Methods seems to be a behavioral synapomorphy shared by all damselfishes and regarded as Study area somewhat conservative and stereotyped The study was conducted at Seixas across species, the extent to which reefs, João Pessoa, PB, Northeastern, Brazil damselfishes are able to defend territories is (Figure 1). It is a shallow (< 6 m) sandstone highly variable (Hata and Kato, 2002). For reef made up by scattered consolidated example, the two most common substrata surrounded by a contiguous damselfishes in tropical reefs of Brazil limestone bottom. Common algae include (Stegastes fuscus and S. variabilis) show Gelidium spp., Caulerpa racemosa, distinct territorial defense patterns, the Halimeda opuntia, Dictyopteris delicatula former being more aggressive towards and Lithothamnium spp. and algal diversity intruders than the latter (Medeiros et al., is high throughout the reef. Common 2010a). The extent to which algae are sessile animals in the area include the deliberately removed from the territories hydrocoral Millepora alcicornis, the (i.e. weeding) also differs among species zoanthids Palythoa caribaeorum and (Ferreira et al., 1998; Ceccarelli et al., Protopalythoa variabilis, and the corals 2001; Hata and Kato, 2002). Therefore, Agarices agaricites and Siderastrea variations in algal composition are mostly stellata. These zoanthids and corals show a due to differences in algal biomass and somewhat patchy distribution, but with richness within territories of different peak abundances in particular species (Ceccarelli et al., 2001; Gobler et microhabitats. al., 2006). In fact, variations may also show Braz. J. Biol. Sci., 2016, v. 3, no. 6, p. 375-384. Local variations in microhabitat use by Stegastes fuscus 377 Figure 1. Location and map of the study area. Microhabitat preferences of still frames where one or more S. fuscus Density of juvenile and adult individuals appeared in the center frame Stegastes fuscus, and benthic composition (total of 33 frames) with 50, randomly were evaluated by means of video surveys chosen frames (with or without S. fuscus). on ten randomly assigned 10 m x 2 m This was done to test the degree of transects (length x width). Data were randomness in microhabitat use and to recorded during daytime by the same diver specific determine composition of algal in January 2013 at depths below 2 m and farms in relation to the overall benthic during low tides. Video was recorded using composition. Therefore, benthic estimates high definition configuration using a Canon were treated in a twofold manner: observed digital camera equipped with appropriate (benthic cover estimated in still frames water housing. In each transect, one diver exclusively associated to S. fuscus; i.e. the swam at constant speed, resulting in actual microhabitat use) versus randomly transects lasting (mean ± SD) 2 min 19 sec expected (benthic cover estimated in all ± 9 sec (average diver speed: 4.32 m.min-¹). microhabitats, associated or not to S. To determine the density of fuscus). Twenty points were randomly S. fuscus, each video was cautiously plotted on each still frame and assigned to examined three times and individuals one of ten categories, as follows: fleshy (settled juveniles and adults) were macroalgae (e.g. Dictyopterys, Caulerpa quantified. Repeated counts were conducted and Sargassum), turf algae (e.g. Gelidium, as a means of minimizing observer error Griffithsia), Lithothamnium, Halimeda, during video quantification. Nonetheless, Protopalythoa variabilis, Palythoa divergences among counts within each caribaeorum, Zoanthus sociatus, video transect were not observed Siderastrea stellata, uncolonized pavement whatsoever. and limestone. Percent cover, therefore, is Given that the video transects the relative proportion of a category relative included all microhabitats of the reef (some to the overall composition. Number of of which were not occupied by S. fuscus), crevices was also estimated on each still we evaluated the degree of microhabitat frame. These procedures were conducted on fidelity by comparing benthic composition Braz. J. Biol. Sci., 2016, v. 3, no. 6, p. 375-384. 378 Medeiros et al. Coral Point Count Software (Kohler and and single benthic categories, benthic Gill, 2006). diversity, and number of crevices. Benthic composition and biomass To compare biomass values from were also evaluated inside and outside the inside and outside the territories, a territories of 10 S. fuscus individuals. First, MANOVA procedure, similar to the one we used focal observations (Lehner, 1996) described above, was conducted. To to determine