Zootaxa 1807: 1–25 (2008) www.mapress.com / zootaxa/ ISSN 1175-5326 (print edition) Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition) ZOOTAXA The reef fish assemblage of the Laje de Santos Marine State Park, Southwestern Atlantic: annotated checklist with comments on abundance, distribution, trophic structure, symbiotic associations, and conservation OSMAR J. LUIZ JR.1,2,7, ALFREDO CARVALHO-FILHO2,3, CARLOS E. L. FERREIRA4, SERGIO R. FLOETER5, JOÃO LUIZ GASPARINI 6 & IVAN SAZIMA1,8 1 Depto. de Zoologia e Museu de História Natural, Universidade Estadual de Campinas, Campinas, SP, Brasil. E-mail: osmarluizjr@terra.com.br; isazima@unicamp.com 2 Instituto Laje Viva, www.lajeviva.org.br. Santos, SP, Brasil 3 Fish Bizz Ltda. Rua Moncorvo Filho 51, São Paulo, SP, Brasil. E-mail: alfie@telnet.com.br 4 Depto. de Biologia Marinha, Universidade Federal Fluminense, Niterói, RJ, Brasil. E-mail: cadu@vm.uff.br 5 Lab. de Biogeografia e Macroecologia Marinha, Universidade Federal de Santa Catarina, Florianópolis, SC, Brasil. E-mail: floeter@ccb.ufsc.br 6 Depto. de Ecologia e Recursos Naturais, Universidade Federal do Espírito Santo, Av. Fernando Ferrari 514, Vitória, ES, Brasil. E-mail: gaspa.vix@terra.com.br 7 Corresponding author 8 Retired and associated as voluntary researcher Abstract A check-list containing 196 species of reef fishes recorded at the Laje de Santos Marine State Park is presented. Most of them occur along the tropical western Atlantic or occur on both sides of the Atlantic Ocean. A minor part ranges to the temperate rocky reefs of Patagonia or are endemics to southeastern Brazil. Moringua edwardsi, Antennarius multiocellatus, Scorpaena dispar, Aulostomus strigosus, Lutjanus buccanella, Mulloidichthys martinicus and Halichoeres penrosei have here their ranges extended southwards to the São Paulo coast. Basic data on species abundance at the study site, distribution of species between habitat types, trophic structure, feeding symbiotic associations, and conservation are commented upon. Key words: Western South Atlantic, Brazil, rocky reefs, reef fishes, species list and distribution, zoogeography, conservation, trophic structure, feeding symbioses Resumo Uma lista com 196 espécies de peixes recifais registradas é apresentada para o Parque Estadual Marinho da Laje de Santos. A maioria das espécies tem ocorrência comum a todo o Atlântico Ocidental tropical ou ocorre nos dois lados do Oceano Atlântico. Uma parte menor das espécies tem distribuição em comum com os recifes rochosos temperados da Patagônia ou é endêmica ao Sudeste do Brasil. Moringua edwardsi, Antennarius multiocellatus, Scorpaena dispar, Aulostomus strigosus, Lutjanus buccanella, Mulloidichthys martinicus and Halichoeres penrosei tem aqui seu limite meridional de distribuição estendido ao Estado de São Paulo. Informações básicas sobre abundância das espécies no local de estudo e sua distribuição entre os diferentes tipos de habitats, estrutura trófica, associações alimentares simbióticas e conservação são aqui comentadas. Accepted by L. A. Rocha: 24 April 2008; published: 23 Jun. 2008 1 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. Introduction The Laje de Santos State Marine Park (Parque Estadual Marinho da Laje de Santos - PEMLS) is located on the southeastern coast of Brazil (24º15´S; 46º10´W), 36 km off the city of Santos, São Paulo State (Fig. 1). It consists of an uninhabited islet and several sparse rocky reefs with extensive intervening sand bottoms. The maximum depth is about 45 m. The subtidal substrate is composed of granitic boulders of varying sizes and shapes that delineate a steep profile (Fig. 2). The rocky substrate is mainly covered with patches of brown and red algae, the zoanthid Palythoa caribeorum, hydrozoans, ascidians, octocorals and sparse colonies of the scleractinian corals Madracis decactis and Mussismilia hispida. The local setting is a transitional tropical-subtropical environment that fits the definition of a high latitude ‘marginal’ reef site (Perry & Larcombe 2003) where hard corals may occur only as isolated colonies on the exposed bedrock. FIGURE 1. The Laje de Santos Marine State Park location in western South Atlantic. Photo: C.L.B. Francini. The area is of particular interest as this is the only Marine Protected Area totally closed to fishing in the most populated and developed region of Brazil, allowing researchers to evaluate fishing impacts on local rocky reefs (Floeter et al. 2006). Since 1993 a 5,000 ha area around the Laje de Santos Island was declared a protected area. After some years as a “paper park” with low or even inexistent effective enforcement, the study site is experiencing a great increase in the effort to eradicate fishing in the last four years. Since 2003, a new official government management policy and the rise of an active NGO dedicated to protect the Park brought new boats, equipment and personnel (volunteer and staff) extensively engaged in the inspection of the Park boundaries. Recreational activities allowed in the Park include scuba diving and boating. Anchoring is not allowed anymore and mooring buoys have been used by dive boats. Despite a few recent publications on reef fish taxonomy (Moura 1995), behavior (Sazima et al. 2000) and a species list of benthic algae (Amado-Filho et al. 2006), the Laje de Santos remains largely unknown biologically and a comprehensive list of reef fish species occurring in the Marine Park is still lacking. The knowl- 2 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. edge of the species composition of a community is instrumental to provide a baseline for future management and a starting point to produce scientific information needed for the process of designing and evaluating Marine Protected Areas, such as biomass, dispersal patterns, recruitment dynamics, trophic interactions and habitat preferences of the resident organisms (Agardy 2000, Craig et al. 2004). The present study provides the first comprehensive checklist of the reef fish species that occur at the Laje de Santos Marine State Park. In addition, patterns of species distribution, abundance, zoogeography, trophic types, representative symbiotic associations, and conservation issues are commented upon. Material and methods The species list results from fishes observed, photographed, and collected by the authors, as well as museums vouchers and reliable literature records. Bony fishes are listed in the phylogenetic order of families following Nelson (2006); elasmobranchs are listed following Compagno (1999). Species are organized in alphabetical order within families. Recent changes in the classification of the Serranidae as proposed by Craig and Hastings (2007) and Smith and Craig (2007) are adopted. Among these are the resurrection of the family Epinephelidae as distinct from Serranidae; the placement of the genus Paranthias within Cephalopholis; the change from genus Epinephelus to Mycteroperca for M. marginata (formerly E. marginatus) and the genus change of deep-bodied groupers of the Epinephelus niveatus complex to the resurrected genus Hyporthodus. In the list, we included the following information for each species: Abundance. an indicator of the relative abundance in the last five years, period when the first author started quantitative visual censuses of fishes using SCUBA in the study area and in more than thirty years of observations on fishes, by SCUBA and fishing, done by the second author; this indicator is based on a diver’s likelihood of recording a species in its normal habitat and depth range on any given dive (modified from Humann & DeLoach 2002, Feitoza et al. 2003), where: AB = abundant (several sightings of many individuals – at least 50 – are expected on nearly every dive), VC = very common (several sightings are expected on nearly every dive, but not necessarily of many individuals), CO = common (sights are frequent, but not necessarily expected on every dive), OC = occasional (sightings are not unusual, but are not expected on a regular basis), UN = unusual (sights occurs less than occasionally), and RA = rare (sights are exceptional). Habitat and distribution. The particular place where a species has usually been found; we arbitrarily stipulated different habitats types based on physiographic factors like substrate type and depth (fig. 2), where: Sh = Shallow reef (rocky substrate from 0 to 12 m depth), RS = Reef slope (rocky substrate from 13 to 20 m depth), SB = Sand bottom (sandy substrate adjacent to the rocky reef slope), WC = Water column (pelagic environment from 0 to 10 m depth, adjacent to the rocky reef but distant at least ~3 m from the bottom), and DR = Deep reef (rocky substrate in the range of 30–45 m depth). Geographic range of the species. namely: Br = Brazilian province (sensu Briggs 1974), CE = Central Atlantic (Islands of St. Helena and Ascension), CT = Circumtropical, EA= Eastern Atlantic, Pat = Patagonian (occur primarily in the temperate rocky reefs south to Argentina), SCa = Southern Caribbean (Coast of Venezuela, Trinidad and Tobago and other islands of the low lesser Antilles), SE = Southeastern Brazil (endemic from the region that encompass 20°S to 27S° in the Western Atlantic), TA = Trans-Atlantic (occur at both sides of the Atlantic Ocean), and WA = Western Atlantic (occur in Northern and Southern West Atlantic). Trophic category. assessed from direct behavioral observations and from the literature (Randall 1967, 1996, Carvalho-Filho 1999, Ferreira et al. 2004), where: CAR = Carnivores (eat a variety of mobile organisms, including invertebrates and fishes), MIF = Mobile invertebrate feeders (feed primarily on small benthic mobile invertebrates like mollusks, crustaceans, worms, etc. associated to the hard- or nearby soft-substrate), OMN = Omnivores (feed on variety of organisms, including animal and vegetal), PIS = Piscivores (feed only or mostly on live fishes), PLK = Planktivores (feed primarily on macro- and micro-zooplankton), ROVH = REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 3 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. Roving herbivores (non-territorial, large herbivores which includes in their diet a rich mass of detritus, turf algae and macroalgae), SIF = Sessile invertebrate feeders (feeds on a array of sessile benthic invertebrates like cnidarians, bryozoans, ascidians and sponges that are most associated to hard substrata), and TERH = Territorial herbivores (with a diet composed mainly by turf algae farmed within a vigorously defended territory). FIGURE 2. Habitat types found at the Laje de Santos Marine State Park. Record type. how the species was recorded, where: COL = Collected, MUS = Museum voucher, LIT = Literature, PHO = Photograph, and SIG = Sighting. For specimens deposited in museum collections, the institution and voucher number are provided in the appendix. We define reef fishes here as those species that are primarily associated with hard substrata, and which depend on the reef or it’s immediate vicinity for shelter and food during any phase of its post-settlement life. We also consider as reef fishes the epipelagic species that regularly visit reefs in search of food, cleaning services, and reproduction. For the purpose of describing the occurrence patterns of fish species in the different habitats, a cluster analysis based on a species presence/absence matrix was performed. The Bray-Curtis dissimilarity index was used, and habitats were clustered according to the UPGMA method (Pielou 1984). The abundance patterns of reef fish in the different habitats, considering variables such as exposure degree, substrate complexity and depth will be presented elsewhere. Results and discussion Species composition A total of 196 reef fish species in 124 genera in 66 families were recorded at the Laje de Santos (Table I). The most species rich families were Carangidae (16), Epinephelidae (12), Labridae (10), Pomacentridae (8) and Scaridae (8). Caranx, Gymnothorax, Halichoeres, Mycteroperca and Sparisoma, with 5 species each were the most species rich genera, followed by Chromis, Haemulon, and Seriola with 4 species. Twelve species were abundant (6%) (fig. 3), 26 (13%) were very common, 45 (23%) were common, 61 (32%) were occasional, 14 (7%) were unusual and 38 (19%) were rare. 4 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE I. List of families and species, habitat, abundance, geographic range, trophic category and record status of the reef fishes recorded at the Laje de Santos Marine State Park. Families of elasmobranchs are arranged following Compagno (1999) and bony fishes are arranged according to Nelson (2006); genera and species are arranged in alphabetical order. Habitat: DR = Deep Reef; RS = Reef Slope; SB = Sand Bottom; SH = Shallow Reef; WC = Water Column. Occurrence: AB = Abundant; CO = Common; OC = Occasional; RA = Rare; UN = Unusual; VC = Very Common. Geographic Range: Br = Brazilian Province; CE = Central Atlantic; CT = Circumtropical. EA = Eastern Atlantic; SCa = Southern Caribbean; SE = Southeastern Brazil; TA = Trans-Atlantic; WA = Western Atlantic. Trophic Category: CAR = Carnivore; MIF = Mobile Invertebrate Feeder; OMN = Omnivore; PIS = Piscivore; PLK = Planktivore; ROVH = Roving Herbivore; SIF = Sessile Invertebrate Feeder; TERH = Territorial Herbivore. Record Type: COL = Collected; LIT = in litteris; MUS = Museum Voucher; PHO = Photograph; SIG = Sighting. Family and species Habitat Occurrence Geog. Range Trophic Category Record Type DR UN CT PIS SIG WC RA CT PIS SIG Carcharhinus brevipinna (Muller & Henle, 1839) WC RA CT PIS COL, PHO Carcharhinus falciformis (Bibron, 1839) WC RA CT PIS COL, PHO Carcharhinus limbatus (Muller & Henle, 1839) WC RA CT PIS COL, PHO Carcharhinus longimanus (Poey, 1861) WC RA CT PIS PHO SB RA WA MIF SIG RS, SB RA TA MIF PHO Dasyatis centroura (Mitchill, 1815) RS, SB OC TA MIF PHO Dasyatis hypostigma Santos & Carvalho, 2004 RS, SB RA SE MIF PHO WC CO CT MIF PHO Manta birostris (Walbaum, 1792) WC OC CT PLK PHO Mobula hypostoma (Bancroft, 1831) WC RA CT PLK MUS, SIG Mobula japanica (Muller & Henle, 1841) WC RA CT PLK SIG Mobula tarapacana (Philippi, 1892) WC RA CT PLK LIT* ODONTASPIDIDAE Carcharias taurus Rafinesque, 1810 ALOPIIDAE Alopias vulpinus (Bonaterre, 1788) CARCHARHINIDAE RHINOBATIDAE Zapteryx brevirostris (Müller & Henle, 1841) GYMNURIDAE Gymnura altavela (L., 1758) DASYATIDAE MYLIOBATIDAE Aetobatus narinari (Euphrasen, 1790) MOBULIDAE * Gadig & Sampaio, 2002 MORINGUIDAE Moringua edwardsi (Jordan & Bollman, 1889) SH, RS UN WA MIF PHO Gymnothorax funebris Ranzani, 1839 SH, RS OC WA CAR MUS, PHO Gymnothorax miliaris (Kaup, 1856) SH, RS CO TA CAR COL, PHO Gymnothorax moringa (Cuvier, 1829) SH, RS CO WA+CE CAR COL, PHO Gymnothorax ocellatus Agassiz, 1831 SB RA WA CAR PHO Gymnothorax vicinus (Castelnaul, 1855) SH, RS CO TA CAR COL, PHO Muraena retifera Goode & Bean, 1882 RS, DR OC WA CAR COL, PHO MURAENIDAE to be continued. REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 5 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Record Type Category OPHICHTHIDAE Ahlia egmontis (Jordan, 1884) SH, RS UN WA MIF COL, PHO Myrichthys ocellatus (LeSueur, 1825) SH CO WA MIF PHO Myrichthys breviceps (Richardson, 1848) SH, RS RA WA MIF PHO Ophichthus ophis (L., 1758) SB RA TA PIS MUS Harengula clupeola (Cuvier, 1829) WC OC WA PLK COL Sardinella janeiro (Eingenmann, 1894) WC OC WA PLK MUS Synodus foetens (L., 1776) SH, RS, SB UN WA PIS MUS Synodus intermedius (Spix & Agassiz, 1829) SH, RS, SB VC WA PIS MUS, COL, PHO Synodus synodus (L., 1758) SH, RS, SB CO TA PIS COL, PHO SB RA SE+Pa CAR MUS SH UN WA CAR PHO RS, SB UN WA CAR PHO WC OC CT PIS MUS, PHO Hemiramphus balao (Lesueur, 1821) WC OC TA OMN SIG Hemiramphus brasiliensis (L., 1758) WC OC TA OMN SIG Holocentrus adscensionis (Osbeck, 1765) SH, RS VC TA MIF MUS, COL, PHO Myripristis jacobus Cuvier, 1829 SH CO TA PLK MUS, COL, PHO Hippocampus reidi Ginsburg, 1933 RS RA WA PLK PHO Micrognathus crinitus (Jenyns, 1842) RS OC WA PLK PHO SH RA TA CAR PHO RS, SB OC TA PIS SIG RS, SB CO TA MIF MUS, COL, PHO Scorpaena dispar Longley & Hildebrand, 1940 SH, RS, DR OC WA CAR PHO Scorpaena isthmensis Meek & Hildebrand, 1928 ? RA WA CAR MUS Scorpaena plumieri Bloch, 1789 SH, RS, DR OC WA+CA CAR COL, PHO Scorpaenodes tredecimspinosus (Metzelaar, 1919) SH, RS CO WA COL, PHO CLUPEIDAE SYNODONTIDAE BATRACHOIDIDAE Porichthys porosissimus (Cuvier, 1829) ANTENNARIIDAE Antennarius multiocellatus (Valenciennes, 1837) OGCOCEPHALIDAE Ogcocephalus vespertilio (L., 1758) BELONIDAE Tylosurus acus (Lacepede, 1803) HEMIRAMPHIDAE HOLOCENTRIDAE SYNGNATHIDAE AULOSTOMIDAE Aulostomus strigosus Wheeler, 1955 FISTULARIIDAE Fistularia tabacaria L., 1758 DACTYLOPTERIDAE Dactylopterus volitans L., 1758 SCORPAENIDAE CAR to be continued. 6 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Record Type Category SERRANIDAE Acanthistius brasilianus (Cuvier, 1828) RS, DR CO SE+Pat CAR MUS, COL, PHO Acanthistius patachonicus (Jenyns, 1840) DR RA SE+Pat CAR COL, PHO Diplectrum formosum (L., 1766) SB UN WA CAR PHO Dules auriga Cuvier, 1829 DR CO SE+Pat CAR COL, PHO Pronotogrammus martinicensis (Guichenot, 1868) DR UN WA PLK SIG Serranus baldwini (Evermann & Marsh, 1899) SH, RS CO WA CAR MUS, PHO Cephalopholis fulva (L., 1758) ? RA WA CAR MUS Cephalopholis furcifer (Valenciennes, 1828) WC CO TA PLK MUS, COL, PHO Epinephelus adscensionis (Osbeck, 1765) SH, RS RA TA CAR MUS, COL, PHO Epinephelus itajara Lichtenstein, 1822 SH, RS OC TA CAR PHO Epinephelus morio (Valenciennes, 1828) RS UN WA CAR COL, PHO Hyporthodus flavolimbatus (Poey, 1865) DR OC WA CAR COL, PHO Hyporthodus niveatus (Valenciennes, 1828) SH, RS, DR CO WA CAR MUS, COL, PHO Mycteroperca acutirostris (Valenciennes, 1828) SH, RS VC WA PIS MUS, COL, PHO Mycteroperca bonaci (Poey, 1860) SH, RS RA WA PIS COL, PHO Mycteroperca interstitialis (Poey, 1860) SH, RS CO WA PIS MUS, COL, PHO Mycteroperca marginata (Lowe, 1834) SH, SB, RS, DR VC SE+Pat+ CAR EA MUS, COL, PHO Mycteroperca venenosa (L., 1758) SH, RS, DR RA WA PIS COL, PHO Cookeolus japonicus (Cuvier, 1829) DR OC CT PLK COL, PHO Heteropriacanthus cruentatus (Lacepède, 1801) SH, RS OC CT PLK COL, PHO Priacanthus arenatus Cuvier, 1829 SH, RS, DR OC TA MIF COL, PHO Apogon americanus Castelnau, 1855 SH, RS CO Br PLK MUS, COL, PHO Apogon pseudomaculatus Longley, 1932 SH, RS CO TA PLK MUS, COL, PHO Caulolatilus chrysops (Valenciennes, 1833) SB RA WA CAR MUS Malacanthus plumieri (Bloch, 1786) SB CO WA+CE CAR MUS, PHO WC OC CT CAR COL, PHO Echeneis naucrates (L., 1758) WC OC CT CAR COL, PHO Remora albescens (Temminck & Schlegel, 1845) WC OC CT CAR PHO Remora remora (L., 1758) WC OC CT MIF PHO SB OC CT CAR COL, PHO EPINEPHELIDAE PRIACANTHIDAE APOGONIDAE MALACANTHIDAE POMATOMIDAE Pomatomus saltatrix (L., 1766) ECHENEIDAE RACHYCENTRIDAE Rachycentron canadum (L., 1766) to be continued. REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 7 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Category Record Type CT CAR COL CORYPHAENIDAE Coryphaena hippurus L., 1758 WC RA CARANGIDAE CT Alectis ciliaris (Bloch, 1787) WC CO CT CAR COL, PHO Caranx bartholomaei (Cuvier, 1833) WC OC WA PIS COL, PHO Caranx crysos (Mitchill, 1815) WC OC TA CAR COL, PHO Caranx hippos (L., 1766) WC RA TA CAR COL, PHO Caranx latus Agassiz, 1831 WC OC TA CAR MUS, PHO Caranx ruber (Bloch, 1793) WC OC WA CAR PHO Decapterus macarellus (Cuvier, 1833) WC OC CT PLK COL, PHO Decapterus punctatus (Cuvier,1829) WC UN TA PLK COL, PHO Pseudocaranx dentex (Bloch & Schneider, 1801) WC, SH, RS, VC SB, DR CT MIF, PLK MUS, COL, PHO Seriola dumerilli (Risso, 1810) WC CO CT CAR MUS, COL, PHO Seriola fasciata (Bloch, 1793) WC UN TA CAR COL, PHO Seriola lalandi Valenciennes, 1833 WC OC CT CAR COL, PHO Seriola rivoliana (Valenciennes, 1833) WC OC CT PIS COL, PHO Trachinotus falcatus (L.,758) WC OC WA CAR PHO Trachinotus goodei Jordan & Evermann,1896 WC VC WA CAR PHO Uraspis secunda (Poey, 1860) WC OC CT CAR COL, PHO Lutjanus analis (Cuvier, 1828) SH, RS VC WA CAR PHO Lutjanus buccanella (Cuvier, 1828) DR RA WA CAR COL, PHO Lutjanus cyanopterus (Cuvier,1828) SH, RS, DR OC WA CAR PHO Ocyurus chrysurus (Bloch, 1791) WC RA WA CAR COL, PHO Pristipomoides aquilonaris Goode & Bean, 1896) DR OC WA PIS COL, PHO Rhomboplites aurorubens (Cuvier, 1829) WC CO WA CAR, PLK MUS, COL, PHO WC OC CT CAR COL, PHO Anisotremus surinamensis (Bloch, 1791) SH, RS VC WA MIF COL, PHO Anisotremus virginicus (L., 1758) SH, RS, WC, DR AB WA MIF COL, PHO Haemulon aurolineatum Cuvier, 1830 SH, RS, WC, DR AB WA MIF COL, PHO Haemulon parra (Desmarest, 1823) RS OC WA MIF COL, PHO Haemulon plumierii (Lacepède, 1801) RS CO WA MIF COL, PHO Haemulon steindachneri (Jordan & Gilbert, 1882) RS CO WA MIF COL, PHO Calamus bajonado (Bloch & Schneider, 1801) RS RA WA MIF COL, PHO Calamus mu Randall & Caldwell, 1966 RS, SB RA SE MIF MUS LUTJANIDAE LOBOTIDAE Lobotes surinamensis (Bloch, 1790) HAEMULIDAE SPARIDAE to be continued. 8 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Record Type Category Calamus pennatula Guichenot, 1868 SB CO WA MIF COL, PHO Diplodus argenteus (Valenciennes, 1830) WC, SH, RS AB WA OMN COL, PHO Pagrus pagrus (L., 1758) SB, DR VC TA MIF MUS, COL, PHO Odontoscion dentex (Cuvier, 1830) SH, RS CO WA CAR PHO Pareques acuminatus (Bloch & Schneider, 1801) SH, RS CO WA CAR MUS, PHO Mulloidichthys martinicus (Cuvier, 1829) RS, SB RA TA MIF PHO Pseudupeneus maculatus (Bloch, 1793) RS, SB VC WA MIF MUS, COL, PHO SH VC WA PLK MUS, PHO Chaetodon sedentarius Poey, 1860 SH, RS UN WA MIF MUS, PHO Chaetodon striatus L., 1758 SH, RS VC WA SIF MUS, PHO Prognathodes brasiliensis Burgess, 2001 DR OC Br MIF PHO Prognathodes guyanensis (Durand, 1960) DR RA WA MIF PHO Centropyge aurantonotus Burgess, 1974 SH, RS OC TA TERH SIG Holacanthus ciliaris (L., 1758) SH, RS OC WA SIF PHO Holacanthus tricolor (Bloch, 1795) SH OC WA SIF PHO Pomacanthus paru (Bloch, 1787) SH, RS, WC VC WA OMN PHO Kyphosus incisor (Cuvier, 1831) SH, RS, WC AB TA ROVH MUS, COL, PHO Kyphosus sectator (L., 1766) SH, RS, WC AB TA ROVH COL, PHO Abudefduf saxatilis (L., 1758) SH, WC AB CT OMN COL, PHO Chromis cf. enchrysura Jordan & Gilbert, 1882 RS, DR OC WA MIF PHO Chromis flavicauda (Günther, 1880) RS CO Br PLK MUS, COL, PHO Chromis jubauna Moura, 1995 RS, DR AB Br+SCa PLK MUS, COL, PHO Chromis multilineata (Guichenot, 1853) WC, SH AB TA PLK MUS, COL, PHO Stegastes fuscus (Cuvier, 1830) SH, RS AB Br TERH COL, PHO Stegastes pictus (Castelnau, 1855) RS CO Br+SCa TERH COL, PHO Stegastes cf. variabilis (Castelnau, 1855) SH, RS CO WA TERH COL, PHO Bodianus pulchellus (Poey, 1860) SH, RS, DR VC TA MIF MUS, COL, PHO Bodianus rufus (L., 1758) SH, RS VC WA MIF MUS, COL, PHO Clepticus brasiliensis Heiser, Moura & Robertson, 2000 WC OC Br PLK MUS, COL, PHO Doratonotus megalepis Günther, 1862 RS RA TA MIF PHO Halichoeres sp n. SB, DR CO SE MIF COL, PHO SCIAENIDAE MULLIDAE PEMPHERIDAE Pempheris schomburgki Müller & Troschel, 1848 CHAETODONTIDAE POMACANTHIDAE KYPHOSIDAE POMACENTRIDAE LABRIDAE to be continued. REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 9 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Record Type Category Halichoeres brasiliensis (Bloch, 1791) SH, RS, SB CO Br MIF COL, PHO Halichoeres dimidiatus (Agassiz, 1831) RS VC Br MIF COL, PHO Halichoeres penrosei (Starks, 1913) SH, RS OC Br MIF SIG Halichoeres poeyi (Steindachner, 1867) SH, RS AB WA MIF MUS, COL, PHO Thalassoma noronhanum (Boulenger, 1890) WC, SH OC Br PLK MUS, PHO Cryptotomus roseus Cope, 1871 RS CO WA ROVH PHO Scarus trispinosus Valenciennes, 1840 SH RA Br ROVH PHO Scarus zelindae Moura, Figueiredo & Sazima, 2001 SH, RS CO Br ROVH PHO Sparisoma amplum (Ranzani, 1842) SH, RS CO Br ROVH PHO Sparisoma axillare (Steindachner, 1878) SH, RS VC Br ROVH COL, PHO Sparisoma frondosum (Agassiz 1831) SH, RS VC Br+SCa ROVH PHO Sparisoma radians (Valenciennes, 1840) RS CO WA ROVH MUS, SIG Sparisoma tuiupiranga Gasparini, Joyeux & Floeter, RS 2003 VC SE ROVH PHO DR OC SE+Pat CAR PHO SH, RS CO WA MIF COL Labrisomus nuchipinnis (Quoy & Gaimard, 1824) SH VC TA MIF MUS, COL, PHO Labrisomus kalisherae (Jordan, 1904) SH OC WA MIF PHO Malacoctenus delalandii (Valenciennes, 1836) SH CO WA MIF PHO Starksia brasiliensis (Gilbert, 1900) SH CO Br MIF PHO SH, RS VC WA MIF MUS, COL, PHO Hypleurochilus fissicornis (Quoy & Gaimard,1824) SH OC Br MIF PHO Hypsoblennius invemar Smith-Vaniz & Acero, 1980 SH CO WA MIF PHO Ophioblennius trinitatis Miranda-Ribeiro,1919 SH RA Br TERH MUS, SIG Parablennius marmoreus (Poey, 1876) SH VC WA MIF PHO Parablennius pilicornis (Cuvier, 1829) SH, RS, DR AB TA MIF MUS, COL, PHO Scartella cristata (L., 1758) SH OC CT TERH MUS, COL, PHO SH,, RS CO WA MIF COL Coryphopterus glaucofraenum Gill, 1863 RS, SB VC WA OMN COL, PHO Ctenogobius saepepallens (Gilbert & Randall, 1968) RS, SB OC WA OMN SIG Elacatinus figaro Sazima, Moura & Rosa, 1997 SH, RS VC Br MIF COL, PHO SCARIDAE PINGUIPEDIDAE Pinguipes brasilianus Cuvier, 1829 TRIPTERYGIIDAE Enneanectes altivelis Rosenblatt, 1960 LABRISOMIDAE CHAENOPSIDAE Emblemariopsis signifera (Ginsburg, 1942) BLENNIIDAE CALLYONIMIDAE Callionymus bairdi Jordan, 1887 GOBIIDAE to be continued. 10 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. TABLE 1. (continued) Family and species Habitat Occurrence Geog. Range Trophic Record Type Category Gnatholepis thompsoni Jordan, 1902 RS, SB OC TA OMN SIG SB CO Br+SCa MIF PHO WC CO WA SIF COL, PHO Acanthurus bahianus Castelnau, 1855 SH CO WA ROVH COL, PHO Acanthurus coeruleus Bloch & Schneider, 1801 SH RA WA ROVH PHO Acanthurus chirurgus (Bloch, 1787) SH, RS AB TA ROVH COL, PHO Acanthurus monroviae Steindachner, 1876 SH, RS OC SE+EA ROVH PHO Sphyraena barracuda (Edwards, 1771) WC RA CT PIS PHO Sphyraena tome Fowler, 1903 WC OC SE PIS PHO WC OC TA CAR PHO Bothus maculiferus (Poey, 1860) SB OC WA CAR PHO Bothus ocellatus (Agassiz, 1831) SB CO WA CAR PHO Balistes vetula L., 1758 SH, RS OC TA MIF PHO Melichthys niger (Bloch, 1786) WC RA CT OMN SIG Aluterus monoceros (L., 1758) SH, RS, WC UN CT CAR PHO Aluterus scriptus (Osbeck, 1765) SH, RS OC CT SIF SIG Cantherhines macrocerus (Hollard, 1853) SH, RS OC WA SIF PHO Cantherhines pullus (Ranzani, 1842) SH UN TA OMN MUS, SIG Stephanolepis hispidus (L., 1766) SH, RS UN TA MIF SIG RS CO WA SIF PHO Canthigaster figueiredoi Moura & Castro, 2002 SH, RS VC Br+SCa MIF MUS, PHO Sphoeroides spengleri (Bloch, 1785) SH, RS VC WA MIF MUS, COL, PHO Chilomycterus spinosus (L., 1758) SH, RS OC WA MIF MUS, SIG Diodon hystrix L., 1758 RS OC CT MIF SIG WC RA CT CAR PHO MICRODESMIDAE Ptereleotris randalli Gasparini, Rocha & Floeter, 2001 EPHIPPIDAE Chaetodipterus faber (Broussonet, 1782) ACANTHURIDAE SPHYRAENIDAE SCOMBRIDAE Euthynnus alleteratus (Rafinesque, 1810) BOTHIDAE BALISTIDAE MONACANTHIDAE OSTRACIIDAE Acanthostracion polygonius Poey, 1876 TETRAODONTIDAE DIODONTIDAE MOLIDAE Mola mola (L., 1758) REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 11 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 3. Some abundant fish species at the Laje de Santos Marine State Park. The tomtate grunt Haemulon aurolineatum (a); adult and juvenile sergeant major Abudefduf saxatilis (b); the brown chromis Chromis multilineata (c); juvenile dusky damselfish Stegastes fuscus (d); intermediate individual of the jubauna reeffish Chromis jubauna (e); the silver porgy Diplodus argenteus (f); juvenile porkfish Anisotremus virginicus (g); the ringneck blenny Parablennius pilicornis (h). Photos: O.J. Luiz Jr, except (e) by L.F. Cassino. 12 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. Geographical Distributions and Zoogeography The relative proportion of geographic distribution types is shown on Fig.4. Forty two percent of the species (86) occur in the entire Western Atlantic, 19% (40) are trans-Atlantic, 18% (37) are circumtropical, and 12% (22) are endemic to the Brazilian coast (Fig. 5), 3% (6) are distributed southwards to temperate Patagonia, 2% (3) are found in the Western Atlantic and the islands of the mid-Atlantic (Ascension and Sta. Helena) and 1% (2) are found in the eastern Atlantic and southeastern coast of Brazil but neither reach northern sites of Brazil and the Northwestern Atlantic. Five species (3%) present a curious distribution pattern: they occur along the Brazilian coast but are limited to the Southern Caribbean in the Northwest Atlantic (coast of Venezuela, Trinidad & Tobago, Barbados and Curacao). It has been hypothesized that these five species have a Brazilian origin and only recently crossed to the north of the Amazon Barrier (Joyeux et al. 2001, Rocha 2003). Perhaps the most surprising and unexpected finding at the Laje the Santos Marine State Park was the occurrence of Acanthurus monroviae individuals (fig. 7a), a surgeonfish thought to be restricted to the Eastern Atlantic. This record was described in detail by Luiz Jr. et al. (2004). FIGURE 4. Relative proportions of geographic distribution types of the species observed at the Laje de Santos Marine State Park. Br = Brazilian Province; CE = Central Atlantic; CT = Circumtropical. EA = Eastern Atlantic; SCa = Southern Caribbean; SE = Southeastern Brazil; TA = Trans-Atlantic; WA = Western Atlantic. Despite the position of the study site at subtropical latitudes, the local reef fish fauna more closely resembles that of the tropical Western Atlantic and Northeastern Brazilian coast than that of the southern temperate reefs (Floeter et al. 2008). Due to the prevailing Brazil Current, which flows southwards from low latitudes, superficial warm waters continuously reach the Laje de Santos Marine Park, providing larval supply and a suitable thermal range for tropical reef species. Colder waters are also present at the study site due to a seasonal upwelling that brings cold (14-18ºC) deep waters from the shelf slope (Campos et al. 1995). Cold waters are usually restricted to the deepest parts of the reefs, providing a suitable environment for temperate species (fig. 6). Thus, the species that range south to Patagonia and the subtropical endemics were observed mostly on, or are even restricted to the deep reefs at our study site. The mixture of different conditions in ecological transitional areas is regarded as a major factor for a high diversity in SE Brazilian reef systems (Floeter et al. 2001). Why are there many more tropical than subtropical/temperate species is a subject of future research. One possible cause is that the deep reefs were poorly sampled and an increase in the sampling effort could result in additional new records of cold water species for the region. Few studies with the use of SCUBA were done by biologists at depths greater than 30 m, but on REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 13 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. the other hand commercial and recreational fisheries were present for many decades in the surrounding areas of the study site, providing a fairly good assessment of fish species from all depths over the study area. Thus, we could expect to find some unrecorded species in the deep reefs that could only be found by SCUBA and not by conventional fishing gear, however, this is not expected to significantly change the zoogeographical affinities presented here. An alternative explanation to poor sampling may be that the larval input from the southern temperate reefs is limited. Waters from Patagonia are known to reach the southeastern coast of Brazil through the Malvinas current, which flows northward from the coast of Argentina and can carry larvae from temperate reefs to the study site (Pereira 1989, Campos et al. 1996). However, dispersal via Malvinas Current may be restricted to species with some tolerance to low salinity waters. The high freshwater outflow from La Plata River and Patos Lake may extend for a great distance offshore (Piola et al. 2000, Pimenta et al. 2005) and can act as a barrier to larval dispersal in the same way as the Amazon River does between the Caribbean and Brazil (Floeter & Gasparini 2000, Rocha et al. 2003). Range Extensions We consider here only species previously unrecorded for the coast of São Paulo State. We use this unnatural, political division because most books and species accounts of fishes in Brazil usually determine the limits of species’ ranges this way. Additionally, we feel that species whose recorded range limits lie only a few kilometers north or south of our study site do not represent an actual range increase. Thus, the Laje de Santos Marine Park position, approximately in the middle of São Paulo’s coast, distant 171 km from Rio de Janeiro State to the north and 215 km from Paraná State to the south, provides a convenient ‘buffer’ distance for range extensions. Species whose ranges are extended in this paper are listed below. Moringua edwardsi: Previously recorded for Trindade Island (Gasparini & Floeter 2001) and on the continental coast south to the State of Bahia (Menezes et al. 2003). Our record is based on a photograph made in July 2006 by Armando de Luca Jr. (fig. 7b). Antennarius multiocellatus: Previous southernmost record is at Arraial do Cabo, Rio de Janeiro State (Carvalho-Filho 1999, Ferreira et al. 2001). Our record is based on a photograph made in November 2004 by Robson Leite (fig. 7c). Scorpaena dispar: Southernmost occurrence previously recorded for the State of Rio de Janeiro (Menezes et al. 2003). Our record is based on a photograph made in July 2001 by Osmar J. Luiz Jr. (fig. 7d). Aulostomus strigosus: Previous southernmost record is the state of Rio de Janeiro (Carvalho-Filho 1999). Our record is based on a photograph made in January 2004 by Renata Linger (fig. 7e). Lutjanus buccanella: Previous southernmost record is Ilhéus, State of Bahia (Carvalho-Filho 1999, Menezes et al. 2003). Our record is based on a specimen 42 cm SL collected in October 1987 by Alfredo Carvalho-Filho (fig. 7f). Mulloidichthys martinicus: Previous southernmost record is the state of Rio de Janeiro (Carvalho-Filho 1999, Menezes et al. 2003). Our record is based on a photograph made in June 2002 by Osmar J. Luiz Jr. (fig. 7g). Halichoeres penrosei: Previous southernmost record is the state of Rio de Janeiro (Carvalho-Filho 1999, Menezes et al. 2003). Our record of this species is based on sightings of several individuals in January 2006 and 2007 by Alfredo Carvalho-Filho. These previously unrecorded species were stray individuals that were seen once or twice at the study site and do not seem to represent resident populations. This reflects the high potential for dispersal among reef fishes and their ability to reach distant areas when associated with spatially and temporally unusual currents. When they do reach new areas, site specific ecological factors and occurrence of competitors or predators could prevent the establishment of new populations, and in our case cold waters seem to prevent the establishment of the above mentioned species. 14 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 5. Selected Brazilian endemic reef fish species that occur at the Laje de Santos Marine State Park. The barber goby Elacatinus figaro (a); the Brazilian yellowcheek wrasse Halichoeres dimidiatus, initial phase (b); the Brazilian wrasse Halichoeres brasiliensis, intermediate phase (c); the Noronha wrasse Thalassoma noronhanum, terminal male (d); the tuiupiranga parrotfish Sparisoma tuiupiranga, initial phase (e); Zelinda’s parrotfish Scarus zelindae, initial phase (f); the reef parrotfish Sparisoma amplum, initial phase (g); the gray parrotfish Sparisoma axillare, terminal male (h). Photos: O.J. Luiz Jr, except (d) by I. Cavas. REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 15 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 6. Selected fish species that dwell in the deep reef (30-45 m) community at the Laje de Santos State Marine Park. The deep-reef wrasse Halichoeres sp.n. (a); the reeffish Chromis cf. enchrysura (b); the red porgy Pagrus pagrus (c); the Brazilian sandperch Pinguipes brasilianus (d); the sea basses Acanthistius brasilianus (e), A. patachonicus (f) and Dules auriga (g); the snowy grouper Hyporthodus niveatus (h). The former species (a) is probably a Brazilian endemic, closely related to the Northwestern Atlantic species H. bathyphilus. The distinctive status from its sister species is supported by molecular mtDNA analysis (L.A. Rocha pers. comm.).The last six species (c-h) ranges southward to temperate Patagonian rocky reefs. Photos: A. Carvalho-Filho (e-g); O.J. Luiz Jr. (a-d, h). 16 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 7. Unusual records of reef fish species for the State of São Paulo. The African surgeonfish Acanthurus monroviae (a), which record was dealt in details by Luiz Jr. et al. (2004) and the previously unrecorded species: the eel Moringua edwardsi (b); the frogfish Antennarius multiocellatus (c); the scorpionfish Scorpaena dispar (d); the trumpetfish Aulostomus strigosus (e); the blackfin snapper Lutjanus buccanella (f); the yellow goatfish Mulloidichthys martinicus (g). Except for (a), all records are southward extensions for these species’ ranges in the West Atlantic. Photos: A. Carvalho-Filho (f); R. Leite (c); R. Linger (e); A. de Luca Jr. (a, b); O.J. Luiz Jr. (d, g). REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 17 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. Trophic Structure More than a half of the species at the study site are carnivores and mobile invertebrate feeders, 28% of the species belonging in each of these categories, followed by planktivores (13%), piscivores (11%), roving herbivores (8%), omnivores (5%), sessile invertebrate feeders (4%), and territorial herbivores (3%). Mobile invertebrate feeders are expected to be the most species-rich trophic category on reef fish assemblages along the Brazilian coast due mostly to their diverse morphology and variety of prey items (Ferreira et al. 2004). Carnivores are mostly epinephelids, carangids and muraenids, three diverse families at the study site. The relatively high proportion of planktivores and piscivores may be attributed to the mid-shelf location of the study site. Distance from the coast is regarded as a good predictor for the occurrence and abundance of planktivorous reef fishes in southwestern Atlantic rocky reef system (Floeter et al. 2007). Most piscivores belong to epipelagic families (carangids, scombrids, pomatomids, sphyraenids) that usually are found far offshore, but this category also includes many species that dwell on sandy bottom, an extensive habitat at the study site. These patterns, however, were not consistent between the different zones of the study site. Habitat specific trophic structure was reflected in the similarity of clusters of species recorded for each habitat (fig. 8). The shallow reef and reef slope zones were almost identical and matched best for the whole assemblage at the study site. However, as expected, in the water column zone, planktivore and carnivore categories are better represented. The deep reef and the sand bottom habitats were similar in trophic structure patterns; both have no herbivores and no sessile invertebrate feeders, probably a result of the reduced algal coverage due to low light levels in the deep reef, and the lack of suitable substrate for attachment of algae. Symbiotic feeding associations As most reef assemblages in tropical or temperate regions (e.g., Fricke 1975, Strand 1988, Côté 2000, Grutter 2005, Sazima et al. 2007), the Laje de Santos has its share of symbiotic feeding relationships (see Losey 1978 for an instrumental definition and examples of symbiotic behavior). One such feeding association is cleaning symbiosis, in which a fish species (the cleaner) remove parasites, necrotic tissue, and mucus from a variety of fishes (the clients) that seek the cleaner’s services (reviews in Côté 2000, Grutter 2005). An obligate cleaner recorded at our study site is the barber goby Elacatinus figaro, which maintains well defined cleaning stations on prominent points of the reef and services a diverse and species-rich fish assemblage (see Sazima et al. 2000). The goby is a bottom-dweller but moves towards mid-water clients that hovers near the cleaning station (fig. 9a). Since cleaning symbiosis between fishes is a strictly diurnal activity, nocturnal species leave their shelters to seek cleaning (fig. 9b). Other cleaners recorded at the Laje de Santos are facultative and clean mostly as juveniles (Côté 2000), such as the porkfish Anisotremus virginicus and the spotfin hogfish Bodianus pulchellus (fig. 9c); this latter, however, occasionally cleans as a small adult as well (fig. 9d). Although occasionally recorded at the Laje de Santos, the versatile Noronha wrasse Thalassoma noronhanum was never recorded as a cleaner there, a role it apparently plays in Brazil’s oceanic islands only (e.g., Gasparini & Floeter 2001, Sazima et al 2005). Two common fishes at our study site, the silver porgy Diplodus argenteus and the sergeant major Abudefduf saxatilis, also remain to be recorded as cleaners there, although they occasionally clean at other coastal sites in southern and southeastern Brazil (Sazima 1986, Krajewski 2007). Apparently, the rarity or even absence of cleaning by these two latter species is related to the presence of more colorful and efficient cleaners such as the goby E. figaro and the wrasse B. pulchellus. The “weak” cleaning role of D. argenteus in southern Brazil seems related to its unattractiveness as compared to more specialized and colorful cleaners there (Krajewski 2007), and a similar idea may be applied to the cleaner guild of the Laje de Santos. Another conspicuous symbiotic association in reef assemblages is following behavior (revision in Sazima et al. 2007). In this association type, a benthic species (the nuclear) attracts other, opportunistic and mostly carnivorous species (the followers) during its feeding activity. Small animals exposed by the substrate-dis- 18 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. turbing activity of the nuclear fish and not taken by it may be preyed upon by the follower fish. Substrate feeders that dig into sandy bottoms are very prone to attract followers, as the sediment clouds are a clue to the latter (see Sazima et al. 2006). The best known and studied example are the goatfishes, family Mullidae, which are regarded as nuclear fishes par excellence (Sazima et al. 2006). Less studied in the following symbiosis is the role of jacks, family Carangidae, which may act both as nuclear and followers (Sazima 1988, Sazima et al. 2007). At our study site, the white trevally Pseudocaranx dentex (fig. 9e) acts as a nuclear species and attracts mostly wrasses, which are known for their follower role (Sazima et al. 2007). Morays and other eels are another fish group that is readily followed during foraging activity (Diamant & Shpigel 1985, Gerhardinger et al. 2006), and attracts mostly groupers (fig. 9f). Less important are several fishes that forage on sandy bottoms but need not raise sediment to attract followers. One such example is the flying gurnard Dactylopterus volitans (fig. 9g) whose moving close to the bottom disturb small fishes and other animals that are preyed upon by the followers (Sazima & Grossman 2005). The hitch-hiking remoras (Echeneididae) exemplify still another type of feeding association. Although widely known as hitch-hikers on larger fishes and other marine vertebrates, and feeding on scraps left by their hosts or cleaning them of parasites (review in O’Toole 2002), there is a less known feeding activity of remoras on their hosts. While attached near the mouth of a filter-feeding, large fish such as the Atlantic manta (fig. 9h) or a whale shark, the remoras engage in filter-feeding (Clarke & Nelson 1997), a role recorded for other species in the family (e.g., Sazima et al. 2006). Conservation Remarks With 196 recorded species, the reef fish assemblage of the Laje de Santos Marine State Park has a relatively high richness when compared to other localities along the Brazilian coast. Despite its higher latitude position (i.e., subtropical), our study site has an equal or even a larger number of species than similarly-sized tropical reefs of Northeastern Brazil (Rosa & Moura 1997, Feitoza 2001, Ferreira & Cava 2001, Rocha & Rosa 2001, Souza et al. 2007). This is probably due to the co-occurrence of tropical and subtropical/temperate species on Brazil’s southeastern coast. Additionally, there are more species at the Laje de Santos than at other similarly-sized coastal reef sites in southeastern and south Brazil (Ferreira et al. 2001, Hostim-Silva et al. 2006, Rangel et al. 2007), which could be explained by the off-shore, mid-shelf location of the Laje de Santos reefs and the consequent occurrence of a relatively large number of epipelagic species. Unlike coral reefs, rocky reefs are highly restricted to the shores of continents and islands (Ebeling & Hixon 1991) with few patchily distributed rocky bottoms at the mid-shelf. Thus, the Laje de Santos Marine State Park is to be regarded as an important biodiversity hot-spot for Brazilian reef fishes. Some species that are targeted and highly prized by spear-fishermen occasionally attain large sizes at the Laje de Santos (fig. 10). However, it would be premature to say that the occurrence of these endangered species is directly related with the protection conferred to the Laje de Santos without further investigation. Acknowledgements We thank Ana Paula Balboni and Guilherme Kodja, representing all the staff of the NGO ‘Instituto Laje Viva’ for their much valuable support and help in the field. Otto Gadig for the identification of D. hypostigma. Carlo Leopoldo B. Francini, Armando de Luca Jr., Maurício Andrade, João Paulo Krajewski, Lara Cheidde, Luis F. Cassino, Robson Leite, Renata Linger and Luciano Eneas for several photographs. Julio Vellardi (Instituto Florestal- SP) for the support to scientific research at the Laje de Santos State Marine Park (COTEC 42.635/ 2006). JLG Thanks Fausto Pires dos Campos and Guilherme Moura for support and help in the field work. Financial support was provided by CNPq (MSc. grant 135112/2007-9 to OJLJr, and productivity grants to CELF and IS). This paper is dedicated to Clóvis B. de Carvalho for his inspiring enthusiasm and effort in the conservation of ‘the big magic rock’. REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 19 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 8. Cluster analysis of habitat types at the Laje de Santos Marine State Park based on the similarity of species composition. The relative distribution of trophic categories in each habitat is shown in the graphs. CAR = Carnivore; MIF = Mobile Invertebrate Feeder; OMN = Omnivore; PIS = Piscivore; PLK = Planktivore; ROVH = Roving Herbivore; SIF = Sessile Invertebrate Feeder; TERH = Territorial Herbivore. 20 · Zootaxa 1807 © 2008 Magnolia Press LUIZ JR. ET AL. TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 9. Selected examples of symbiotic associations between reef fishes recorded at the Laje de Santos Marine State Park. The barber goby Elacatinus figaro cleans the head of the jubauna reeffish Chromis jubauna hovering close to the goby’s cleaning station (a); the same cleaner species inspects the back of the nocturnal squirrelfish Holocentrus adscensionis that approached its cleaning station (b); juvenile spotfin hogfish Bodianus pulchellus cleans the mouth of the spotted moray Gymnothorax moringa (c); adult of the same hogfish species cleans the head of the jubauna reeffish (d); the wrasse Halichoeres sp. n. follows a group of the white trevally Pseudocaranx dentex, which stir sediment clouds while feeding on the sandy bottom (e); the dusky grouper Mycteroperca marginata closely follows the goldspotted snake eel Myrichthys ocellatus that nudges its head in rocky crevices (f); the spotfin hogfish follows the flying gurnard Dactylopterus volitans moving close to the bottom (g); two diskfish Remora remora attached near the mouth of the Atlantic manta Manta birostris (h).Photos: M. Andrade (h); A. Carvalho Filho (e-f); J. P. Krajewski (a); O.J. Luiz Jr. (c-d, g); A. de Luca Jr. (b). REEF FISHES OF LAJE DE SANTOS MARINE PARK Zootaxa 1807 © 2008 Magnolia Press · 21 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. FIGURE 10. Targeted and endangered top-predators recorded at the Laje de Santos Marine State Park. The dusky groupers Mycteroperca marginata are very common in the area, but attain unusual large size and are largely unafraid of divers, contrarily to what happens at other unprotected sites (a); the goliath grouper Epinephelus itajara (b) and the cubera snapper Lutjanus cyanopterus (c). Several individuals of these two latter species have been seen at in the Laje de Santos in the last two years, after a period of more than ten years over which they remained unrecorded at the site. Photos: A. Carvalho-Filho (c); L. Cheidde (b); A. Valente (a). References Agardy, T. 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Appendix Museum vouchers of specimens from the Laje de Santos Marine State Park. Species arranged in alphabetical order. Institutions names and acronyms are the follow: California Academy of Sciences (CAS); Museu de Zoologia, Universidade de São Paulo (MZUSP); National Museum of Natural History, Smithsonian Institution (USNM); Museu de História Natural, Universidade Estadual de Campinas (ZUEC). Abudefduf saxatilis—MZUSP 50989; Acanthistius brasilianus—MZUSP 14890, 70738, 70743; Acanthurus bahianus— MZUSP 45659; Acanturus chirurgus—MZUSP 45661; Anisotrenus virginicus—MZUSP 45662, 67870, 67873; Apogon americanus—MZUSP 45641; Apogon pseudomaculatus—MZUSP 45635, 45643; Bodianus pulchellus— MZUSP 44589, 66230, 66238, 66242; ZUEC 3394, 4255, 4427; Bodianus rufus— MZUSP 44596, 66240; Calamus mu—MZUSP 69958, 70039; Calamus penna—MZUSP 14891, 14892; Cantherhines pullus—MZUSP 44658, 44660, 72924; Canthigaster figueiredoi—MZUSP 44594 (paratype), 44595; USNM 357498 (paratype); Caranx latus—MZUSP 43487; Caulolatilus chrysops—MZUSP 14878, 44650; Cephalopholis fulva—MZUSP 47436; Chaetodon sedentarius—MZUSP 45654; Chaetodon striatus— MZUSP 45644, 45649; Chilomycterus spinosus— MZUSP 5246, 5247, 5248, 5249, 71370; Chromis flavicauda MZUSP 44624, 44625, 44626; Chromis jubauna— MZUSP 44631 (holotype), 44632- 44635 (paratypes); ZUEC 4331, 4426; Chromis multilineata—MZUSP 44619, 44620; ZUEC 3916, 6310; Clepticus brasiliensis—CAS 99821-99822 (paratypes); MZUSP 44590, 44644, 44657, 47151 (paratype), 53271, 9982199822 (paratypes); Dactylopterus volitans—MZUSP 46986; Diplodus argenteus— MZUSP 45651; Dules auriga—MZUSP 70845; Elacatinus figaro—ZUEC 3898, 3902, 3903, 3911, 3912, 3914, 3915; Emblemariopsis signifera—MZUSP 44608; Epinephelus adscensionis—MZUSP 47440; Epinephelus morio – MZUSP 71106; Gymnothorax funebris - MZUSP 14877; Halichoeres sp n.—MZUSP 46825, 47152; Halichoeres poeyi—MZUSP 44584, 44585, 44586, 47435; Halichoeres brasiliensis—MZUSP 47437, 47438, 47441; Holocentrus adscensionis—MZUSP 43489, 47434, 45657, 47434; Hyporthodus niveatus—MZUSP 43490, 70937, 70949; Kyphosus incisor—43495, 44656; Labrisomus nuchipinnis—MZUSP 44614, 44615, 44616, 44617, 44618, 66796; ZUEC 6309; Lobotes surinamensis—MZUSP 72733; Malacanthus plumieri—MZUSP 14870; Mobula hypostoma – 13402 (head); Mycteroperca acutirostris—MZUSP 43494; Mycteroperca interstitialis—MZUSP 43497; 47150, 70975, 70976; Mycteroperca marginatus—MZUSP 43491, 51248, 70913; Myripristis jacobus—MZUSP 43488, 44645; Ophichthus ophis—MZUSP 44649; Ophioblennius trinitatis— MZUSP 44607; Pagrus pagrus—MZUSP 45647, 70094; Parablennius pilicornis—MZUSP 44599, 44601, 63869; Paranthias furcifer—MZUSP 43496, 71045, 71046; Pareques acuminatus—MZUSP 43493, 44647; Pempheris schomburgki—MZUSP 43492, 45645; Porichthys porosissimus—MZUSP 44651; Priacanthus arenatus—MZUSP 69931; Pseudocaranx dentex—MZUSP 14871, 45655; Pseudupeneus maculatus— MZUSP 41992, 43486; Remora albescens—MZUSP 69754; Rhomboplites aurorubens—MZUSP 44582; Sardinella janeiro—MZUSP 11411; Scartella cristata—MZUSP 44602; Scarus zelindae—USNM 357500 (paratype); Scorpaena isthmensis—MZUSP 43481; Serranus baldwini—MZUSP 43478, 43479; Seriola dumerili—MZUSP 14880, 46989; Seriola fasciata—MZUSP 46990; Sparisoma amplum— MZUSP 46444; Sparisoma axillare—MZUSP 46817; Sparisoma frondosum—MZUSP 46802, 46818; Sparisoma radians – MZUSP 46440; Sparisoma tuipiranga—MZUSP 46441, 46442, 46443; Sphoeroides spengleri—MZUSP 44579; Stegastes fuscus—MZUSP 45775, 45780, 45782, 45784, 45787, 45789, 49065; ZUEC 6309; Stegastes pictus—MZUSP 45786; Stegastes variabilis—MZUSP 45776, 45779, 45781, 45783, 45791; Synodus foetens— MZUSP 43483; Synodus intermedius—MZUSP 43484; Thalassoma noronhanum—MZUSP 45633, 45639; ZUEC 3146; Tylosurus acus- MZUSP 14879; Uraspis secunda—MZUSP 65901. 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