Atlantic Reef Fish Biogeography and Evolution

Home , Bathyanthias, Enneacampus, Enneanectes, Epinephelus marginatus, Gastropsetta, Gobiesocidae

Journal of Biogeography (J. Biogeogr.) (2008) 35, 22–47

SPECIAL Atlantic reef ﬁsh biogeography and PAPER evolution S. R. Floeter1,2*, L. A. Rocha3, D. R. Robertson4, J. C. Joyeux5, W. F. Smith- Vaniz6, P. Wirtz7, A. J. Edwards8, J. P. Barreiros9, C. E. L. Ferreira10, J. L. Gasparini5, A. Brito11, J. M. Falco´n11, B. W. Bowen3 and G. Bernardi12

1National Center for Ecological Analysis and ABSTRACT Synthesis, University of California, 735 State Aim To understand why and when areas of endemism (provinces) of the tropical St., 300, Santa Barbara, CA, 93101, USA, 2Departamento de Ecologia e Zoologia, Lab. de Atlantic Ocean were formed, how they relate to each other, and what processes Biogeografia e Macroecologia Marinha, have contributed to faunal enrichment. Universidade Federal de Santa Catarina, Location Atlantic Ocean. Floriano´polis, SC 88010-970, Brazil, 3Hawaii Institute of Marine Biology, University of Methods The distributions of 2605 species of reef fishes were compiled for Hawaii, Kaneohe, HI, 96744, USA, 25 areas of the Atlantic and southern Africa. Maximum-parsimony and distance 4Smithsonian Tropical Research Institute, analyses were employed to investigate biogeographical relationships among those Balboa, Unit 0948, APO AA 34002, Panama, areas. A collection of 26 phylogenies of various Atlantic reef fish taxa was used to 5Departamento de Ecologia e Recursos assess patterns of origin and diversification relative to evolutionary scenarios Naturais, Universidade Federal do Espı´rito based on spatio-temporal sequences of species splitting produced by geological Santo, Vito´ria, ES, 29060-900, Brazil, 6US and palaeoceanographic events. We present data on faunal (species and genera) Geological Survey, Florida Integrated Science richness, endemism patterns, diversity buildup (i.e. speciation processes), and Center, Gainesville, FL, 32653, USA, 7Centro evaluate the operation of the main biogeographical barriers and/or filters. de Cieˆncias do Mar, Universidade do Algarve, Faro, P-8000-117, Portugal, 8School of Biology, Results Phylogenetic (proportion of sister species) and distributional (number Newcastle University, Newcastle upon Tyne, of shared species) patterns are generally concordant with recognized NE1 7RU, UK, 9Departamento de Cieˆncias biogeographical provinces in the Atlantic. The highly uneven distribution of Agra´rias, Universidade dos Açores, 9701-851, species in certain genera appears to be related to their origin, with highest species Portugal, 10Departamento de Biologia richness in areas with the greatest phylogenetic depth. Diversity buildup in Marinha, Universidade Federal Fluminense, Atlantic reef fishes involved (1) diversification within each province, (2) isolation Niteroí, RJ, 24001-970, Brazil, 11Dpto. Biologıá as a result of biogeographical barriers, and (3) stochastic accretion by means of Animal (Ciencias Marinas), Universidad de La dispersal between provinces. The timing of divergence events is not concordant Laguna, La Laguna, Tenerife, Islas Canarias, among taxonomic groups. The three soft (non-terrestrial) inter-regional barriers 38206, Spain, 12Department of Ecology and (mid-Atlantic, Amazon, and Benguela) clearly act as ‘filters’ by restricting Evolutionary Biology, University of California, dispersal but at the same time allowing occasional crossings that apparently lead Santa Cruz, CA, 95060, USA to the establishment of new populations and species. Fluctuations in the effectiveness of the filters, combined with ecological differences among provinces, apparently provide a mechanism for much of the recent diversification of reef fishes in the Atlantic. Main conclusions Our data set indicates that both historical events (e.g. Tethys closure) and relatively recent dispersal (with or without further speciation) have had a strong influence on Atlantic tropical marine biodiversity and have contributed to the biogeographical patterns we observe today; however, examples of the latter process outnumber those of the former. *Correspondence: S. R. Floeter, Departamento de Ecologia e Zoologia, Lab. de Biogeografia e Keywords Macroecologia Marinha, Universidade Federal Atlantic Ocean, biodiversity, biogeographical barriers, biogeographical prov- de Santa Catarina, Floriano´polis, SC 88010-970, Brazil. inces, historical biogeography, marine biogeography, phylogeography, specia- E-mail: fl[email protected] tion, Tethys Sea.

22 www.blackwellpublishing.com/jbi ª 2007 The Authors doi:10.1111/j.1365-2699.2007.01790.x Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

‘To some scholars, the search for patterns is a fishing Brazilian and Caribbean reef habitats, and which was estab- expedition, an unworthy exercise… To me, however, it lished c. 11 Ma (Hoorn et al., 1995). The final closure of the provides the critical raw data that enable us to propose and Isthmus of Panama (IOP) c. 3.1 Ma (Coates & Obando, 1996) to constrain possible explanations. Patterns help us to separated the Caribbean from the Tropical Eastern Pacific. identify… critical circumstances of events that could affect Finally, the tropical reef fauna of the Atlantic has been isolated diversity’. (Vermeij, 1996). from the southern Indian Ocean by the cold Benguela Current for at least 2 Myr (the Benguela barrier; Shannon, 1985; Marlow et al., 2000). These geographic and oceanographic features are INTRODUCTION regarded as the primary barriers or filters producing diver- Marine biogeographers have used the geography of endemism gences and speciation among the regional faunas of the Atlantic. to divide the biosphere into major provinces/regions (Ekman, The Atlantic provinces proposed by Briggs (1974) were 1953; Briggs, 1974; Vermeij, 1978), each of which is charac- based on fewer (and less reliable) species distributional data, terized by a number of species found only in limited and he did not have access to modern statistical methods. Here geographical areas, known as areas of endemism. For simplic- we assemble the first comprehensive distribution data base for ity we will use the term ‘province’ to identify major areas of the entire tropical and subtropical Atlantic, including the endemism (Briggs, 1974, 1995). In characterizing biogeo- southwestern Indian Ocean. Recent field and taxonomic graphical provinces, it is necessary to recognize the types of research in understudied areas, such as tropical West Africa barriers or boundaries that separate them (Briggs, 1973, 1974; (e.g. Afonso et al., 1999; Edwards et al., 2003; Wirtz et al., Floeter & Gasparini, 2000; Joyeux et al., 2001; Rocha, 2003). It 2007), the Brazilian coast (e.g. Floeter et al., 2001; Rocha & is also a general aim to understand why and when these Rosa, 2001; Menezes et al., 2003; Dutra et al., 2005), and provinces were formed, and how they relate to each other associated oceanic islands (e.g. Santos et al., 1997; Gasparini & (Vermeij, 2001). In order to achieve these goals we need to Floeter, 2001; Feitoza et al., 2003), revealed new species (e.g. learn more about richness and endemism patterns, diversity Sazima et al., 1998; Rocha & Rosa, 1999; Heiser et al., 2000; buildup (i.e. speciation processes), ecological and palaeoeco- Gasparini et al., 2003), and greatly advanced the knowledge of logical settings, and the operation of biogeographical barriers geographical distributions of tropical reef fishes. Concurrently, or filters. new data were made available on the palaeoceanographic Here we discuss the geographic distribution and evolution of characteristics of the tropical Atlantic (e.g. Stramma & Schott, Atlantic ‘reef fishes’, a group that includes any shallow 1999; Marlow et al., 2000; Peeters et al., 2004). Many mor- (< 100 m) tropical/subtropical benthic or benthopelagic fishes phological and molecular phylogenies of Atlantic reef fishes that consistently associate with hard substrates of coral, algal, or (see Methods) have recently been published and can help to rocky ‘reefs’ or occupy adjacent sand substrate. The distribu- elucidate historical affinities among endemism areas, as well as tion of the great majority of reef fishes is dependent on the the origins and diversification patterns of Atlantic reef fishes. dispersal of eggs and larvae in ocean currents, since these fishes This paper has a number of purposes, namely: (1) to are relatively sedentary during adult stages (Leis, 1991; Victor, summarize current information on diversity and endemism in 1991). Reef fishes are highly diverse; they are not limited to Atlantic reef fishes; (2) to evaluate relationships among the tropical reef areas (like reef-building corals) and thus are ideal main Atlantic biogeographical regions, analysing the effect of for comparisons involving centres of high diversity and the various barriers or filters on these relationships; and (3) to peripheral, lower-diversity areas; they are relatively easy to review published phylogenies (mainly derived from recent identify compared with other groups (e.g. sponges, ascidians); molecular studies) and assess origins and diversification and, finally, a number of reef fish phylogenies (mainly patterns of Atlantic reef fishes by addressing the following molecular) have been published in the last decade, providing questions: (a) what is the geographical pattern of sister taxa insights into speciation patterns within a geographic context. among provinces? (b) are area relationships based on phylo- The tropical Atlantic reef fauna encompasses four primary genetic and distributional patterns concordant? (c) is there a provinces: Brazil, the ‘Greater’ Caribbean, the Mid-Atlantic congruent pattern of divergences resulting from historical Ridge, and the Eastern Atlantic (Table 1; Briggs, 1974). Five events? (d) what is the relative importance of vicariant events major barriers define these regions. The first is the > 3500-km- and dispersal across soft barriers on contemporary biodiversity? wide Atlantic itself, which gradually began to form as a deep ocean barrier (the mid-Atlantic Barrier, MAB) following the METHODS separation of Africa and South America c. 84 Ma (Pittman et al., 1993). The second barrier emerged c. 12–18 Ma with the Atlantic reef fish data base uplift of the Red Sea land bridge, separating the tropical faunas of the Atlantic from the Indian Ocean and closing the Tethys Species lists were compiled for 25 areas of the Atlantic Ocean Seaway in the Terminal Tethyan Event (TTE; Steininger & Ro¨gl, through literature searches of peer-reviewed checklists and 1984; Bellwood & Wainwright, 2002). The third barrier is the published field guides, and from extensive observations by the freshwater outflow from the Orinoco and Amazon rivers, which authors (see Table 1 for area definition and references used). spans 2300 km of the NE coast of South America, separating This data base represents the most exhaustive data set on the

Journal of Biogeography 35, 22–47 23 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Table 1 Deﬁnitions of the studied areas of the Atlantic Ocean and references of checklists used to compile the data set. Regions were based on the results of the maximum-parsimony analysis. Entries in italics signify two or more areas pertaining to the same geographic group.

Province Area Geographical deﬁnition References

Northwestern Atlantic (NWA) Carolinian Carolinian From Cape Kennedy to Smith-Vaniz et al. (1999), North Carolina Quattrini et al. (2004) Greater Caribbean Bermuda Smith-Vaniz et al. (1999)

Western Caribbean Continental area from Cervigo´n (1982), Smith-Vaniz et al. (1999), Florida to Tobago and Smith (1997), Randall (1996), Venezuela (including Schmitter-Soto et al. (2000), Gulf of Mexico) Smith et al. (2002, 2003) Eastern Caribbean Insular area from Bo¨hlke & Chaplin (1993), Bahamas, Cuba to Smith-Vaniz et al. (1999), Dominica and Barbados Claro & Parenti (2001), Smith et al. (2002), Smith-Vaniz et al. (2006)

Southwestern Atlantic (SWA) Brazilian Northeastern Brazil From the Mouth of the Carvalho-Filho (1999), Rocha et al. (1998), Amazon to southern Bahia Feitoza (2001), Rocha & Rosa (2001), Menezes et al. (2003), Feitoza et al. (2005), Dutra et al. (2005), S.R. Rosa, C.R.R. Nunes & L.A. Rocha (unpublished manuscript)

Southeastern Brazil From Espı´rito Santo to Menezes et al. (2003), Floeter et al. (2007), Santa Catarina Hostim-Silva et al. (2006) Atol das Rocas Fernando de Noronha Ridge Rosa & Moura (1997), Menezes et al. (2003), Sampaio et al. (2006) Fernando de Noronha Fernando de Noronha Ridge Menezes et al. (2003), Sampaio et al. (2006) St Paul’s Rocks Lubbock & Edwards (1981), Feitoza et al. (2003), Sampaio et al. (2006) Trindade Is. Gasparini & Floeter (2001), Menezes et al. (2003), Sampaio et al. (2006)

Eastern South America ‘Argentina’ From Uruguay to Menni et al. (1984), Patagonia Cousseau & Perrota (2004)

Mid-Atlantic Ridge (MAR) Ascension Lubbock (1980), D.R. Robertson (personal observation)

St Helena Edwards & Glass (1987a,b), Edwards (1990), D.R. Robertson (personal observation)

Northeastern Atlantic (NEA) Western Mediterranean Gibraltar to the Strait Jennings (2000), Hureau (1998) of Messina

Eastern Mediterranean Levant to the Strait Jennings (2000), Hureau (1998) of Messina Azores Macaronesia Santos et al. (1997) Madeira Macaronesia Que´ro et al. (1990), Wirtz (2001), Wirtz (unpublished data) Canaries Macaronesia Brito et al. (2002)

Tropical Eastern Atlantic (TEA) Northwest Africa Gibraltar to Cape Que´ro et al. (1990), Almada et al. (2001), Verde (Senegal) Falcoń et al. (2002) Cape Verde Is. Reiner (1996), Brito et al. (1999), Menezes et al. (2004) Saõ Tome´ & Prıńcipe Includes Annobon Afonso et al. (1999), Wirtz et al. (2007) Tropical West Africa Cape Verde to Que´ro et al. (1990), Reiner (2001), Moçamedes (Angola) Edwards et al. (2003)

24 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Table 1 continued.

Province Area Geographical deﬁnition References

Southwest Africa (SA) Benguela From Moc¸amedes Smith & Heemstra (1995), Heemstra & (Angola) to the Heemstra (2004), L.E. Beckley, P.A. Hulley, & Cape of Good Hope P.H. Skelton (unpublished manuscript), D. Polack (personal communication)

Agulhas From the Cape of Smith & Heemstra (1995), Good Hope to Heemstra & Heemstra (2004), L.E. Beckley, Kei River P.A. Hulley, & P.H. Skelton (unpublished manuscript), D. Polack (personal communication)

Southwestern Indian South Africa’s From Kei River to Smith & Heemstra (1995), Ocean (SWIO) Indian Ocean Kosi Bay Heemstra & Heemstra (2004), L.E. Beckley, P.A. Hulley & P.H. Skelton (unpublished manuscript), D. Polack (personal communication) distributions of Atlantic reef fishes compiled to date (Table S1, 2002). Geographical areas were characterized by the presence/ Supplementary Material). absence of 2605 reef fish taxa. MP analyses were performed with heuristic searches having starting trees obtained by means of the tree-bisection-reconnection (TBR) branch-swapping Atlantic reef fish phylogenies algorithm. Support for each node from the majority consensus The following phylogenies were used for analysis and infer- tree from the data set was investigated with 1000 bootstrap ences: Abudefduf (Quenouille et al., 2004), Acanthostracion/ pseudo-replicates (Felsenstein, 2004). Lactophrys (Klassen, 1995), Acanthurus (Rocha et al., 2002), To evaluate relationships among areas based on reef fish Alphestes (Craig et al., 2004, 2006), Anisotremus (Bernardi & phylogenies, we calculated the ratio (proportion; given in each Lape, 2005), Aulostomus (Bowen et al., 2001), Centropyge arrow) between the number of nodes in which two areas form a ‘xiphipops’ complex (Bowen et al., 2006a), chaenopsids (Has- sister pair in a given phylogeny (when the terminal clade was a tings, 2000), Chromis (Quenouille et al., 2004; C. chromis clade polytomy, the relationships were counted for all the areas – Domingues et al., 2005; C. ‘multilineata’ complex – L.A. involved, i.e. overlap) and the number of times the same pairs Rocha, D.R. Robertson, A.J. Edwards & B.W. Bowen, in of areas were available for evaluation in the studied phyloge- preparation), Clepticus (Heiser et al., 2000), Dermatolepis nies. The number of studied pairs (possibilities of interactions) (Craig et al., 2004), Diplodus (Summerer et al., 2001), Elac- was around 300 for the seven areas in the studied phylogenies. It atinus (Taylor & Hellberg, 2005), Gnatholepis (Rocha et al., is important to note that not all phylogenies include taxa 2005a), Gobiesocinae (Briggs, 1955), Hippocampus (Casey present in all the areas, and some have more than one pair of et al., 2004), Halichoeres (Rocha, 2004; Barber & Bellwood, sister areas; we thus worked with proportions of the areas in 2005), Holacanthus (Bellwood et al., 2004; G. Bernardi, question (to avoid bias related to unequal coverage of areas). S.R. Floeter & D.R. Robertson, in preparation), Lethrinus In order to summarize the relationships among the areas, (Galbo et al., 2002), Ophioblennius (Muss et al., 2001), cluster analyses (Bray-Curtis dissimilarity index and UPGMA Pomacanthus (Bellwood et al., 2004), Rypticus (Guimara˜es, method) were performed using mvsp software based on: (1) 1999; Carlin et al., 2003), Sparisoma/Nicholsina (Robertson the number of reef fish species (or genera) common to the et al., 2006), Sparidae (Orrell et al., 2002), Strongylura (Ban- pairs of areas, and (2) the matrix of ratios indicating the ford et al., 2004), and Thalassoma (Costagliola et al., 2004). importance of the phylogenetic connections. Varying the When interpreting the phylogenies we assumed that no clustering routine had no effect on the grouping of the sites, extinctions have occurred, and followed estimates (and except when indicated in the figure legend. associated errors) of the timing of speciation events provided and discussed in the original publications. A minimum age of RESULTS AND DISCUSSION 3.1 Ma was assumed for splits related to the closure of the Isthmus of Panama. Geographical distributions

Data analyses Diversity and endemism Maximum-parsimony (MP) analyses of 25 geographical areas In total, our data base covers 1720 teleost reef ﬁsh species from in the Atlantic Ocean were performed using paup* (Swofford, 465 genera and 69 families from the Atlantic, plus another 885

Journal of Biogeography 35, 22–47 25 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Table 2 Species richness, number of endemics and endemism species richness of its northwest counterpart, and a some- (%) of Atlantic areas and South Africa’s Indian Ocean*. what lower, but still substantial, rate of endemism (25%). This endemism rate of the Brazilian coast plus its oceanic Richness Endemics Percentage islands provides the basis for distinguishing the ‘Brazilian Western Atlantic (WA) 1023 881 86.1 Province’, first proposed by Briggs (1974, 1995; see also Northwestern Atlantic (NWA) 831 473 56.9 Floeter & Gasparini, 2000; Joyeux et al., 2001). When only Caribbean 814 272 33.4 the Brazilian coast is considered, the endemism level drops Southwestern Atlantic (SWA) 471 118 25.1 to c. 10% (see also Rocha, 2003). Within coastal Brazil, the Brazilian coast 437 46 10.5 northeast coast has nine endemic species (2.3% of the total Mid-Atlantic Ridge (MAR) 111 29 26.1 richness), and the southeast coast has eight endemics (2.1%). Eastern Atlantic (EA) 551 352 63.9 The SWA contains 117 genera, 78% of the Caribbean Tropical Eastern Atlantic (TEA) 388 117 30.6 number, with only three endemic genera (2.6%), vs. 16% of Southwestern Indian Ocean (SWIO) 1141 – – endemic genera in the Caribbean. *See Table 1 for definitions of the areas. The Mid-Atlantic Ridge (MAR) islands of Ascension and St Helena harbour 111 reef fish species, with 29 (26.1%) endemics, justifying the province proposed by Briggs (1974, 1995). The Eastern Atlantic (EA; from the Mediterranean to the Benguela region) has 551 species (124 genera), with 64% of these species (31.5% of genera) restricted to this side of the Atlantic (Tables 2 and 3). The tropical area of the EA (Cape Verde Islands, Saõ Tome´, and the West African coast from Cape Verde in Senegal to Moçamedes in Angola) includes 388 species (106 genera), and the endemism rate is c. 30% for species (3.8% for genera). Thus, the species and genera richness in the tropical EA (TEA) is considerably lower than that of the tropical Western Atlantic (WA), which is consistent with earlier literature (e.g. Briggs, 1974, 1995; Paulay, 1997). In comparison, the species richness of the tropical Eastern Pacific (TEP) is approximately 89% of the richness of the Caribbean (Robertson, 1998; Fig. 1). The number of species of reef fishes found in tropical South Africa is about 1140, but only 20% of these are also found in the Atlantic Ocean, mainly Figure 1 Reef fish richness patterns in the Atlantic Ocean and in the Agulhas area. Approximately 5% are found in tropical adjacent areas. Circle diameters are sized in proportion to the areas of the Atlantic. number of species in each area. The largest circle inside the The relative impoverishment of the Atlantic fauna compared Atlantic represents 750 species in the Western Caribbean plus with that of the Indo-West Pacific (Paulay, 1997; Bellwood & Florida. Wainwright, 2002) probably reflects the much smaller geographical scale of the region and the historical changes that species from South Africa’s tropical Indian Ocean, resulting in have impacted reefs, especially in the wider Caribbean a total of 2605 species. Encompassing the Greater Caribbean, (Vermeij, 2001, 2005; Johnson et al., 2007). There is no doubt the Northwestern Atlantic (NWA) has the highest number of that the reef biota of this region has been subject to wide-scale species and genera in the Atlantic (831 species – Table 2 and disturbances and episodic extinctions, with the consequence Fig. 1; 150 genera – Table 3, based on selected speciose that the present-day assemblages are less diverse than those of families). This number is dominated by Caribbean species with the Miocene (Budd, 2000). Based on skeletonized groups, it tropical affinities (814). The endemism figures are high for seems that Pliocene extinctions were severe in the Western these areas (species: 57% NWA and 33% Caribbean; genera: Atlantic, where many groups underwent 20 to 30% genus-level 16% for both the Caribbean and the NWA). When considered extinction (Vermeij, 2001, 2005). The closure of the Isthmus separately, the western Caribbean plus Florida (‘Caribbean also modified the oceanographic regime of the western Province’ sensu Briggs, 1974) and the eastern Caribbean (‘West Caribbean, with a change from cooler upwelling to a warmer Indian Province’ sensu Briggs, 1974) have essentially identical oligotrophic regime (Johnson et al., 2007). Moreover, Pleisto- levels of endemism, each with c. 50 (or c. 7%) endemic species. cene glaciations that squeezed the tropical regions and the area This is much lower than the 19% endemism rate of the ‘West of shallow-water habitats (Bellwood & Wainwright, 2002) are Indian Province’ found back in 1974 (Briggs, 1974). also thought to be related to the relative paucity of species in The Southwestern Atlantic (SWA; Brazilian oceanic islands the tropical Atlantic provinces, even when compared with the and Argentina included) has slightly over half (471) of the south-western Indian Ocean (Table 2). However, in the

26 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Table 3 Selected speciose families – total number of genera per family and genera restricted to single regions (in parentheses). NW, New World (TEP + WA); TEP, Tropical Eastern Paciﬁc; WA, Western Atlantic; Braz., Brazil; Carib., Caribbean; EA, Eastern Atlantic; Med, Mediterranean; Cape + SWIO, South African’s Cape + Indian Ocean. Indones., Indonesia (based on Allen & Adrim, 2003) is included for comparison purposes.

Family NW TEP1 WA2 Braz.3 Carib. EA4 TEA Med Cape+IO* Indones.

Apogonidae 3 (1) 1 3 (1) 3 3 2 2 1 12 (11) 14 Balistidae 6644444112(6)10 Batrachoididae 7 (7) 3 (2) 5 (5) 5 (1) 5 (1) 2 2 1 5 (2) 2 Blenniidae 10 (4) 7 (2) 7 (1) 6 7 (1) 13 (3) 10 (1) 9 21 (18) 28 Carangidae 10 (1) 9 10 (1) 10 (1) 8 11 (1) 9 7 9 (1) 14 Chaenopsidae 13 (13) 10 (3) 10 (3) 3 10 (2) – – – – – Dactyloscopidae 8 (8) 6 (1) 7 (2) 4 (1) 6 (1) – – – – – Gobiesocidae 6 (6) 3 6 (6) 5 6 (1) 5 (3) 5 (1) 5 (1) 6 (3) 5 Gobiidae 26 (14) 13 25 (14) 15 25 (8) 19 (16) 14 19 (8) 50 (43) 50 Haemulidae 333333(1)322(1)2 Labridae 8 (2) 7 8 (2) 7 8 (1) 11 (5) 9 8 29 (25) 35 Labrisomidae 7 (3) 6 (2) 5 (1) 4 5 (1) 2 2 – – – Lutjanidae 7 (3) 2 6 (3) 5 6 3 3 – 7 (4) 8 Muraenidae 11 (1) 9 (1) 8 7 8 8 7 4 12 (6) 12 Pomacentridae 7 (3) 7 (3) 4 4 4 5 (1) 5 (1) 1 13 (10) 19 Pseudochromidae ––––––––6(6)11 Scaridae 4 (1) 3 4 (1) 4 4 3 3 1 6 (4) 7 Serranidae 27 (16) 15 27 (15) 15 23 (7) 12 (1) 11 5 25 (15) 21 Sparidae 6 (3) 2 6 (1) 4 5 12 (4) 12 (1) 10 21 (13) – Syngnathidae 10 (3) 5 9 8 9 (1) 8 (3) 4 4 15 (10) 18 Tripterygiidae 4 (3) 4 (3) 1 1 1 1 (1) 1 1 5 (5) 5 No. genera (endemic) 183 (92) 121 (17) 158 (56) 117 (3) 150 (24) 124 (39) 106 (4) 79 (9) 256 (177*) 261 Percent endemic genera 50.3 14.0 35.4 2.6 16.0 31.5 3.8 11.4 69.1* –

1: Eastward-migrant tropical transpacific Pacific genera that occur in the TEP but not the WA or SWIO are not counted as TEP genera. 2: Genera known only from southern Brazil are included in counts of TWA-restricted genera. 3: Including Argentina. 4: Including Macaronesian and Mediterranean (some extending north in the Northeastern Atlantic) genera, except endemic Mediterranean genera. *Numbers in parentheses do not represent strict endemism, but genera not found in the Atlantic and the TEP. Coral reef fishes only. absence of a good fossil record, there is almost no direct speciation. In the Tropics, the intermediate endemism rates of evidence of extinction in reef fishes. 3–6% at Saõ Tome´ and Prıńcipe, Fernando de Noronha Ridge Apart from the high endemism of Ascension and St Helena and Trindade Island probably result from the proximity (Table 4), endemism rates on oceanic islands vary from zero in (< 1000 km) to continental margins, and/or connections to Madeira (although there are two endemic species shared with mainland areas via seamounts. In contrast, the high ende- the Canaries and one with the Azores), to 1.1% in the Azores, mism at the Cape Verde Islands may be related to: (1) greater 1.8% in Bermuda, 1.6% in the Canaries, c. 3% in Saõ Tome´ isolation (from the mainland and between islands), (2) high and Prıńcipe (plus Annobon), 5% in Atol das Rocas plus heterogeneity of habitats (Pe´rez-Ruzafa et al., 2005), and (3) Fernando de Noronha, 5.7% at Trindade Island, 8.3% in the maintenance of warm tropical waters in glacial periods. The Cape Verde Islands, and 9.3% at St Paul’s Rocks. genus Diplodus is thought to have originated in this Various combinations of island age, geographic isolation, archipelago (Summerer et al., 2001), as well as the only area of available habitat and patterns of oceanic currents endemic Atlantic damselfish genus (Similiparma) and the connecting these islands to potential source areas are probably monotypic genus Virididentex (Sparidae). The Cape Verde involved in producing the relatively low endemism in the Islands are also known for high diversity and endemism in non-MAR islands (Table 4; Briggs, 1966; Smith-Vaniz et al., tropical marine invertebrates, including several endemic 1999; Gasparini & Floeter, 2001; Rocha, 2003; Wirtz et al., species in the gastropod genus Conus (Cunha et al., 2005; 2007). The low level of endemism at Bermuda, the Canaries, Duda & Rolań, 2005). The high endemism of St Paul’s Rocks and other high-latitude locations seems to indicate extirpation is related to geographic isolation and overall low species during Pleistocene glacial periods and recent recolonization richness (five of the 54 species are endemics, whereas 17 of (Briggs, 1966; Smith-Vaniz et al., 1999; Rocha, 2004; Domin- the 205 Cape Verde species are endemics). Despite having gues et al., 2006, 2007), with little time for subsequent three species shared only with Ascension and St Helena on the

Journal of Biogeography 35, 22–47 27 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Table 4 Endemism rates of reef ﬁshes and characteristics of the oceanic islands of the Atlantic Ocean.

Distance Isolation Maximum from from distance inside Shelf Diversity Endemism Richness mainland nearest reef the archipelago surface Age Ice age of Island (%) (n species) Latitude (km) habitat (km) (km) (km2) (Ma) effects* habitats

St Helena 13.2 76 16 S 1870 1290 – 156 10 No Low Ascension 11.0 82 8 S 1536 1290à – 106 1.5 No Low St. Paul’s Rocks 9.3 52 1 N 1010 630§ – 1? 9.3 No Low Cape Verde Is. 8.3 207 16 N 560 Mainland 365 13,991 10.3 No High Trindade Is. 5.7 85 20 S 1160 300– 47 32 3.6 No Low F. de Noronha Ridge 5.0 120 4 S 350 Mainland 150 27** 12 No Interm. Sao Tome´ & Prı´ncipe 3.0 221 1 N 250 210 140 860 31 No High Bermuda 1.6 280 32 N 1049 Mainland – 1204 60 Yes High Canaries 1.6 182 28 N 100 Mainland 490 8027 20.5 Yes High Azores 1.1 95 38 N 1300 740àà 675 5363 0.2–7 Yes High Madeira 0.0 133 33 N 630 285§§ 96 810 15 Yes? Interm.

*Only temperature effects. To Ascension. àTo St Helena. §To F. de Noronha. –To seamounts of the Vito´ria-Trindade Chain. **Noronha only. To Fernando Poo (=Bioko). ààTo Madeira. §§To Selvagem Islands.

MAR, all the other components of St Paul’s Rocks are shared absent; and Labrisomidae has only two species). The MAR with the Brazilian coast or the Fernando de Noronha Ridge islands are more similar to the EA than to the WA. Two (Feitoza et al., 2003). Thus, all the Brazilian oceanic islands pantropical families are relatively (percentage of the total are considered here as impoverished outposts of the Brazilian fauna) more speciose in the MAR than elsewhere in the Province (Fig. 4; Edwards & Lubbock, 1983; Floeter & Atlantic, namely Muraenidae and Carangidae. Gasparini, 2000). (3) The southwestern Indian Ocean (SWIO) exhibits a highly distinctive pattern, with several families being more important there than in the Atlantic areas (e.g. Acanthuridae, Composition of the faunas Apogonidae, Clinidae, Labridae, Pomacentridae); several Species richness varies greatly among biogeographical regions, families are unrecorded in the Atlantic (e.g. Caesionidae, not only in terms of total numbers, but also in the relative Nemipteridae, Pseudochromidae, Siganidae); and some fam- contributions from each taxonomic family (Fig. 2). Some ilies that are important in the Atlantic faunas are lacking in features are immediately apparent. the SWIO (Chaenopsidae, Dactyloscopidae, Grammatidae, (1) Overall patterns of species richness per family in the Labrisomidae). NWA and SWA are similar but each family is consistently Some genera have highly skewed distributions (Fig. 3). For more diverse in the NWA, especially families of small example, 156 genera occur in the New World (NW - both sides cryptobenthic fishes. Three of the most speciose families of of tropical America) but not in the EA (e.g. Anisotremus, Neotropical reef fishes have far more species in the NWA than Calamus, Elacatinus, Haemulon, Halichoeres (‘Iridio’ clade), in the SWA (Gobiidae with 91 vs. 27; Labrisomidae with 46 vs. Lythrypnus, Ogcocephalus, and Opistognathus). Other genera, 16; and Chaenopsidae with 51 vs. 4). This is not the case with such as Malacoctenus, Mycteroperca, Sparisoma and Stegastes other species-rich groups that are not restricted to the (Fig. 3), are much more diverse in the WA (and the TEP) than Neotropics: the Labridae, Scorpaenidae, Haemulidae and in the EA. However, Diplodus, Parablennius and Scartella are Carangidae have comparable numbers in both areas, and more speciose in the EA than in the WA. Eighty-four genera Pinguipedidae has more species in the SWA. occurring in the EA (not considering southern Africa) are not (2) The EA areas (TEA and NEA) have very similar patterns found in the WA (e.g. Gobius, Lipophrys, Pagellus, Spicara, but differ greatly from the WA areas. Two speciose families, Symphodus, Thorogobius, Wheelerigobius). Eighteen EA genera Sparidae and Blenniidae, are markedly more speciose in both occur in the Indo-Pacific but not in the WA or TEP (e.g. parts of the EA than in the WA (40 vs. 21 and 43 vs. 24). Branchiostegus, Callionymus, Coris, Lethrinus, Periophthalmus, However, two of the four most speciose families in the WA are Plectorhinchus), indicating a historical connection via southern absent or very poorly represented in the EA (Chaenopsidae is Africa or Tethys relicts.

28 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Figure 2 Species richness, by family, for the ﬁve major Atlantic biogeographical regions and the Southwestern Indian Ocean. NWA: Northwestern Atlantic, SWA: Southwestern Atlantic, MAR: Mid-Atlantic Ridge (Ascension + St Helena), EA: Eastern Atlan- tic, TEA: Tropical Eastern Atlantic, NEA: Northeastern Atlantic, SWIO: South Africa’s tropical Indian Ocean (from Kei River to Kosi Bay). The ordering of families in each region follows the ranking of those families in the NWA. Coloured bars (and bold below) indicate families that are disproportionally more important in one or more speciﬁc regions. Families: 1, Gobiidae; 2, Serranidae; 3, Chaenopsidae; 4, Labrisomidae; 5, Paralichthyidae; 6, Apogonidae; 7, Gobiesocidae; 8, Labridae; 9, Muraenidae; 10, Syngnathidae; 11, Carangidae; 12, Scorpaenidae, 13, Sparidae; 14, Haemulidae; 15, Lutjanidae; 16, Opistognathidae; 17, Pomacen- tridae; 18, Blenniidae; 19, Dactyloscopidae; 20, Batrachoididae, 21, Ophidiidae; 22, Scaridae; 23, Bothidae; 24, Grammatidae; 25, Holocentridae; 26, Ogcocephalidae; 27, Ophichthidae; 28, Mal- acanthidae; 29, Monacanthidae; 30, Synodontidae; 31, Tetra- odontidae; 32, Tripterygiidae; 33, Antennariidae; 34, Bythitidae; 35, Chaetodontidae; 36, Diodontidae; 37, Pomacanthidae; 38, Balistidae; 39, Congridae; 40, Sciaenidae; 41, Callionymidae; 42, Microdesmidae; 43, Ostraciidae; 44, Chlopsidae; 45, Mullidae; 46, Priacanthidae; 47, Acanthuridae; 48, Ptereleotridae; 49, Carapi- dae; 50, Fistulariidae; 51, Kyphosidae; 52, Moringuidae; 53, Pempheridae; 54, Uranoscopidae; 55, Aulostomidae; 56, Cirrhiti- dae; 57, Dactylopteridae; 58, Caesionidae; 59, Caracanthidae; 60, Centracanthidae; 61, Chanidae; 62, Cheilodactylidae; 63, Clinidae; 64, Kuhliidae; 65, Lethrinidae; 66, Myrocongridae; 67, Nemip- teridae; 68, Pentacerotidae; 69, Pinguipedidae; 70, Platycephali- dae; 71, Plesiopidae; 72, Pseudochromidae; 73, Siganidae; 74, Synanceiidae; 75, Trichonotidae; 76, Xenisthmidae; 77, Zanclidae.

(further subdivided into the NEA, TEA, Benguela, MAR and Cape) and the South African Indian Ocean (Fig. 4). Despite the Amazon filter, the SWA and NWA have the greatest number of species and genera in common (348 and 201, respectively, including a total of 55 families; Fig. 5). These species represent 42% of the NWA total reef fish fauna and 74% of the SWA fauna (Fig. 5). If closely related species pairs (n = 30; Table S2) are included in the calculations, their representation increases to 45% and 82%, respectively (Fig. 6). Another way to look at the effectiveness of Atlantic A few genera show disjunct distributions that indicate either biogeographical barriers/filters is to examine the proportional extinction in the WA or colonization of the EA from the IO. For representation of trans-barrier species within each family example, Cirrhitus, Girella and Prionurus are found in the EA, (Figs 6–8). Notably, families Serranidae and Carangidae have Indo-West Pacific and TEP but not in the WA. Tyler (1997) the greatest number of species crossing the Amazon filter. With reported a Caribbean fossil of the putative sister taxon of the the exception of gobiids, the families containing trans-Ama- surgeonfish genus Naso, providing direct evidence for the zonian species are mostly pelagic spawners with medium to extinction scenario (today, no member of the Naso lineage is large body sizes. Demersal-spawning and brooder families with found in the Atlantic). The relationship between these patterns small-bodied cryptobenthic species seem to be the most of diversity and the origin of genera will be discussed later. affected by the Amazon freshwater and sediment discharge (e.g. Chaenopsidae, Gobiidae). Most of these families are disproportionately more diverse on Caribbean coral reefs than Relationships among areas and biogeographical in the SWA (Floeter & Gasparini, 2000; Moura & Sazima, barriers/filters 2003; Rocha, 2003). The second highest exchange is between The maximum-parsimony analysis using the presence/absence the EA and the WA (NWA and SWA areas with c. 106 species species distribution data base illustrates relationships among in each link). In total, we recorded 112 species with trans- the WA (further subdivided into the NWA and SWA), the EA Atlantic distributions (Fig. 7). This corresponds to approxi-

Journal of Biogeography 35, 22–47 29 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Figure 3 Richness patterns of six selected genera in the Atlantic Ocean and adjacent areas. Circle diameters are sized proportionally to the number of species in each area. The maximum number of species is given (in the largest circle) for comparison. mately three times fewer trans-Atlantic species than trans- species are considered, 16 families in the TEA are composed Amazon species (n = 348). Relative to the total fauna, these only of these wide-ranging taxa (Fig. 7). Detailed statistical species comprise 11.8% of the WA fauna (23% of the SWA and analyses involving several biological attributes of reef fishes 13% of the NWA), 20% of the EA, and 45% of the MAR crossing barriers in the Atlantic will be published elsewhere. islands fish fauna. The weakest connection across a soft barrier corresponds to When the TEA is considered separately, the proportion of the Benguela Current filter, represented by 47 species (38 trans-Atlantics rises to 27% of the fauna. If putative sister shared with the TEA – plus 22 putative sister species, and 9 species are included as trans-Atlantic units (species that have shared with the SWA). Relative to the total fauna of each side, their closest relatives on the other side of the Atlantic Ocean; the 60 species and sister species in common comprise 16% of Table S2), the proportion of trans-Atlantic fish taxa in the TEA TEA fishes but only 5% of SWIO. These numbers indicate that increases to 38% of the fauna, a proportion close to that of the the strength of the connection between the Indo-Pacific and MAR. The number of reef fish species common to the MAR the EA has previously been underestimated (Briggs, 1974, (Ascension + St Helena) and the EA (n = 64) or common to 1995). Examples from other taxonomic groups with a good the MAR and the SWA (n = 71) constitute c. 60% of the MAR fossil record reinforcing the relationships between the IO (and/ fish fauna (Fig. 5). or the IWP) and the TEA include gastropods, bivalves, and The majority of trans-Atlantic species belong to families of opisthobranch molluscs (Edmunds, 1977; Oliver and von pelagic-spawners with long pelagic larval durations (PLDs; e.g. Cosel, 1992; Vermeij & Rosenberg, 1993). Muraenidae, Serranidae; Victor, 1991) and/or fishes that are The link between the IO and SWA has been under- known rafters (e.g. Balistidae, Carangidae, Diodontidae, appreciated in previous analyses (but see Vermeij & Snyder, Monacanthidae). Entire families in the MAR and TEA are 2003 for gastropod examples). The nine shared species are few composed of trans-Atlantic species (e.g. Diodontidae, Holo- in comparison to the total number of species in each area, but centridae, Priacanthidae, Synodontidae). If trans-Atlantic sister reveal a possible connection across the South Atlantic. The

30 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Figure 4 Relationships of 25 areas in the Atlantic Ocean based on maximum-parsimony analysis using the presence/absence distribution data base of 2605 reef ﬁsh species. Values at nodes represent bootstrap support (1000 replicates). Consistency index = 0.499, homoplasy index = 0.501 and retention index = 0.620.

II WT

IV ET NWA 13%

42% I III 105 27% 16% 64 58% 15% 71 TEA 348 64% MAR 28% 10% 107 74% 38 SWA 23% 2% 9 V 3% SWIO 0.8% 4% 47

NWA SWA MAR TEA SWIO 0

0.16

0.32

0.48

0.64

0.8

Figure 5 Biogeographical barriers and relationships among areas based on the number of reef ﬁsh species common to each pair of areas. Hard barriers: (I) Isthmus of Panama, 3.1–3.5 Ma (II) Terminal Tethyan Event (Red Sea land bridge), 12–18 Ma. Soft barriers: (III) Amazon discharge, 6–10 Ma; (IV) Mid-Atlantic Barrier, 0–60 Ma; (V) Benguela Barrier, 2 Ma. Areas/provinces: WT, West Tethys; ET, East Tethys. NWA, Northwestern Atlantic (Caribbean + Bermuda + Carolinian); SWA, Southwestern Atlantic (Brazil + Brazilian oceanic islands + Argentina); MAR, Mid-Atlantic Ridge (Ascension + St Helena); EA, Eastern Atlantic (Macaronesia + Gulf of Guinea); SA, South Africa (tropical Indian Ocean). Arrow widths are sized proportionally to the number of species common to the areas. The proportion of the number of species common to a pair of areas relative to its entire reef ﬁsh fauna is given at each end of the arrows. In order to summarize the relationships among the areas, a cluster analysis (Bray-Curtis dissimilarity index; UPGMA method) based on the matrix of the number of shared species among areas indicating the importance of the biogeographical connections is shown at the bottom. Varying the clustering routine has no effect on the grouping of the sites. The link between the SWIO and the Atlantic (47 species) is further divided into two: 38 with the TEP and 9 with the SWA.

Journal of Biogeography 35, 22–47 31 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

80 90 51 50 Chaenopsidae

Gobiidae Caribbean Labrisomidae 40

30 Syngnathidae Apogonidae Gobiesocidae Opstognathidae Blenniidae

20 Pomacentridae Batrachoididae Tripterygiidae 10 Bythitidae Microdesmidae

40 Brazilian Coast Others Number of species Sister-species Brazil + Caribbean 30

0 Mullidae Labridae Scaridae Bothidae Sparidae Gobiidae Balistidae Bythitidae Cirrhitidae Carapidae Congridae Lutjanidae Blenniidae Ophidiidae Serranidae Sciaenidae Chlopsidae Ostraciidae Carangidae Kyphosidae Muraenidae Haemulidae Diodontidae Apogonidae Fistulariidae Moringuidae Ophichthidae Acanthuridae Pempheridae Labrisomidae Tripterygiidae Gobiesocidae Priacanthidae Syngnathidae Synodontidae Holocentridae Scorpaenidae Antennariidae Ptereleotridae Callionymidae Chaenopsidae Malacanthidae Paralichthydae Batrachoididae Pomacentridae Uranoscopidae Tetraodontidae Microdesmidae Monacanthidae Dactylopteridae Pomacanthidae Opstognathidae Chaetodontidae Ogcocephalidae Dactyloscopidae

Figure 6 Distributions of species in families of ﬁshes across the Amazon Barrier: numbers of species found on both sides of the barrier (Brazil + Caribbean); numbers of species that have their closest relative on the other side of the barrier (sisters); and numbers of species found on one side that have no known sisters on the other side of the barrier (other). Demersal-spawning and brooder families are indicated with arrows.

recent record of the starry toadfish Arothron firmamentum in MAR to the EA (Fig. 5), in a similar fashion to the maximum- Argentinian waters (Dıáz de Astarloa et al., 2003) and at St parsimony analysis (Fig. 4). Paul’s Rocks (L. Fritzsche, personal communication) exempli- The number of species shared only between the NWA and fies this route via the South Atlantic gyre. Two recent the SWA (n = 225) is the highest among all pairwise compari- population genetic surveys support the direct recent (post sons; these species are proportionately more dominant in the Benguela-establishment) colonization of the SWA by an IO SWA than in the NWA (48% vs. 27%). Eleven species are found goby (genus Gnatholepis, Rocha et al., 2005a) and angelfish only in the EA and the MAR (e.g. Acanthostracion notacanthus, (genus Centropyge, Bowen et al., 2006a). There are also Gymnothorax unicolor, Myroconger compressus, Xyrichtys sanc- examples of an IO genus found at St Helena and Ascension taehelenae), comprising 10% of the total MAR fauna. but nowhere else in the Atlantic (Helcogramma sp.; Hadley- The SWIO and the TEA share 15 species (plus 11 putative Hansen, 1986; Heemstra & Heemstra, 2004), and there is an sister-species pairs totalling c. 10% of the EA fauna), for assumed invasion of the Atlantic by Chaetodon sedentarius example Gymnothorax margaritophorus and Rhabdosargus from the western IO C. dolosus clade at around 1.6 Ma (Fessler globiceps (both with wider distributions in the Indo-Pacific), & Westneat, 2007). and Lithognathus mormyrus and Sarpa salpa (with a putative A cluster analysis based on the shared species synthesized the EA origin owing to their wider distributions in this area). An overall pattern closely linking the NWA to the SWA, and the additional 52 species with subtropical affinities are distributed

32 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

88 98 59 35 Tropical Western Atlantic 30

0 Tropical Eastern Atlantic 30

25 Others Sister-species 20 Amphiatlantic

Number of species 5

Mid-Atlantic Ridge Figure 7 Distributions of species in families of ﬁshes across the Mid-Atlantic Barrier, 10 and at the Mid-Atlantic Ridge islands (Ascension and St Helena): numbers of species found on both sides of the barrier (Amphiatlantic); numbers of species that 5 have their closest relative on the other side of the barrier or on the Mid-Atlantic Ridge Islands (sisters); and numbers of species found on one side that have no known sisters 0 on the other side of the barrier (other). Mullidae Labridae Bothidae Scaridae Gobiidae Sparidae Balistidae Lutjanidae Congridae Blenniidae Ophidiidae Serranidae Chlopsidae Ostraciidae Carangidae Kyphosidae

Species recorded as vagrants were also Muraenidae Diodontidae Apogonidae Ophichthidae Acanthuridae Labrisomidae Aulostomidae Syngnathidae Synodontidae Priacanthidae Holocentridae Antennariidae Scorpaenidae Callionymidae Pinguipedidae Tetraodontidae Pomacentridae Microdesmidae Monacanthidae Pomacanthidae Dactylopteridae included. Chaetodontidae along the South African coast from the Agulhas to the the TEP, 11.4% in the Mediterranean, 3.8% in the TEA, and Benguela region (Namibia and southern Angola). However, 2.6% in the SWA. these species do not reach tropical waters. A few are also found in the NEA and/or the Mediterranean, thus representing anti- Subtropical and temperate areas tropical distributions (e.g. Sparidae). A cluster analysis sum- marizing the overall pattern of shared-only taxa (not shown) The subtropical and warm-temperate areas included in our closely linked the NWA to the SWA as in Fig. 5, but with the analysis are impoverished outposts of adjacent tropical areas. IO and the EA closely related, and the MAR between the two For example, the Carolinian Province (sensu Smith-Vaniz clusters. et al., 1999) and Argentina (see Irigoyen et al., 2005) clustered A cluster analysis of genera (Fig. 9) revealed a pattern with oceanic islands of the nearest contiguous tropical region similar to that of the species cluster analysis (Fig. 4). At this (Fig. 4). Despite some recognized endemism (although taxo- taxonomic level, however, we were able to include the TEP, nomically restricted to eight Gobiidae genera plus one which clustered with the WA as expected. The TEP and the Gobiesocidae genus; Table 3), the Mediterranean Sea has a NWA have 155 genera in common, the NWA and the SWA marine fauna related to the Macaronesian Islands and the have 144, the TEA and WA have 110, and the TEA and South northeastern Atlantic coast (Gonc¸alves et al., 1993, 2005; Africa’s Indian Ocean have 94. The generic endemism rate is Briggs, 1995; Almada et al., 2001). South Africa’s Benguela (the very high (50.3%) in the New World (TEP + WA) and in the ‘Southwest Africa’ province of Briggs, 1974) and Agulhas EA as a whole (31.5%; Table 3). Within these major regions, provinces grouped with Ascension and St Helena, possibly as a 16% of the Caribbean genera are endemic, followed by 14% in result of their common low diversity rather than of their

Journal of Biogeography 35, 22–47 33 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

121 90 South Africa 75 Indian Ocean

Tropical Eastern Atlantic Others Number of species 30 Sister-species Indian Ocean + TEA

Figure 8 Distributions of species in families 10 of ﬁshes across the Benguela Barrier: numbers of species found on both sides of the barrier (Indian Ocean + Tropical Eastern Atlantic); numbers of species that have their 0 closest relative on the other side of the barrier (sisters); and numbers of species found on Labridae Bothidae Sparidae Gobiidae Balistidae Cirrhitidae Blenniidae Serranidae Sciaenidae Lethrinidae Carangidae Kyphosidae Muraenidae Haemulidae Diodontidae Fistulariidae one side that have no known sisters on the Acanthuridae Gobiesocidae Syngnathidae Synodontidae Antennariidae Scorpaenidae Callionymidae Malacanthidae Paralichthydae Batrachoididae Tetraodontidae Monacanthidae

Centracanthidae other side of the barrier (other). biogeographical affinities (i.e. long-branch attraction; Felsens- What is the geographical pattern of distributions of sister taxa tein, 2004). These areas are a mix of EA and IO species, as among provinces? expected from their geographic locations. Notwithstanding this, there are a number of endemics in southwestern Africa The three strongest sister-species relationships among regions (38 species representing 12% of the fauna), such as the genus (Fig. 11) are: (1) the TEA and MAR, with 86% of the species Clinus with 10 endemic species. being sisters (e.g. Thalassoma sanctaehelenae/ascensionis – newtoni (Fig. 12), Sparisoma strigatum – cretense); (2) NWA and SWA [78%; e.g. Thalassoma bifasciatum – noronhanum, Patterns of historical relationships using phylogenies Sparisoma griseorubra – frondosum, Diplodus argenteus – holbrooki/bermudensis, Halichoeres (Fig. 12), Acanthurus ‘bahi- Origins and diversification of Atlantic reef fishes anus’ clades]; and (3) the TEP and NWA [69%; e.g. Abudefduf Owing to the rarity of reef fish fossils from studied areas (the taurus – concolor, Anisotremus virginicus – taeniatus (Fig. 13)]. Mediterranean being the exception; e.g. Carnevale, 2006), While outside the domain of this study, we also observed historical relationships are recovered from published strong relationships between the IO and WP [75%; e.g. phylogenetic hypotheses (morphological or molecular-based). Hippocampus ‘kuda’ complex, Pomacanthus (Fig. 12)]. Although controversial (cf. Bromham & Penny, 2003; Pulque´- The relatively recent formation of the Isthmus of Panama is rio & Nichols, 2007), molecular phylogenies can provide some reflected in the number of endemic genera (Fig. 9) and sister idea of the age of cladogenic events through the use of taxa shared between the WA and the TEP (e.g. Rosenblatt, ‘molecular clocks’ (Avise, 2000). With a collection of 26 1967; Knowlton & Weigt, 1998; Bernardi & Lape, 2005). Many available phylogenies involving Atlantic reef fish taxa (refer- Caribbean reef fishes are phylogenetically related to the EP ences in Methods), we assess the origins and diversification [69%; e.g. Abudefduf taurus – concolor, Anisotremus virginicus – (i.e. faunal enrichment) patterns of Atlantic reef fishes using taeniatus (Fig. 13)]. However, it is important to note that the evolutionary scenarios shown in Fig. 10. These scenarios Robertson et al. (2004) reviewed the literature and showed that are based on the expected timing and sequence of cladogenic 10 out of 16 EP species (0.63 ratio) are more closely related to events from geological and palaeoceanographic records. Indo-Pacific than to NWA lineages. Thus, the relationship

34 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

between the EP and the NWA (69%) is about the same as its relation to the WP (63%; e.g. Thalassoma lucasanum; Aulo- stomus). The Caribbean reef fishes are more closely related to the TEP (69%) than to the EA [47%; e.g. Hippocampus erectus – hippocampus (Fig. 12)]. The Mid-Atlantic Barrier seems quite permeable, at least in Macaronesia Tropical E Atlantic Mediterranean Tropical E Pacific Caribbean SW Atlantic Carolinian S Africa Benguela S Africa Agulhas SW Indian Ocean terms of the relationship between Brazil and the Gulf of 19 15 Guinea. Brazilian species are sisters to EA species in 67% of the 18 cases [e.g. Clepticus brasiliensis – africanus, Hippocampus reidi – 18 5 algiricus (Fig. 12)]. The percentage of sisters between the 3 20 33 9 0 0 9 6 0 5 203 Caribbean and the EA is lower (47% of cases) than the Brazil- 0 EA link, and the same pattern applies to the MAR (63% Brazil- 0.16 MAR and 44% Caribbean-MAR). It is interesting that 64 WA Abudefduf taurus is found in the Caribbean and in the TEA, 0.32 10 NEA 122 NW but not in Brazil; however, Muraena melanotis is found in St 65 0.48 EA 270 Paul’s Rocks and in the TEA, but not in the Caribbean. Recent Dissimilarity index records of strays of Sargocentron hastatum in the Caribbean 0.64 187 287 (Wilk et al., 2005) and Acanthurus monroviae in Brazil (Luiz 0.80 et al., 2004) indicate that this barrier is breached occasionally. Figure 9 Cluster analysis based on the presence/absence of Relationships between the Atlantic reef fish fauna and that genera in different areas of the Atlantic and Southern Africa and of the Indian Ocean are also strong, 67% of the nodes being the number of endemic genera in each area (Bray-Curtis, UP- sisters in comparisons with the TEA (e.g. Aulostomus chinensis GMA). Horizontal lines show disjunct distributions (or distribu- – strigosus (Fig. 12); Lethrinus atlanticus), and 50% with the tions not related to the cluster analysis) of the genera. Numbers Caribbean, Brazil and the MAR [e.g. Dermatolepis inermis – opposite the horizontal lines represent the total number of genera striolata, Gnatholepis, Centropyge (Fig. 12)]. Craig & Hastings related to each distribution. Line thickness is proportional to (2007) showed that two EA Cephalopholis species (taeniops and the number of genera. Bold numbers represent endemism. Itali- nigri) are closely related to species present in the IO, instead of cized numbers do not represent strict endemism, but genera found in the WA. Other examples of putative sistership between the in the assigned areas plus the IWP. NW: New World; WA: IO and the EA are the Anthiinae Meganthias carpenteri (EA) Western Atlantic; NEA: Northeastern Atlantic; EA: Eastern and M. natalensis (IO) (Anderson, 2006), the butterflyfishes Atlantic.

Figure 10 Evolutionary scenarios for Atlantic reef fishes based on expected timing and sequence of cladogenetic events derived from geological and palaeoceanographic records. Colour coding represents geographical areas: magenta, New World (NW); blue, Eastern Pacific (EP); red, Western Atlantic (WA); green, Caribbean (CA); brown, Brazil (BR); orange, Eastern Atlantic (EA); pink, mid-Atlantic Ridge (MAR); black: Indo-West Pacific (IWP) origin or splits of unknown geography. IP, Indo-Pacific; IO, Indian Ocean. Splits resulting from the Amazon Barrier are also responsible for faunal enrichment within the Atlantic Basin (see scenarios discussed in Rocha, 2003). These splits result in trans-Amazon sister species (see Figs 6, 12 & 14 for examples).

Journal of Biogeography 35, 22–47 35 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

I NWA 0.47 EP 0.69 III WP 0.63* 0.86 0.63 EA 0.78 MAR WP 0.67 0.67 SWA 0.50 V 0.75 IO

NWA SWA MAR EA IO WP 0

0.12

0.24

0.36

0.48

0.6

Figure 11 Biogeographical barriers and relationships among areas based on reef fish phylogenies. Legends of the barriers are the same as in Fig. 1. Areas/Provinces: WT, West Tethys; ET, East Tethys. WP, West Pacific; EP, Eastern Pacific; NWA, Northwestern Atlantic (Caribbean + Bermuda + Carolinian); SWA, Southwestern Atlantic (Brazil + Brazilian oceanic islands + Argentina); MAR, Mid-Atlantic Ridge (Ascension + St Helena); EA, Eastern Atlantic (Mediterranean + Macaronesia + Gulf of Guinea + Benguela); IO, Indian Ocean. Arrow width is sized proportionally to the ratio (proportion; numbers are given in each arrow) between the number of nodes in which two areas form a sister pair in a given phylogeny and the number of times the same pair of areas are available for evaluation in the studied phylogenies (see Methods). In order to summarize the relationships among the areas, a cluster analysis (Bray-Curtis; UPGMA) based on the matrix of ratios indicating the importance of the phylogenetic connections is shown at the bottom. Varying the clustering routine has no effect on the grouping of the sites, except for the EP, which was further excluded from the analysis owing to its inconsistent position in the dendrogram with different routines (sometimes grouping with the Western Atlantic sites and sometimes grouping with the Western Pacific). *: ratio based on the compilation of genetic studies by Robertson et al. (2004). The arrows connecting to the IO represent separate numbers for the EA-IO and WA-IO relationships.

Chaetodon hoefleri (EA) and C. marleyi (IO), and the and thus although sympatric speciation may have occurred it Drepanidae Drepane africana (EA) and D. longimana (IO) seems to be of less importance in marine faunal enrichment. (Heemstra & Heemstra, 2004). All these cases indicate a clear phylogenetic connection of the TEA to (sub)tropical southern Are phylogenetic and distributional patterns concordant with Africa. ‘traditionally’ accepted provinces? Another interesting aspect among sister species is the percentage of putative sympatric (or microallopatric) vs. The overall picture is that phylogenetic and distributional allopatric speciation. From a total of 50 sister pairs in the patterns are concordant. In spite of the Amazon filter (Floeter available phylogenies we found only nine examples (18%) of & Gasparini, 2000; Joyeux et al., 2001; Rocha, 2003), a strong range overlap (i.e. where sympatric speciation might be relationship between the NWA and the SWA exists in terms of inferred). Williams & Reid (2004) reported five examples of both shared species (Fig. 5) and sister species (Figs 6 & 11). It range overlap between sister species among 19 studied pairs in is interesting to note that many ‘Brazilian’ fishes were recently snails of the genus Echinolittorina. Among these cases, we discovered in the southern tip of the Caribbean (e.g. St. could not be sure that sympatric speciation occurred because Vincent, Venezuela and Tobago; see Joyeux et al., 2001; Rocha, they could also reflect secondary contact after allopatric split, 2003; Robertson et al., 2006). New genetic evidence has also

36 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Figure 12 Phylogenies related to Tethys relicts, recent invasions from the Indian Ocean, and diversification within the Atlantic scenarios. Colour coding represents geographic ranges: blue, Eastern Pacific (EP); red, Western Atlantic (WA); green, Carib- bean (CA); brown, Brazil (BR); orange, Eastern Atlantic (EA); pink, mid-Atlantic Ridge (MAR); black, IWP origin or splits of unknown geography. Numbers (or confi- dence intervals) represent the estimated timing of speciation events in Ma as given in the original publications. The minimum age of 3.1 Ma for the splits related to the final closure of the Isthmus of Panama is assumed. Age estimates for Pomacanthidae were cali- brated using fossil evidence (see Bellwood et al., 2004). Coalescence analysis was used to estimate divergence times in Centropyge and Gnatholepis (Rocha et al., 2005a; Bowen et al., 2006a). The complete phylogeny for the genus Sparisoma is given in Fig. 14.

Figure 13 Phylogenies related to the origin in the New World scenario. See Fig. 12 legend for colour codes and inferred timing of speciation events. revealed examples of closer evolutionary links between ‘Bra- The link between the EA and MAR is found both in zilian’ and Venezuelan ﬁshes rather than between Brazil and phylogenetic and in distributional patterns (Figs 5 & 11). The other areas of the NWA [e.g. Sparisoma griseorubra – links between the WA and EA and between the Atlantic basin frondosum (Robertson et al., 2006); samples of Nicholsina usta and Indian Ocean are also concordant. It was not possible to (Brazil) closer to Venezuela than to Florida (Fig. 12)]. assess the relationships between the TEP and the Atlantic by

Journal of Biogeography 35, 22–47 37 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al. species distributions (they share only a few species), but the related to the TEA in overall diversity than to the Macarone- relationships at the genus level (Fig. 9) and sister-species sian Islands or the Mediterranean. In an analysis of the numbers reflect this historical connection (e.g. Rosenblatt, blenniid fauna, Almada et al. (2001) found that Mauritania 1967; Knowlton & Weigt, 1998; Fig. 13). clustered with the TEA, but Morocco grouped with the NEA. A general agreement with the biogeographical provinces A´ vila (2000) found that Moroccan molluscs were affiliated described by Briggs (1974, 1995) was observed in all analyses. with both the Cape Verde Islands and the NEA. In addition, Both the maximum-parsimony (Fig. 4) and the distance there is recent genetic evidence for both endemism in the (Figs 5 & 10) analysis largely agreed with earlier proposed Mediterranean (e.g. Aurelle et al., 2003; Bargelloni et al., 2003; provinces (Briggs, 1974, 1995; Vermeij, 1978). The relationship Table 3) and genetic continuity along the Mediterranean and of the Brazilian oceanic islands with the Brazilian Province also the Atlantic coastline (e.g. Bargelloni et al., 2003; Costagliola supported Briggs (1974, 1995), Lubbock & Edwards (1981), et al., 2004), as well as cases where one of two sister species are Gasparini & Floeter (2001), Rocha (2003) and Feitoza et al. present in the Mediterranean and adjacent Atlantic, while the (2003) in that these islands are impoverished outposts of the other is found in tropical west Africa and the Macaronesian Brazilian Province. However, the Brazilian Province islands (e.g. Chromis chromis – limbata, Domingues et al., should be extended to Santa Catarina at 28 S (Floeter et al., 2005; Parablennius sanguinolentus – parvicornis, Almada et al., 2001). 2005). Briggs (1974) divided the Greater Caribbean into the western Caribbean plus Florida (Caribbean Province) and Is there a general relationship between diversity and origin the eastern Caribbean (West Indian Province). Recent genetic of clades? and biogeographical surveys have demonstrated an important partition at the Mona Passage – between Puerto Rico and The uneven distribution of species in selected genera in Hispaniola (Baums et al., 2005; Dennis et al., 2005; Taylor & different provinces (Fig. 3) seems to be related to their putative Hellberg, 2006), while others found genetic breaks between origin. For example, the genera Anisotremus, Elacatinus, Florida and the Bahamas (Carlin et al., 2003; Rocha et al., Rypticus, Serranus, and Sparisoma all have several species in 2005b). Vermeij (2005), working with fossils, also found a deep the NW (New World, i.e. both sides of tropical America) and split between Florida and the Caribbean for molluscs. There is are now thought to have originated and diversified in the NW thus some justification for partitioning within the NWA (e.g. (Fig. 13; Guimara˜es, 1999; Bernardi & Lape, 2005; Taylor & Gilbert, 1972). However, genetic analyses of wide-ranging fish Hellberg, 2005; Robertson et al., 2006), some with subsequent species in the NWA revealed no significant partitions in most invasion of the EA (Fig. 13; Carlin et al., 2003; Serranus –M. cases (Shulman & Bermingham, 1995; Muss et al., 2001; Rocha Kochzius & A. Dominici-Arosemena, personal communica- et al., 2002; Bowen et al., 2006b). Using the 10% endemism tion). In contrast, the genus Diplodus is more speciose in the level proposed by Briggs (1974) as well as similarity and EA than in the WA, and its molecular phylogeny indicates an parsimony analyses (Figs 4 & 5), the recognition of a ‘Greater EA origin (Fig. 12; Summerer et al., 2001). Distributional data Caribbean’ Province including the eastern and western Carib- indicate that the same pattern should be expected for bean plus Florida and Bermuda is supported (Burgess et al., Parablennius and Scartella. (Both lack molecular phylogenies; 1994; Smith-Vaniz et al., 1999). however, these groups are currently being analysed by As expected, the mid-Atlantic islands of Ascension and V. Almada and collaborators.) St Helena are intermediate in faunal composition between The link between species richness and area of origin is found in the WA and EA, consistent with their intermediate location. the genus Lethrinus, which occurs in the EA and the Indo-Pacific Seventy-one species are shared with the west and 64 with the but not in the WA. The only Atlantic species (L. atlanticus)is east, but, as a result of the greater similarity of their total nested deeply within the Indo-Pacific Lethrinus (Galbo et al., richness with the EA (7.5% of the WA fauna vs. 16% for the 2002), indicating that this genus originated and diversified in the EA), they clustered with the EA in Figs 5 & 11 and with south- Indo-Pacific, and that the Atlantic species is the result of an west Africa and the EA in Fig. 4. A reinforcement of this invasion around 5–6 Ma, assuming the standard divergence of pattern is the stronger phylogenetic relationship of Ascension 2% per million years for cytochrome b (Avise, 2000). Even and St Helena and the EA [Fig. 11; e.g. Ophioblennius and considering a divergence rate of 1% per million years, this Thalassoma (Fig. 13)]. However, genetic analyses of the genera invasion probably occurred via southern Africa well after the Acanthurus and Centropyge in the Atlantic support a close Tethyan Terminal Event (c.12–18 Ma). relationship between the MAR and the SWA (Rocha et al., In summary, it seems that the greatest species richness for 2002; Bowen et al., 2006a). reef fish genera is found at their locations of origin. The division of the EA into three sub-areas (the TEA, the warm-temperate NEA, and southwestern Africa – Fig. 4) was Is there a congruent pattern linked to historical events? Is the predicted by sea-surface-temperature isotherm bunching at timing of divergence consistent among the various taxa? capes along the African coast. The difference between our analysis and that of Briggs (1974, 1995; i.e. ‘Lusitania’ It is clear from the phylogenies that all major biogeographical Province) is that the northwest African coast is more closely barriers/filters left their marks (Figs 12–14). Nevertheless,

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Figure 14 Comparative chronogram of cladogenic events based on seven distinct reef fish phylogenies (references in Methods). Time-scales are based on conventional rates of divergence for fish cytochrome b sequences (1% per Ma at the top and 2.5% per Ma at the bottom). The grey band highlights the diversification that occurred during the late Miocene to the Pliocene (peaking from 8 to 2 Ma if the 2.5% per Ma time-scale is assumed). Colour coding represents geographic ranges as in Figs 12 and 13. speciation events are not simply predicted by geological The timing of cladogenic events is not concordant among or oceanographic events. Both episodic extinction and genera (Figs 12 & 13). For example, trans-Atlantic divergence cladogenic events influence patterns of faunal richness. Owing times range from c. 5.5 Ma (Fig. 13; Thalassoma, Costagliola to the lack of a fossil record for reef fishes, we could et al., 2004; Nicholsina usta clade, Robertson et al., 2006) to not infer extinctions; instead, we could only explore c. 3 Ma (Fig. 13; Sparisoma ‘rubripinne’ clade, Robertson et al., relationships between speciation and large-scale historical 2006; Hippocampus ‘erectus’ clade, Casey et al., 2004), as well as events. £ 1 Ma (Fig. 13; Hippocampus reidi – algiricus, Casey et al.,

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2004) to a few tens of thousands of years (Gnatholepis, Rocha ecological speciation in the genus Halichoeres, and a very et al., 2005a). The relatively recent time frame of partitions similar pattern was observed in gobies of the genus Bathygobius (e.g. Robertson et al., 2006) eliminates vicariance following the (Lima et al., 2005). opening the Atlantic (65 Ma), and there is no suggestion of simultaneous interruptions of gene flow among species What is the relative importance of vicariant persistence and resulting from a single palaeoceanographic event. dispersal routes on contemporary biodiversity? The lack of congruent divergences is also found across the Amazon barrier [Figs 13 & 14: from 6 Ma in Ophioblennius Our data indicate that both vicariant events, such as the macclurei – trindadensis (Muss et al., 2001); to 4 Ma in closure of the Isthmus of Panama, and dispersal events across Thalassoma bifasciatum – noronhanum (Costagliola et al., the three primary Atlantic biogeographical filters have con- 2004); to c. 1Main Acanthurus bahianus (Rocha et al., tributed to the distribution patterns that we observe today. 2002), Halichoeres radiatus – brasiliensis (Rocha, 2004) and Theories concerning the origins of the NW reef biota Sparisoma atomarium – tuiupiranga (Robertson et al., 2006); involve two primary hypotheses: relicts of Tethyan fauna vs. to very recent in Acanthurus coeruleus (Rocha et al., 2002) and colonization from the IWP (Veron, 1995; Barber & Bellwood, Sparisoma viride – amplum (Robertson et al., 2006)]. Likewise, 2005). The relict hypothesis is based on the separation of the the dispersal events from the Indian Ocean to the Atlantic vary western Atlantic from the eastern Indian and Pacific oceans, widely through time (Fig. 13; from 4.6 Ma in the Chromis resulting in two isolated reef systems after the final closure of ‘chromis’ clade to 0.13 Ma in Gnatholepis, Rocha et al., 2005a). the Red Sea land bridge c. 12–18 Ma (Steininger & Ro¨gl, These invasions seem to have occurred both before the 1984). The alternative hypothesis is colonization from the Benguela upwelling c. 2 Ma and relatively recently (Bowen Pacific across the East Pacific Barrier (EPB) and into the et al., 2001, 2006a; Rocha et al., 2005a). Caribbean prior to the closure of the Isthmus of Panama, In summary, the Benguela, mid-Atlantic, and Amazon filters and/or colonization of the Atlantic from the Indian Ocean via have restricted dispersal but also allowed occasional crossings southern Africa. that resulted in the establishment of new populations and These theories yield clear predictions. The relict theory species. Ocean-current dynamics, which are likely to affect predicts that the age of the Atlantic/East Pacific fauna must dispersal across these barriers, have varied considerably with exceed 12 Myr, with origins derived from IO lineages (Fig. 10, changing sea-levels over the past 10 Myr (Kaneps, 1979; scenario A). Colonization via the opened Isthmus predicts that Rocha, 2003; Robertson et al., 2006). The effectiveness of the NW species must be older than 3.1 Myr (Coates & Obando, Amazon barrier, in particular, is thought to have been strongly 1996), and that they will have sister taxa with central and influenced by changing sea-levels not only through variation in eastern Pacific distributions (Fig. 10, scenario B). Recent current speeds (Kaneps, 1979), which affect larval transit times, invasions via South Africa predict that Atlantic species must but also through variations in the salinity of that barrier and be younger than 3.1 Myr and will have Indian Ocean sister the availability of marine habitats beneath the freshwater species (Fig. 10, scenario C1). A vicariant scenario (C2) plume (Rocha, 2003). Fluctuations in the effectiveness of the involving the closure of the Isthmus of Panama and the three filters provide a mechanism for much of the recent intensification of the Benguela upwelling system could involve diversification of reef fishes in the Atlantic. the following steps: (1) an original large geographical range Despite the lack of a consistent temporal signal for encompassing the IO, WA and TEP; (2) isolation of the TEP speciation events, an important diversification occurred during population with the rise of the Isthmus (c. 3.1 Ma); and (3) the late Miocene to the Pliocene (peaking from 8 to 2 Ma; isolation of the WA and the IO by the Benguela Barrier (c. Fig. 14). This phenomenon is probably related to the closing of 2 Ma). the Isthmus of Panama and its effects on the direction and Other possible scenarios for the origins and diversification speed of the Atlantic currents (see Muss et al., 2001 and of the Atlantic reef fauna would be origin in the NW, references therein). A second burst of speciation events probably after W-E Tethys separation deep in the phylo- occurred well after the Isthmus closure (Figs 13 & 14), many genies – i.e. ‘sister-genera’ relationships (Fig. 10, scenario D), of them £ 1.5 Ma and possibly related to fluctuations in the and/or dispersal within the Atlantic basin (Fig. 10, scenarios effectiveness of the three filters or recent diversification across E and F). ecological gradients (Rocha, 2004; Rocha et al., 2005a,b; Taylor From our compilation of phylogenies (Figs 12–14), Atlantic & Hellberg, 2005; Robertson et al., 2006). For example, some reef fishes fit all of the proposed scenarios. Supporting the Caribbean species are present in the Brazilian offshore islands Tethys relict scenario, we have examples of the 18-Ma major of Fernando de Noronha and Atol das Rocas (South of the split in the Halichoeres phylogeny (Figs 12 & 14), and the Amazon barrier) but do not occur on the adjacent Brazilian c. 20-Ma splits in Holacanthus and Pomacanthus, all dividing continental shoreline, which imposes very different ecological the NW and IWP clades. Moreover, there is the morphological conditions (high sediment loads, low water transparence, phylogeny of Scaridae (Bellwood, 1994), where Sparisoma is continental freshwater runoff, etc.). In some cases, these thought to be a Tethyan relict (Streelman et al., 2002). species are replaced by closely related sister taxa along the A similar case is the 20-Ma split of the EA clade Chaetodon coast. This observation led Rocha et al. (2005b) to propose robustus-hoefleri (Chaetodontidae) from IWP clades (Fessler &

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Westneat, 2007). Orrell et al. (2002) also suggested a clear Sea must have affected the evolution of Atlantic reef biota. Tethyan partition within the sparid evolution. Some elements of the Atlantic reef fauna pre-date this event, Supporting colonization via the opened Isthmus hypothesis including corals (Veron, 1995), the endemic lineages of (Fig. 10, scenario B), we have the diversification of a NWA parrotfishes (Bellwood, 1994; Streelman et al., 2002), some clade within Halichoeres (Barber & Bellwood, 2005) and wrasses (Barber & Bellwood, 2005), and angelfishes (Bellwood Thalassoma (Costagliola et al., 2004) (Fig. 14). Bellwood et al., 2004). The three soft barriers or filters have clearly (1994) and Gomon (1997) discuss similar cases for Scarus restricted dispersal and have promoted the development of and Bodianus, respectively. new species in allopatry and/or parapatry. Since habitat In recent years, there has been mounting evidence that late characteristics vary greatly among regions, ecological specia- Pliocene or Pleistocene (< 3 Ma) invasions via southern Africa tion may also have been important in diversification. Fluctu- (Fig. 10, scenario C1) contribute to diversity in the Atlantic. ations in the effectiveness of the soft barriers provide a Examples in Fig. 12 include the genera Aulostomus, Centropyge, mechanism for much of the recent faunal exchanges in the Gnatholepis, and the Hippocampus ‘kuda’ clade (see Casey Atlantic. Thus, vicariant events (e.g. the closure of the Isthmus et al., 2004; Rocha et al., 2005a; Bowen et al., 2006a). The of Panama), relatively recent dispersal (with further speciation evolution of the genus Dermatolepis fits with scenario C2. or not), and adaptation to novel habitats seem to have Relatively recent diversification within the NW (Fig. 10, combined to regulate Atlantic tropical marine biodiversity, all scenario D – with or without ancient separation of clades having contributed to the biogeographical patterns we observe related to Tethys) is found in the Abudefduf ‘declivifrons’ clade, today. Processes originating from dispersal across barriers or Alphestes, Anisotremus, Chaenopsidae, Chromis ‘cyanea’ clade, colonization of novel habitats seem to outnumber vicariant Elacatinus, Gobiescocinae, and Rypticus. It is interesting to events. However, it will be necessary to analyse additional note that the number of geminate pairs separated by the higher-level phylogenies (family and genus levels; e.g. Orrell Isthmus of Panama is relatively low, compared with the total et al., 2002), when they become available, to separate out the number of species in the Neotropical clades. Speciation events relative importance of Tethys relicts vs. later colonization. The within the EP or the WA are much more common than plurality of biogeographical histories and speciation patterns in previously expected, and may correspond to instances of the marine tropics found here agrees with the recent findings sympatric and ecological speciation. of Rocha (2003) for fishes, Meyer (2003) for cowries, and Within the Atlantic basin, the available phylogenies indicate Williams & Reid (2004) for littorinid snails. invasions from the WA to the EA in most cases: Clepticus, Finally, there are undeniable roles for humanity in contem- Hippocampus ‘erectus’ clade, Nicholsina, Ophioblennius, Spari- porary biogeographical processes. These include three obvious soma ‘rubripinne’ clade, and Thalassoma (Fig. 10, scenario E). factors involving the transport and introduction of exotic Invasions from the EA to the WA are rare (e.g. Diplodus). species, and probably several others that elude our grasp. First, However, further studies of other genera with distribution the aquarium trade has introduced Indo-Pacific species into patterns like that of the genus Parablennius (see Fig. 3) may the Atlantic, most notably the Pacific lionfishes in the NWA reveal more cases of westward migrants (see Luiz et al., 2004; (Whitfield et al., 2002). Second, ballast water in cargo vessels is Fig. 10, scenario F). considered a primary threat; however, the screens on ballast In summary, historical relationships among Atlantic prov- tanks mostly exclude fish larvae, and the smaller invertebrate inces are complex and reticulated (sensu Brooks, 2004), with larvae are a much greater hazard (Wonham et al., 2000). provinces comprising a mixture of residents (species that Third, the Suez Canal has essentially reopened the Tethys Sea evolved within province boundaries – after the closure of the connection between the Red Sea and the Mediterranean. The Tethys seaway) and colonizers (species that evolved outside the invasions to date have been for the most part limited by the province and dispersed across a variety of time frames). cooler subtropical conditions in the Mediterranean [but see Moreover, this work provides additional evidence for the Edwards (1987) for examples of some species that have spread importance of founder speciation by means of dispersal in the their distribution ranges since 1902 and Bariche & Saad (2005) evolution of reef fishes (see also Paulay & Meyer, 2002 and for a recent example], but this may change if predictions of Queiroz, 2005). global warming are accurate (see Brito et al., 2005 and Bianchi, 2007 on the recent tropicalization of the littoral ichthyofauna of the Canary Islands and the Mediterranean). Both the Suez Conclusions and future prospects connection and the southern African connection may be Here we have attempted to elucidate the role of biogeograph- enhanced by global warming, and the consequences for inter- ical hard barriers (Tethys and Panama) as well as of more ocean faunal exchanges are uncertain. porous biogeographical filters (Benguela, mid-Atlantic, and Amazon barriers) in generating and maintaining Atlantic ACKNOWLEDGEMENTS biodiversity. Most of the contemporary reef fish families were established by the lower Eocene (50 Ma), and most modern We thank the following people for helping us cross-check genera were present by the mid-Miocene (10 Ma) (Bellwood & data, allowing us to use unpublished data, and/or sharing Wainwright, 2002). As a consequence, the closure of the Tethys their expert knowledge in their study families: L. Beckley,

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P.A. Hulley, P.H. Skelton, D. Polack, J.L. Van Tassell, J. Leis, Barber, P.H. & Bellwood, D.R. (2005) Biodiversity hotspots: R. Fricke, J.E. Randall, W.N. Eschmeyer, G.R. Allen, P. Lobel, evolutionary origins of biodiversity in wrasses (Halichoeres: M. Biscoito, A.G. Floeter, R. Reeves, O.J. Luiz Jr, J.C. Briggs, Labridae) in the Indo-Pacific and New World tropics. R.R. Warner, A. Carvalho-Filho, J. Williams and K. Matsu- Molecular Phylogenetics and Evolution, 35, 235–253. ura. We also thank colleagues who reviewed earlier versions Bargelloni, L., Alarcorn, J.A., Alvarez, M.C., Penzo, E., of the manuscript: G.J. Vermeij, J.C. Briggs, V. Faria, Magoulas, A., Reis, C. & Patarnello, T. (2003) Discord in the V. Almada, V. Domingues and H. Choat. Essential financial family Sparidae (Teleostei): divergent phylogeographical support was provided by the Smithsonian Tropical Research patterns across the Atlantic-Mediterranean divide. Journal of Institute (STV grants to S.R.F. and C.E.L.F.), the National Evolutionary Biology, 16, 1149–1158. Geographic Society (grants 7937-05 to S.R.F. and 7708-04 to Bariche, M. & Saad, M. (2005) Settlement of the lessepsian L.A.R.), and the Universidade Federal de Santa Catarina. This blue-barred parrotfish Scarus ghobban (Teleostei: Scaridae) work was partly conducted while S.R.F. was a postdoctoral in the eastern Mediterranean. Journal of the Marine Biological associate at the National Center for Ecological Analysis and Association UK.2, Biodiversity Records Ref. No. 5049, 1–3. Synthesis, a center funded by the NSF (grant DEB-0072909) Baums, I.B., Miller, M.W. & Hellberg, M.E. (2005) Regionally and the University of California, Santa Barbara. We also isolated populations of an imperiled Caribbean coral, acknowledge the access to many institutional ichthyological Acropora palmata. Molecular Ecology, 14, 1377–1390. collections and the cooperation of their curatorial staffs – Bellwood, D.R. (1994) A phylogenetic study of the parrotfishes without their help the distributional data for this study would family Scaridae (Pisces: Labroidei), with a revision of genera. have been inadequate. Records of the Australian Museum, Supplement, 20, 1–86. Bellwood, D.R. & Wainwright, P.C. (2002) The history and biogeography of fishes on coral reefs. Coral reef fishes. REFERENCES Dynamics and diversity on a complex ecosystem (ed. by P.F. Afonso, P., Porteiro, F.M., Santos, R.S., Barreiros, J.P., Worms, Sale), pp. 5–32. Academic Press, New York, NY. J. & Wirtz, P. (1999) Coastal marine fishes of Saõ Tome Bellwood, D.R., van Herwerden, L. & Konow, N. (2004) Island (Gulf of Guinea). Arquipe´lago Life and Marine Sci- Evolution and biogeography of marine angelfishes (Pisces: ences, 17A, 65–92. Pomacanthidae). Molecular Phylogenetics and Evolution, 33, Allen, G.R. & Adrim, M. (2003) Coral reef fishes of Indonesia. 140–155. Zoological Studies, 42, 1–72. Bernardi, G. & Lape, J. (2005) Tempo and mode of speciation Almada, V.C., Oliveira, R.F., Gonçalves, E.J., Almeida, A.J., in the Baja California disjunct fish species Anisotremus Santos, R.S. & Wirtz, P. (2001) Patterns of diversity of the davidsonii. Molecular Ecology, 14, 4085–4096. north-eastern Atlantic blenniid fish fauna (Pisces: Blennii- Bianchi, C.N. (2007) Biodiversity issues for the forthcoming dae). Global Ecology and Biogeography, 10, 411–422. tropical Mediterranean Sea. Hydrobiologia, 580, 7–21. Almada, F., Almada, V.C., Domingues, V.S., Brito, A. & Santos, Bo¨hlke, J.E. & Chaplin, C.C.G. (1993) Fishes of the Bahamas R.S. (2005) Molecular validation of the specific status of and adjacent tropical waters. University of Texas Press, Parablennius sanguinolentus and Parablennius parvicornis Austin, TX. (Pisces: Blenniidae). Scientia Marina, 69, 519–523. Bowen, B.W., Bass, A.L., Rocha, L.A., Grant, W.S. & Robertson, Anderson, W.D., Jr (2006) Meganthias carpenteri, new species D.R. (2001) Phylogeography of the trumpetfish (Aulo- of fish from the eastern Atlantic Ocean, with a key to eastern stomus): ring species complex on a global scale? Evolution, 55, Atlantic Anthiinae (Perciformes: Serranidae). Proceedings of 1029–1039. the Biological Society of Washington, 119, 404–417. Bowen, B.W., Muss, A.J., Rocha, L.A. & Grant, W.S. (2006a) Aurelle, D., Guillemaud, T., Afonso, P., Morato, T., Wirtz, P., Shallow mtDNA coalescence in Atlantic pygmy angelfish Santos, R.S. & Cancela, M.L. (2003) Genetic study of Coris (genus Centropyge) indicates a recent invasion from the julis (Osteichtyes, Perciformes, Labridae) evolutionary his- Indian Ocean. Journal of Heredity, 97, 1–12. tory and dispersal abilities. Comptes Rendus Biologies, 326, Bowen, B.W., Bass, A.L., Muss, A.J., Carlin, J. & Robertson, 771–785. D.R. (2006b) Phylogeography of two Atlantic squirrelfishes A´ vila, S.P. (2000) Shallow-water marine mollusks of the (family Holocentridae): exploring pelagic larval duration Azores: biogeographical relationships. Arquipe´lago Life and and population connectivity. Marine Biology, 149, 899– Marine Sciences, 2A, 99–131. 913. Avise, J.C. (2000) Phylogeography: the history and formation of Briggs, J.C. (1955) A monograph of the clingfishes (Order species. Harvard University Press, Cambridge, MA. Xenopterygii). Stanford Ichthyological Bulletin, 6, 1–224. Banford, H.M., Bermingham, E. & Collette, B.B. (2004) Briggs, J.C. (1966) Oceanic islands, endemism, and marine Molecular phylogenetics and biogeography of transisth- paleotemperatures. Systematic Zoology, 15, 153–163. mian and amphi-Atlantic needlefishes (Belonidae: Strong- Briggs, J.C. (1973) Operation of zoogeographic barriers. Sys- ylura and Tylosurus): perspectives on New World marine tematic Zoology, 23, 248–256. speciation. Molecular Phylogenetics and Evolution, 31, 833– Briggs, J.C. (1974) Marine zoogeography. McGraw-Hill, New 851. York.

42 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Briggs, J.C. (1995) Global biogeography. Developments in pale- Nacional de Investigacioń y Desarrollo Pesquero, Mar del ontology and stratigraphy, 14, Elsevier, Amsterdam. Plata, Argentina. Brito, A., Herrera, R., Falcoń, J.M., Garcıá-Charton, J.A., Craig, M.T. & Hastings, P.A. (2007) A molecular phylogeny of Barquıń, J. & Pe´rez-Ruzafa, A. (1999) Contribuciońal the groupers of the subfamily Epinephelinae (Serranidae) conocimiento de la ictiofauna de las islas de Cabo Verde. with a revised classification of the Epinephelini. Ichthyolog- Revista de la Academia Canaria de Ciencias, 11, 27–41. ical Research, 54, 1–17. Brito, A., Pascual, P.J., Falcon, J.M., Sancho, A. & Gonzalez, A. Craig, M.T., Hastings, P.A. & Pondella, D.J. (2004) Speciation (2002) Peces de las Islas Canarias. Francisco Lemus Editor, in the Central American Seaway: the importance of taxon Tenerife. sampling in the identification of trans-isthmian geminate Brito, A., Falcoń, J.M. & Herrera, R. (2005) About the recent pairs. Journal of Biogeography, 31, 1085–1091. tropicalisation of the littoral ichthyofauna of the Canary Craig, M.T., Bartsch, P., Wirtz, P. & Heemstra, P.C. (2006) Islands and its relationship with environmental changes and Redescription and validation of Alphestes afer (Bloch 1793) human activities. Vieraea, 33, 515–525. as an amphi-Atlantic grouper species (Perciformes: Serra- Bromham, L. & Penny, D. (2003) The modern molecular nidae). Cybium, 30, 327–331. clock. Nature Genetics, 4, 216–224. Cunha, R.L., Castilho, R., Ru¨ber, L. & Zardoya, R. (2005) Brooks, D.R. (2004) Reticulations in historical biogeography: Patterns of cladogenesis in the venomous marine gastropod the triumph of time over space in evolution. Frontiers in genus Conus from the Cape Verde Islands. Systematic biogeography (ed. by M. Lomolino and L. Heaney), pp. 125– Biology, 54, 634–650. 144. Sinauer Press, Sunderland, MA. Dennis, G.D., Smith-Vaniz, W.F., Colin, P.L., Hensley, D.A. & Budd, A.F. (2000) Diversity and extinction in the Cenozoic McGehee, M.A. (2005) Shore fishes known from islands of history of Caribbean reefs. Coral Reefs, 19, 25–35. the Mona Passage, Greater Antilles with comments on their Burgess, G.H., Smith, S.H. & Lane, E.D. (1994) Fishes of the zoogeography. Caribbean Journal of Science, 41, 716–743. Cayman Islands. The Cayman Islands: natural history and Dıáz de Astarloa, J.M., Figueroa, D.E. & Reta, R. (2003) First biogeography (ed. by M.A. Brunt and J.E. Davies), pp. 199– documented occurrence of the starry toadfish Arothron fir- 228. Kluwer Academic Publishers, Dordrecht. mamentum (Teleostei: Tetraodontidae) in the south-west Carlin, J.L., Robertson, D.R. & Bowen, B.W. (2003) Ancient Atlantic. Journal of the Marine Biological Association of the divergences and recent connections in two tropical Atlantic United Kingdom, 83, 879–880. reef fishes: Epinephelus adscensionis and Rypticus sapona- Domingues, V.S., Bucciarelli, G., Almada, V.C. & Bernardi, G. ceous (Percoidei: Serranidae). Marine Biology, 143, 1057– (2005) Historical colonization and demography of the 1069. Mediterranean damselfish, Chromis chromis. Molecular Carnevale, G. (2006) Morphology and biology of the Miocene Ecology, 14, 4051–4063. butterflyfish Chaetodon ficheuri (Teleostei: Chaetodontidae). Domingues, V.S., Santos, R.S., Brito, A. & Almada, V. (2006) Zoological Journal of the Linnean Society, 146, 251–267. Historical population dynamics and demography of the Carvalho-Filho, A. (1999) Peixes: Costa Brasileira, 3rd edn. eastern Atlantic pomacentrid Chromis limbata (Valenciennes, Melro, Saõ Paulo. 1833). Molecular Phylogenetics and Evolution, 40, 139–147. Casey, S.P., Hall, H.J., Stanley, H.F. & Vincent, A.C.J. (2004) Domingues, V.S., Almada, V.C., Santos, R.S., Brito, A. & The origin and evolution of seahorses (genus Hippocampus): Bernardi, G. (2007) Phylogeography and evolution of the a phylogenetic study using the cytochrome b gene of triplefin Tripterygion delaisi (Pisces, Blennioidei). Marine mitochondrial DNA. Molecular Phylogenetics and Evolution, Biology, 150, 509–519. 30, 261–272. Duda, T.F.J. & Rolań, E. (2005) Explosive radiation of Cape Cervigoń, F. (1982) Los Peces Marinos de Venezuela. Fundacion Verde Conus, a marine species flock. Molecular Ecology, 14, Cientifica Los Roques, Caracas, Venezuela. 267–272. Claro, R. & Parenti, L.R. (2001) The marine ichthyofauna of Dutra, G.F., Allen, G.R., Werner, T. & McKenna, S.A. (eds) Cuba. Ecology of the marine fishes of Cuba (ed. by R. Claro, (2005) A rapid marine biodiversity assessment of the Ab- K.C. Lindeman and L.A. Parenti), pp. 21–57. Smithsonian rolhos Bank, Bahia, Brazil. RAP Bulletin of Biological Institution Press, Washington, DC. Assessment, 38. Conservation International, Washington, Coates, A.G. & Obando, J.A. (1996) The geologic evolution of DC. the Central American isthmus. Evolution and environments in Edmunds, M. (1977) Larval development, oceanic currents, tropical America (ed. by J.B.C. Jackson, A.F. Budd and A.G. and origins of the Opisthobranch fauna of Ghana. Journal of Coates), pp. 21–56. University of Chicago Press, Chicago. Molluscan Studies, 43, 301–308. Costagliola, D., Robertson, D.R., Guidetti, P., Stefanni, S., Edwards, A.J. (1987) Zoogeography of Red Sea fishes. Key Wirtz, P., Heiser, J.B. & Bernardi, G. (2004) Evolution of environments: Red Sea (ed. by A.J. Edwards and S.M. Head), coral reef fish Thalassoma spp. (Labridae). 2. Evolution of pp. 279–286. Pergamon Press, Oxford. the eastern Atlantic species. Marine Biology, 144, 377–383. Edwards, A.J. (1990) Fish and fisheries of Saint Helena Island. Cousseau, M.B. & Perrota, R.G. (2004) Peces Marinos de Centre for Tropical Coastal Management Studies, University Argentina: Biologia, Distribucioń, Pesca. INIDEP. Instituto of Newcastle upon Tyne, UK.

Journal of Biogeography 35, 22–47 43 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Edwards, A.J. & Glass, C.W. (1987a) The fishes of St Helena Labroidae: Scaridae) from Brazil, with comments on the Island, South Atlantic Ocean. I. The shore fishes. Journal of evolution of the genus. Zootaxa, 384, 1–14. Natural History, 21, 617–686. Gilbert, C.R. (1972) Characteristics of the western Atlantic Edwards, A.J. & Glass, C.W. (1987b) The fishes of St Helena reef-fish fauna. Quarterly Journal of Florida Academy of Island, South Atlantic Ocean. II. The pelagic fishes. Journal Sciences, 35, 130–144. of Natural History, 21, 1367–1394. Gomon, M.F. (1997) Relationships of fishes of the labrid tribe Edwards, A.J. & Lubbock, R. (1983) Marine zoogeography of Hypsigenyini. Bulletin of Marine Science, 60, 789–871. St. Paul’s Rocks. Journal of Biogeography, 10, 65–72. Gonçalves, E.J., Almada, V.C., Almeida, A.J. & Oliveira, R.F. Edwards, A.J., Gill, A.C. & Abohweyere, P.O. (2003) A revision (1993) On the occurrence of Parablennius sanguinolentus of F. R. Irvine’s Ghanaian marine fishes in the collections of (Pisces: Blenniidae) on the Portuguese coast. Journal of the The Natural History Museum, London. Journal of Natural Marine Biological Association of the United Kingdom, 73, History, 37, 2213–2267. 465–467. Ekman, S. (1953) Zoogeography of the sea. Sigwick and Jackson, Gonçalves, E.J., Beldade, R. & Henriques, M. (2005) Opea- London. togenys gracilis (Pisces: Gobiesocidae): an overlooked species Falcoń, J.M., Brito, A. & Gonza´lez, G. (2002) Peces de la laguna or another ‘Mediterranean endemism’ found in Atlantic de Khnifiss (Sa´hara, NW de A´ frica) y de los sectores costeros waters? Journal of Fish Biology, 67, 481–489. pro´ximos. Revista de la Academia Canaria de Ciencias, 14, Guimara˜es, R.Z.P. (1999) Revision, phylogeny and comments 139–152. on biogeography of soapfishes of the genus Rypticus (Tele- Feitoza, B.M. (2001) Composiçaõ e estrutura da comunidade de ostei: Serranidae). Bulletin of Marine Science, 65, 337–379. peixes recifais da Risca do Zumbi, Rio Grande do Norte. MSc Hadley-Hansen, P.E. (1986) Revision of the tripterygiid fish Thesis. Universidade Federal da Paraı´ba, Joaõ Pessoa, Brazil. genus Helcogramma, including descriptions of four new Feitoza, B.M., Rocha, L.A., Luiz, O.J., Jr, Floeter, S.R. & species. Bulletin of Marine Science, 38, 313–354. Gasparini, J.L. (2003) Reef fishes of St. Paul’s Rocks: new Hastings, P.A. (2000) Biogeography of the tropical eastern records and notes on biology and zoogeography. Aqua pacific: distribution and phylogeny of chaenopsid fishes. Journal of Ichthyology and Aquatic Biology, 7, 61–82. Zoological Journal of the Linnean Society, 128, 319–335. Feitoza, B.M., Rosa, R.S. & Rocha, L.A. (2005) Ecology and Heemstra, P.C. & Heemstra, E.(2004) Coastal fishes of Southern zoogeography of deep reef fishes in northeastern Brazil. Africa. NISC-SAIAB, Grahamstown, South Africa. Bulletin of Marine Science, 76, 725–742. Heiser, J.B., Moura, R.L. & Robertson, D.R. (2000) Two new Felsenstein, J. (2004) Inferring phylogenies. Sinauer Associates, species of creole wrasse (Labridae: Clepticus) from opposite Sunderland, MA. sides of the Atlantic. Aqua Journal of Ichthyology and Aquatic Fessler, J.L. & Westneat, M.W. (2007) Molecular phylogenetics Biology, 4, 67–76. of the butterflyfishes (Chaetodontidae): Taxonomy and Hoorn, C., Guerrero, J., Sarmiento, G.A. & Lorente, M.A. biogeography of a global coral reef fish family. Molecular (1995) Andean tectonics as a cause for changing drainage Phylogenetics and Evolution, doi:10.1016/j.ympev.2007. patterns in Miocene northern South America. Geology, 23, 05.018 237–240. Floeter, S.R. & Gasparini, J.L. (2000) The southwestern Hostim-Silva, M., Andrade, A.B., Machado, L.F., Gerharding- Atlantic reef fish fauna: composition and zoogeographic er, L.C., Daros, F.A., Barreiros, J.P. & Godoy, E.A.S. (2006) patterns. Journal of Fish Biology, 56, 1099–1114. Peixes de Costaõ Rochoso de Santa Catarina: I. Arvoredo. Floeter, S.R., Guimara˜es, R.Z.P., Rocha, L.A., Ferreira, C.E.L., Editora da Universidade do Vale do Itajaı´, Itajaı´, Brazil. Rangel, C.A. & Gasparini, J.L. (2001) Geographic variation Hureau, J.-C. (1998) Fishes of the North-Eastern Atlantic and in reef-fish assemblages along the Brazilian coast. Global the Mediterranean: Mac Version (World Biodiversity Data- Ecology and Biogeography, 10, 423–433. base CD-ROM Series). UNESCO, Paris. Floeter, S.R., Krohling, W., Gasparini, J.L., Ferreira, C.E.L. & Irigoyen, A.J., Galvań, D.E. & Venerus, L.A. (2005) Occurrence Zalmon, I. (2007) Reef fish community structure on coastal of dusky grouper Epinephelus marginatus (Lowe, 1834) in islands of southeastern Brazil: the influence of exposure gulfs of northern Patagonia, Argentina. Journal of Fish and benthic cover. Environmental Biology of Fishes, 78, 147– Biology, 67, 1741–1745. 160. Jennings, G. (2000) Mediterranean Fishes. A modern taxonomic Galbo, A.M.L., Carpenter, K.E. & Reed, D.L. (2002) Evolution checklist. Calypso Publications, London. http://www.ca- of trophic types in emperor fishes (Lethrinus, Lethrinidae, lypso.org.uk/MedFish2000.pdf. Percoidei) based on cytochrome b gene sequence variation. Johnson, K.G., Todd, J.A. & Jackson, J.B.C. (2007) Coral reef Journal of Molecular Evolution, 54, 754–762. development drives molluscan diversity increase at local and Gasparini, J.L. & Floeter, S.R. (2001) The shore fishes of regional scales in the late Neogene and Quaternary of the Trindade Island, southwestern Atlantic. Journal of Natural southwestern Caribbean. Paleobiology, 33, 24–52. History, 35, 1639–1656. Joyeux, J.C., Floeter, S.R., Ferreira, C.E.L. & Gasparini, J.L. Gasparini, J.L., Floeter, S.R. & Joyeux, J.C. (2003) Sparisoma (2001) Biogeography of tropical reef fish: the South Atlantic tuiupiranga, a new species of parrotfish (Perciformes: puzzle. Journal of Biogeography, 28, 831–841.

44 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Kaneps, A.G. (1979) Gulf Stream: velocity fluctuations during Sparidae (Perciformes: Percoidei) from cytochrome b the late Cenozoic. Science, 204, 297–301. sequences. Copeia, 2002, 618–631. Klassen, G.J. (1995) Phylogeny and biogeography of the Ost- Paulay, G. (1997) Diversity and distribution of reef organisms. raciinae (Tetraodontiformes: Ostraciidae). Bulletin of Mar- Life and death of coral reefs (ed. by C. Birkeland), pp. 298– ine Science, 57, 393–441. 353. Chapman & Hall, New York. Knowlton, N. & Weigt, L.A. (1998) New dates and new rates Paulay, G. & Meyer, C.P. (2002) Diversification in the tropical for divergence across the Isthmus of Panama. Proceeding of Pacific: comparisons between marine and terrestrial systems the Royal Society B: Biological Sciences, 265, 2257–2263. and the importance of founder speciation. Integrative and Leis, J.M. (1991) The pelagic stage of reef fishes: the larval Comparative Biology, 42, 922–934. biology of coral reef fishes. The ecology of fishes on coral reefs Peeters, F.J.C., Acheson, R., Brummer, G.J.A., de Ruijter, (ed. by P.F. Sale), pp. 183–230. Academic Press, San Diego. W.P.M., Schneider, R.R., Ganssen, G.M., Ufkes, E. & Kroon, Lima, D., Freitas, J.E.P., Araujo, M.E. & Sole-Cava, A.M. D. (2004) Vigorous exchange between the Indian and (2005) Genetic detection of cryptic species in the frillfin Atlantic oceans at the end of the past five glacial periods. goby Bathygobius soporator. Journal of Experimental Marine Nature, 430, 661–665. Biology and Ecology, 320, 211–223. Pe´rez-Ruzafa, A., Marcos, C. & Bacallado, J.J. (2005) Bio- Lubbock, R. (1980) The shore fishes of Ascension Island. diversidad marina en archipie´lagos e islas: patrones de Journal of Fish Biology, 17, 283–303. riqueza especı´fica y afinidades faunı´sticas. Vieraea, 33, 455– Lubbock, R. & Edwards, A. (1981) The fishes of Saint Paul’s 475. Rocks. Journal of Fish Biology, 18, 135–157. Pittman, W.C., III, Cande, S., LaBrecque, J. & Pindell, J. (1993) Luiz, O.J., Jr, Floeter, S.R., Gasparini, J.L., Ferreira, C.E.L. & Fragmentation of Gondwana: the separation of Africa from Wirtz, P. (2004) The occurrence of the African surgeonfish South America. Biological relationships between Africa and Acanthurus monroviae (Perciformes: Acanthuridae) in the South America (ed. by P. Goldblatt), pp. 15–34. Yale Uni- southwestern Atlantic, with comments on other eastern versity Press, New Haven. Atlantic reef fishes occurring in Brazil. Journal of Fish Bio- Pulque´rio, M.J.F. & Nichols, R.A. (2007) Dates from the logy, 65, 1173–1179. molecular clock: how wrong can we be? Trends in Ecology Marlow, J.R., Lange, C.B., Wefer, G. & Rosell-Mele, A. (2000) and Evolution, 22, 180–184. Upwelling intensification as part of the Pliocene-Pleistocene Quattrini, A.M., Ross, S.W., Sulak, K.J., Necaise, A.M., Cas- climate transition. Science, 290, 2288–2291. azza, T.L. & Dennis, G.D. (2004) Marine fishes new to Menezes, N.A., Buckup, P.A., Figueiredo, J.L. & Moura, R.L. continental United States waters, North Carolina, and the (eds) (2003) Cata´logo das Espećies de Peixes Marinhos do Gulf of Mexico. Southeastern Naturalist, 3, 155–172. Brasil. Museu de Zoologia da Universidade de Saõ Paulo, Queiroz, A. (2005) The resurrection of oceanic dispersal in Saõ Paulo. historical biogeography. Trends in Ecology and Evolution, 20, Menezes, G.M., Tariche, O., Pinho, M.R., Duarte, P.N., Fer- 68–73. nandes, A. & Aboim, N.A. (2004) Annotated list of fishes Quenouille, B., Bermingham, E. & Planes, S. (2004) Molecular caught by the R/V Arquipe´lago off the Cape Verde Archi- systematics of the damselfishes (Teleostei: Pomacentridae): pelago. Arquipe´lago Life and Marine Sciences, 21 A, 57–71. Bayesian phylogenetic analyses of mitochondrial and nuclear Menni, R.C., Ringuelet, R.A. & Aramburu, R.H. (1984) Peces DNA sequences. Molecular Phylogenetics and Evolution, 31, Marinos de la Argentina y Uruguay. Cata´logo crı´tico ilust- 66–88. rado. Claves para identificacioń de familias, geńeros y Que´ro, J.C., Hureau, J.C., Karrer, C., Post, A. & Saldanha, especies. Nombres vulgares. Glossario. Editorial Hemisferio L.(eds.) (1990) Check-list of the fishes of the eastern tropical Sur, Buenos Aires. Atlantic (CLOFETA), 3 volumes. JNICT, Lisbon; SEI, Paris; Meyer, C.P. (2003) Molecular systematics of cowries (Gastro- UNESCO, Paris. poda: Cypraeidae) and diversification patterns in the tro- Randall, J.E. (1996) Caribbean reef fishes, 3rd edn. TFH, pics. Biological Journal of the Linnean Society, 79, 401–459. Neptune City, NJ. Moura, R.L. & Sazima, I. (2003) Species richness and ende- Reiner, F. (1996) Cata´logo dos Peixes do Arquipe´lago de Cabo mism levels of the Brazilian reef fish fauna. Proceedings of the Verde. Publicaço˜es Avulsas do IPIMAR, 2, Lisboa. 9th International Coral Reef Symposium, 9, 956–959. Reiner, F. (2001) Peixes da Guine´-Bissau. Centro Portugueˆsde Muss, A., Robertson, D.R., Stepien, C.A., Wirtz, P. & Bowen, Estudo dos Mamı´feros Marinhos, Lisboa. B.W. (2001) Phylogeography of Ophioblennius: the role of Robertson, D.R. (1998) Do coral-reef fish faunas have a dis- ocean currents and geography in reef fish evolution. Evo- tinctive taxonomic structure? Coral Reefs, 17, 179–186. lution, 55, 561–572. Robertson, D.R., Grove, J.S. & McCosker, J.E. (2004) Tropical Oliver, P.G. & von Cosel, R. (1992) Taxonomy of tropical west transpacific shore fishes. Pacific Science, 58, 507–565. African Bivalves. IV. Arcidae. Bulletin Museúm National Robertson, D.R., Karg, F., Moura, R.L., Victor, B.C. & Ber- d’Histoire Naturelle Paris, 14, 293–381. nardi, G. (2006) Mechanisms of speciation and faunal Orrell, T.M., Carpenter, K.E., Musick, J.A. & Graves, J.E. enrichment in Atlantic parrotfishes. Molecular Phylogenetics (2002) Phylogenetic and biogeographic analysis of the and Evolution, 40, 795–807.

Journal of Biogeography 35, 22–47 45 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd S. R. Floeter et al.

Rocha, L.A. (2003) Patterns of distribution and processes of Shannon, L.V. (1985) The Benguela ecosystem. Part I. Evolution speciation in Brazilian reef fishes. Journal of Biogeography, of the Benguela physical features and processes. Oceanogra- 30, 1161–1171. phy and Marine Biology Annual Reviews, 23, 105–182. Rocha, L.A. (2004) Mitochondrial DNA and color pattern Shulman, M.J. & Bermingham, E. (1995) Early life histories, variation in three Western Atlantic Halichoeres (Labridae), ocean currents, and population genetics of Caribbean reef with the revalidation of two species. Copeia, 2004, 770– fishes. Evolution 49, 897–910. 782. Smith, C.L. (1997) Tropical marine fishes of the Caribbean, the Rocha, L.A. & Rosa, I.L. (1999) New species of Haemulon Gulf of Mexico, Florida, the Bahamas, and Bermuda. Alfred (Teleostei: Haemulidae) from the NE Brazilian Coast. A. Knopf, Inc., New York. Copeia, 1999, 447–452. Smith, M.M. & Heemstra, P.C. (1995) Smiths’ sea fishes. Rocha, L.A. & Rosa, I.L. (2001) Baseline assessment of reef fish Southern Book Publishers, Johannesburg. assemblages of Parcel Manuel Luiz Marine State Park, Smith, M.L., Carpenter, K.E. & Waller, R.W. (2002) An Maranhaõ, north-east Brazil. Journal of Fish Biology, 58, introduction to the oceanography, geology, biogeography, 985–998. and fisheries of the tropical and subtropical western central Rocha, L.A., Rosa, I.L. & Rosa, R.S. (1998) Peixes recifais da Atlantic. The living marine resources of the western central costa da Paraı´ba, Brasil. Revista brasileira de Zoologia, 15, Atlantic (ed. by K.E. Carpenter), pp. 1–23. FAO species 553–566. identification guide for fishery purposes and American Rocha, L.A., Bass, A.L., Robertson, D.R. & Bowen, B.W. (2002) Society of Ichthyologists and Herpetologists Special Publi- Adult habitat preferences, larval dispersal, and the com- cation No. 5, Vol. 1, FAO, Rome. parative phylogeography of three Atlantic surgeonfishes Smith, C.L., Tyler, J.C., Davis, W.P., Jones, R.S., Smith, D.G. & (Teleostei: Acanthuridae). Molecular Ecology, 11, 243– Baldwin, C.C. (2003) Fishes of the Pelican Cays, Belize. Atoll 252. Research Bulletin, 497. National Museum of Natural History, Rocha, L.A., Robertson, D.R., Rocha, C.R., Van Tassell, J.L., Smithsonian Institution, Washington, DC. Craig, M.T. & Bowen, B.W. (2005a) Recent invasion of the Smith-Vaniz, W.F., Collette, B.B. & Luckhurst, B.E.(1999) tropical Atlantic by an Indo-Pacific coral reef fish. Molecular Fishes of Bermuda: history, zoogeography, annotated checklist, Ecology, 14, 3921–3928. and identification keys. ASIH, Special Publication Number 4, Rocha, L.A., Robertson, D.R., Roman, J. & Bowen, B.W. (2005b) Lawrence, KS. Ecological speciation in tropical reef fishes. Proceedings of the Smith-Vaniz, W.F., Jelks, H.L. & Rocha, L.A. (2006) Relevance Royal Society B: Biological Sciences, 272, 573–579. of cryptic fishes in biodiversity assessments: a case study at Rosa, R.S. & Moura, R.L. (1997) Visual assessment of reef fish Buck Island Reef National Monument, St. Croix. Bulletin of community structure in the Atol das Rocas Biological Marine Science, 79, 17–48. Reserve, off northeastern Brazil. Proceedings of the 8th Steininger, F.F. & Ro¨gl, F. (1984) Paleography and palinspastic International Coral Reef Symposium, 1, 983–986. reconstruction of the Neogene of the Mediterranean and Rosenblatt, R.H. (1967) The zoogeographic relationships of the Paratethys. The geological evolution of the eastern Mediter- marine shore fishes of tropical America. Studies in Tropical ranean (ed. by J.E. Dixon and A.H.F. Robertson), pp. 659– Oceanography, 5, 579–592. 668. Blackwell Scientific Publications, Oxford. Sampaio, C.L.S., Carvalho-Filho, A., Feitoza, B.M., Ferreira, Stramma, L. & Schott, F. (1999) The mean flow field of the C.E.L., Floeter, S.R., Gasparini, J.L., Rocha, L.A. & Sazima, I. tropical Atlantic Ocean. Deep-Sea Research II, 46, 279–303. (2006) Peixes recifais endeˆmicos e ameaçados das Ilhas Streelman, J.T., Alfaro, M., Westneat, M.W., Bellwood, D.R. & Oceaˆnicas Brasileiras e do Complexo Recifal de Abrolhos. Karl, S.A. (2002) Evolutionary history of the parrotfishes: Ilhas Oceaˆnicas Brasileiras – da Pesquisa ao Manejo (ed. by biogeography, ecomorphology, and comparative diversity. R.J.V. Alves and J.W. de Alencar Castro), pp. 217–234. Evolution, 56, 961–971. MMA, Brası´lia. Summerer, M., Hanel, R. & Sturmbauer, C. (2001) Mito- Santos, R.S., Porteiro, F.A. & Barreiros, J.P. (1997) Marine chondrial phylogeny and biogeographic affinities of sea fishes of the Azores. Annotated checklist and bibliography. breans of the genus Diplodus (Sparidae). Journal of Fish Arquipe´lago, Life and Marine Sciences, Bulletin of the Uni- Biology, 59, 1638–1652. versity of the Azores, Supplement 1. Swofford, D.L. (2002) PAUP*: Phylogenetic Analyses using Sazima, I., Gasparini, J.L. & Moura, R.L. (1998) Gramma Parsimony (*and Other Methods), Version 4. Sinauer brasiliensis, a new basslet from the western South Atlantic Associates, Sutherland, MA. (Perciformes: Grammatidae). Aqua Journal of Ichthyology Taylor, M.S. & Hellberg, M.E. (2005) Marine radiations at and Aquatic Biology, 3, 39–43. small geographic scales: speciation in neotropical reef gobies Schmitter-Soto, J.J., Va´squez-Yeomans, L., Aguilar-Perera, A., (Elacatinus). Evolution, 59, 374–385. Curiel-Mondragoń, C. & Caballero-Va´zquez, J.A. (2000) Taylor, M.S. & Hellberg, M.E. (2006) Comparative phylogeo- Lista de peces marinos del Caribe mexicano. Anales del graphy of a genus of coral reef fishes: biogeographical and Instituto de Biologıá Universidad Autońoma de Me´xico. Serie genetical concordance in the Caribbean. Molecular Ecology, Zoologıá, 71, 143–177. 15, 695–707.

46 Journal of Biogeography 35, 22–47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Atlantic reef ﬁsh biogeography and evolution

Tyler, J.C. (1997) The Miocene fish Marosichthys, a putative Wirtz, P. (2001) Madeira, marine life. Francisco Ribeiro & tetraodontiform, actually a perciform surgeon fish (Acan- Filhos LDA, Funchal, Portugal. thuridae) related to recent Naso. Beaufortia, 47, 1–10. Wirtz, P., Ferreira, C.E.L., Floeter, S.R., Fricke, R., Gasparini, Vermeij, G.J. (1978) Biogeography and adaptation: patterns of J.L., Iwamoto, T., Rocha, L.A., Sampaio, C.L. & Schliewen, marine life. Harvard University Press, Cambridge, MA. U. (2007) Coastal fishes of Saõ Tome´ and Prıńcipe – an Vermeij, G.J. (1996) Marine biological diversity: muricid gas- update. Zootaxa, 1523, 1–48. tropods as a case study. Evolutionary paleobiology (ed. by D. Wonham, M.J., Carlton, J.T., Ruiz, G.M. & Smith, L.D. (2000) Jablonski, D.H. Erwin and J.H. Lipps), pp. 355–375. Uni- Fish and ships: relating dispersal frequency and success in versity of Chicago Press, Chicago. biological invasions. Marine Biology, 136, 1111–1121. Vermeij, G.J. (2001) Community assembly in the sea: geological history of the living shore biota. Marine community SUPPLEMENTARY MATERIAL ecology (ed. by M.D. Bertness, S.D. Gaines and M.E. Hay) pp. 39–60. Sinauer Associates, Sunderland, MA. The following supplementary material is available for this Vermeij, G.J. (2005) One-way traffic in the western Atlantic: article online: causes and consequences of Miocene to early Pleistocene Table S1 Atlantic reef fish data base. molluscan invasions in Florida and the Caribbean. Paleobio- Table S2 Sister-species table and references. logy, 31, 624–642. This material is available as part of the online article Vermeij, G.J. & Rosenberg, G. (1993) Giving and receiving – from: http://www.blackwell-synergy.com/doi/abs/10.1111/j. the tropical Atlantic as donor and recipient for invading 1365-2699.2007.01790.x species. American Malacological Bulletin, 10, 181–194. Please note: Blackwell Publishing is not responsible for the Vermeij, G.J. & Snyder, M.A. (2003) The fasciolariid gastropod content or functionality of any supplementary materials genus Benimakia: New species and a discussion of Indo- supplied by the authors. Any queries (other than missing Pacific genera in Brazil. Proceedings of the Academy of Nat- material) should be directed to the corresponding author for ural Sciences of Philadelphia, 153, 15–22. the article. Veron, J.E.N. (1995) Corals in space and time: the biogeography and evolution of the Scleractinia. University of New South Wales Press, Sydney. Victor, B.C. (1991) Settlement strategies and biogeography of BIOSKETCH reef fishes. The ecology of fishes on coral reefs (ed. by P.F. Sale), pp. 231–260. Academic Press, San Diego. Sergio R. Floeter has held a position as professor in Wilk, K., Wilk, K. & Greenfield, D.W. (2005) First record vertebrate zoology at the Universidade Federal de Santa of the Eastern Atlantic Squirelfish Sargocentron hasta- Catarina in southern Brazil since 2006. His main research tum (Holocentridae) from the Western Atlantic Ocean. interests are reef fish macroecology, biogeography, evolution, Proceedings of the California Academy of Sciences, 56, 86–87. and conservation. He also plays the guitar. Williams, S.T. & Reid, D.G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution, 58, 2227–2251. Editor: David Bellwood

Journal of Biogeography 35, 22–47 47 ª 2007 The Authors. Journal compilation ª 2007 Blackwell Publishing Ltd Table S1 Atlantic reef fish database - Floeter et al. A A C I D Y I n A R A R a A C F I N e U O L N A c R A S L G E _ A F O E F K U

T U E A _ N n S C R N E G _ N a A E U N i O P A N T E S A R _ d A R A E S R W A D N A E n R A A N _ E _ E C L B I R L É H R A C I B I E I S S E N L N _ _ _ O A B E I E N I Z W M Z O T B A L E N E D A A A A I I T D B I T I _ V S S I A I O A O R L R U A C C C C D L _ R I U R N E H N I I I I T _ D R R R R E A E D E A E T O O A _ E R M P L E R R R B A O B H P N D C R M R B N G F _ C F F O R O O O I _ _ _ M C N _ _ A A E S Z A A R E O T T A A A E Ã _ _ T R _ R S M S C N A A C S W S S C S A B T N W E E F A S Family Genus Species Authority & Year T ACANTHURIDAE Acanthurus bahianus Castelnau, 1855 1 1 1 1 1 1 1 1 1 1 1 ACANTHURIDAE Acanthurus chirurgus (Bloch, 1787) 1 1 1 1 1 1 1 1 1 1 ACANTHURIDAE Acanthurus coeruleus Bloch & Schneider, 1801 1 1 1 1 1 1 1 1 1 1 ACANTHURIDAE Acanthurus dussumieri Valenciennes, 1835 1 1 ACANTHURIDAE Acanthurus monroviae Steindachner, 1876 1 1 1 1 1 1 1 1 1 ACANTHURIDAE Acanthurus triostegus (Linnaeus, 1758) 1 1 ACANTHURIDAE Naso brevirostris (Cuvier, 1829) 1 1 ACANTHURIDAE Prionurus biafraensis (Blache & Rossignol, 1961) 1 1 ACANTHURIDAE Zebrasoma gemmatum (Valenciennes, 1835) 1 1 ANTENNARIIDAE Antennarius bermudensis Schultz, 1957 1 1 1 ANTENNARIIDAE Antennarius hispidus (Bloch & Schneider, 1801) 1 1 ANTENNARIIDAE Antennarius multiocellatus (Valenciennes, 1837) 1 1 1 1 1 1 1 1 1 1 ANTENNARIIDAE Antennarius nummifer (Cuvier, 1817) 1 1 1 1 1 1 ANTENNARIIDAE Antennarius ocellatus (Bloch & Schneider, 1801) 1 1 1 1 ANTENNARIIDAE Antennarius pardalis (Valenciennes, 1837) 1 1 1 ANTENNARIIDAE Antennarius pauciradiatus Schultz, 1957 1 1 1 1 ANTENNARIIDAE Antennarius radiosus Garman, 1896 1 1 1 1 ANTENNARIIDAE Antennarius senegalensis Cadenat, 1959 1 1 1 1 1 1 ANTENNARIIDAE Antennarius striatus (Shaw, 1794) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ANTENNARIIDAE Histrio histrio (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 APOGONIDAE Apogon affinis (Poey, 1875) 1 1 1 1 1 1 1 APOGONIDAE Apogon americanus Castelnau, 1855 1 1 1 1 1 1 APOGONIDAE Apogon anisolepis Böhlke & Randall, 1968 1 APOGONIDAE Apogon aurolineatus (Mowbray, 1927) 1 1 1 1 APOGONIDAE Apogon axillaris Valenciennes, 1832 1 1 APOGONIDAE Apogon binotatus (Poey, 1867) 1 1 1 APOGONIDAE Apogon gouldi Smith-Vaniz, 1977 1 1 1 1 APOGONIDAE Apogon imberbis (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 APOGONIDAE Apogon kallopterus Bleeker, 1856 1 1 APOGONIDAE Apogon lachneri Böhlke, 1959 1 1 APOGONIDAE Apogon leptocaulus Gilbert, 1972 1 1 APOGONIDAE Apogon maculatus (Poey, 1860) 1 1 1 1 APOGONIDAE Apogon mosavi Dale, 1977 1 1 APOGONIDAE Apogon phenax Böhlke & Randall, 1968 1 1 APOGONIDAE Apogon pillionatus Böhlke & Randall, 1968 1 1 APOGONIDAE Apogon planifrons Longley & Hildebrand, 1940 1 1 1 1 1 1 APOGONIDAE Apogon pseudomaculatus Longley, 1932 1 1 1 1 1 1 1 APOGONIDAE Apogon quadrisquamatus Longley, 1934 1 1 1 1 APOGONIDAE Apogon robbyi Gilbert & Tyler, 1997 1 1 1 1 APOGONIDAE Apogon robinsi Böhlke & Randall, 1968 1 1 1 APOGONIDAE Apogon townsendi (Breder, 1927) 1 1 1 APOGONIDAE Astrapogon alutus (Jordan & Gilbert, 1882) 1 1 1 APOGONIDAE Astrapogon puncticulatus (Poey, 1867) 1 1 1 1 1 1 APOGONIDAE Astrapogon stellatus (Cope, 1867) 1 1 1 1 1 1 1 1 APOGONIDAE Phaeoptyx conklini (Silvester, 1915) 1 1 1 1 APOGONIDAE Phaeoptyx pigmentaria (Poey, 1860) 1 1 1 1 1 1 1 1 1 1 APOGONIDAE Phaeoptyx xenus (Böhlke & Randall, 1968) 1 1 1 AULOSTOMIDAE Aulostomus chinensis (Linnaeus, 1766) 1 1 AULOSTOMIDAE Aulostomus maculatus Valenciennes, 1837 1 1 1 AULOSTOMIDAE Aulostomus strigosus Wheeler, 1955 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Abalistes stellatus (Anonymous, 1798) 1 1 BALISTIDAE Balistes capriscus Gmelin, 1789 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Balistes punctatus Gmelin, 1789 1 1 1 1 1 1 1 BALISTIDAE Balistes vetula Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Canthidermis maculata (Bloch, 1786) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Canthidermis sufflamen (Mitchill, 1815) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Melichthys niger (Bloch, 1786) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 BALISTIDAE Odonus niger (Rüppell, 1836) 1 1 BALISTIDAE Rhinecanthus aculeatus (Linnaeus, 1758) 1 1 BALISTIDAE Rhinecanthus rectangulus (Bloch & Schneider, 1801) 1 1 BALISTIDAE Xanthichthys lineopunctatus (Hollard, 1854) 1 1 BALISTIDAE Xanthichthys ringens (Linnaeus, 1758) 1 1 1 1 1 BALISTIDAE Xenobalistes punctatus Heemstra & Smith, 1983 1 1 BATRACHOIDIDAE Amphichthys cryptocentrus (Valenciennes, 1837) 1 1 BATRACHOIDIDAE Austrobatrachus foedus (Smith, 1947) 1 1 BATRACHOIDIDAE Batrichthys albofasciatus Smith, 1934 1 BATRACHOIDIDAE Batrichthys apiatus (Valenciennes, 1837) 1 1 1 BATRACHOIDIDAE Batrichthys felinus Smith, 1952 1 1 BATRACHOIDIDAE Chatrabus hendersoni (Smith, 1952) 1 BATRACHOIDIDAE Chatrabus melanurus (Barnard, 1927) 1 1 BATRACHOIDIDAE Halobatrachus didactylus (Bloch & Schneider, 1801) 1 1 1 BATRACHOIDIDAE Opsanus beta (Goode & Bean, 1880) 1 1 BATRACHOIDIDAE Opsanus dichrostomus Collette, 2001 1 BATRACHOIDIDAE Opsanus pardus (Goode & Bean, 1880) 1 1 BATRACHOIDIDAE Opsanus phobetron Walters & Robins, 1961 1 1 BATRACHOIDIDAE Opsanus tau (Linnaeus, 1766) 1 BATRACHOIDIDAE Perulibatrachus elminensis (Bleeker, 1863) 1 1 1 BATRACHOIDIDAE Perulibatrachus rossignoli (Roux, 1957) 1 1 BATRACHOIDIDAE Porichthys kymosemeum Gilbert, 1968 1 1 BATRACHOIDIDAE Porichthys oculofrenum Gilbert, 1968 1 1 1 BATRACHOIDIDAE Porichthys pauciradiatus Caldwell & Caldwell, 1963 1 1 BATRACHOIDIDAE Porichthys plectrodon Jordan & Gilbert, 1882 1 1 1 1 BATRACHOIDIDAE Porichthys porosissimus (Cuvier, 1829) 1 1 BATRACHOIDIDAE Sanopus astrifer (Robins & Starck, 1965) 1 BATRACHOIDIDAE Sanopus barbatus (Meek & Hildebrand, 1928) 1 BATRACHOIDIDAE Sanopus greenfieldorum Collette, 1983 1 BATRACHOIDIDAE Sanopus johnsoni Collette & Starck, 1974 1 BATRACHOIDIDAE Sanopus reticulatus Collette, 1983 1 BATRACHOIDIDAE Sanopus splendidus (Berg, 1893) 1 BATRACHOIDIDAE Thalassophryne montevidensis (Berg, 1893) 1 BATRACHOIDIDAE Thalassophryne nattereri Steindachner, 1876 1 1 BATRACHOIDIDAE Thalassophryne punctata Steindachner, 1876 1 1 BATRACHOIDIDAE Triathalassothia argentina (Berg, 1897) 1 BATRACHOIDIDAE Triathalassothia gloverensis Greenfield & Greenfield, 1973 1 BATRACHOIDIDAE Triathalassothia lambaloti Menezes & de Figueiredo, 1998 1 BLENNIIDAE Aidablennius sphynx (Valenciennes, 1836) 1 1 1 BLENNIIDAE Antennablennius australis Fraser-Brunner, 1951 1 1 BLENNIIDAE Antennablennius bifilum (Günther, 1861) 1 1 BLENNIIDAE Aspidontus dussumieri (Valenciennes, 1836) 1 1 BLENNIIDAE Bathyblennius antholops (Springer & Smith-Vaniz, 1970) 1 BLENNIIDAE Blennius normani Poll, 1949 1 1 BLENNIIDAE Blennius ocellaris Linnaeus, 1758 1 1 1 1 BLENNIIDAE Chalaroderma capito (Valenciennes, 1836) 1 1 BLENNIIDAE Chalaroderma ocellata (Gilchrist & Thompson, 1908) 1 BLENNIIDAE Chasmodes bosquianus (Lacepède, 1800) 1 BLENNIIDAE Chasmodes longimaxilla Williams, 1983 1 BLENNIIDAE Chasmodes saburrae Jordan & Gilbert, 1882 1 BLENNIIDAE Coryphoblennius galerita (Linnaeus, 1758) 1 1 1 1 1 1 BLENNIIDAE Entomacrodus cadenati Springer, 1967 1 1 1 1 BLENNIIDAE Entomacrodus nigricans Gill, 1859 1 1 1 1 1 BLENNIIDAE Entomacrodus sp.n. 1 BLENNIIDAE Entomacrodus textilis (Valenciennes, 1836) 1 1 BLENNIIDAE Entomacrodus vomerinus (Valenciennes, 1836) 1 1 1 1 BLENNIIDAE Hypleurochilus aequipinnis (Günther, 1861) 1 1 1 BLENNIIDAE Hypleurochilus bananensis (Poll, 1959) 1 1 1 1 1 BLENNIIDAE Hypleurochilus bermudensis Beebe & Tee-Van, 1933 1 1 1 1 BLENNIIDAE Hypleurochilus caudovittatus Bath, 1994 1 BLENNIIDAE Hypleurochilus fissicornis (Quoy & Gaimard, 1824) 1 1 1 BLENNIIDAE Hypleurochilus geminatus (Wood, 1825) 1 1 1 BLENNIIDAE Hypleurochilus langi (Fowler, 1923) 1 1 BLENNIIDAE Hypleurochilus multifilis (Girard, 1858) 1 BLENNIIDAE Hypleurochilus pseudoaequipinnis Bath, 1994 1 1 1 1 1 1 BLENNIIDAE Hypleurochilus springeri Randall, 1966 1 1 BLENNIIDAE Hypsoblennius exstochilus Böhlke, 1959 1 BLENNIIDAE Hypsoblennius hentz (Lesueur, 1825) 1 1 BLENNIIDAE Hypsoblennius invemar Smith-Vaniz & Acero P., 1980 1 1 1 1 BLENNIIDAE Hypsoblennius ionthas (Jordan & Gilbert, 1882) 1 1 BLENNIIDAE Istiblennius edentulus (Forster & Schneider, 1801) 1 1 BLENNIIDAE Lipophrys trigloides (Valenciennes, 1836) 1 1 1 1 1 1 1 BLENNIIDAE Microlipophrys adriaticus (Steindachner & Kolombatovic, 1883) 1 BLENNIIDAE Microlipophrys basiliscus (Valenciennes, 1836) 1 1 BLENNIIDAE Microlipophrys bauchotae (Wirtz & Bath, 1982) 1 BLENNIIDAE Microlipophrys caboverdensis (Wirtz & Bath, 1989) 1 BLENNIIDAE Microlipophrys canevae (Vinciguerra, 1880) 1 1 BLENNIIDAE Microlipophrys dalmatinus (Steindachner & Kolombatovic, 1883) 1 1 BLENNIIDAE Microlipophrys nigriceps (Vinciguerra, 1883) 1 1 BLENNIIDAE Microlipophrys pholis (Linnaeus, 1758) 1 1 1 1 1 BLENNIIDAE Microlipophrys velifer (Norman, 1935) 1 1 1 1 BLENNIIDAE Omobranchus woodi (Gilchrist & Thompson, 1908) 1 1 BLENNIIDAE Ophioblennius atlanticus (Valenciennes, 1836) 1 1 1 1 BLENNIIDAE Ophioblennius macclurei (Silvester, 1915) 1 1 1 1 BLENNIIDAE Ophioblennius sp. 1 1 BLENNIIDAE Ophioblennius sp.EA 1 1 1 BLENNIIDAE Ophioblennius trinitatis Miranda-Ribeiro, 1919 1 1 1 1 1 1 BLENNIIDAE Parablennius cornutus (Linnaeus, 1758) 1 1 1 1 BLENNIIDAE Parablennius dialloi Bath, 1990 1 BLENNIIDAE Parablennius gattorugine (Linnaeus, 1758) 1 1 1 BLENNIIDAE Parablennius goreensis (Valenciennes, 1836) 1 1 BLENNIIDAE Parablennius incognitus (Bath, 1968) 1 1 1 1 1 1 1 BLENNIIDAE Parablennius marmoreus (Poey, 1876) 1 1 1 1 1 1 BLENNIIDAE Parablennius parvicornis (Valenciennes, 1836) 1 1 1 1 1 1 BLENNIIDAE Parablennius pilicornis (Cuvier, 1829) 1 1 1 1 1 1 1 1 1 BLENNIIDAE Parablennius rouxi (Cocco, 1833) 1 1 BLENNIIDAE Parablennius ruber (Valenciennes, 1836) 1 1 BLENNIIDAE Parablennius salensis Bath, 1990 1 BLENNIIDAE Parablennius sanguinolentus (Pallas, 1814) 1 1 1 BLENNIIDAE Parablennius sierraensis Bath, 1990 1 BLENNIIDAE Parablennius tentacularis (Brünnich, 1768) 1 1 BLENNIIDAE Parablennius verryckeni (Poll, 1959) 1 1 BLENNIIDAE Parablennius zvonimiri (Kolombatovic, 1892) 1 1 BLENNIIDAE Plagiotremus rhinorhynchos (Bleeker, 1852) 1 1 BLENNIIDAE Plagiotremus tapeinosoma (Bleeker, 1857) 1 1 BLENNIIDAE Salaria basilisca (Valenciennes, 1836) 1 1 BLENNIIDAE Salaria pavo (Risso, 1810) 1 1 1 1 BLENNIIDAE Scartella caboverdiana Bath, 1990 1 BLENNIIDAE Scartella cristata (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 BLENNIIDAE Scartella emarginata (Günther, 1861) 1 1 1 1 BLENNIIDAE Scartella nuchifilis (Valenciennes, 1836) 1 BLENNIIDAE Scartella poiti Rangel, Gasparini & Guimarães, 2004 1 BLENNIIDAE Scartella sp.n. 1 1 BLENNIIDAE Scartella springeri (Bauchot, 1967) 1 BLENNIIDAE Spaniblennius clandestinus Bath & Wirtz, 1989 1 BLENNIIDAE Spaniblennius riodourensis (Metzelaar, 1919) 1 BLENNIIDAE Xiphasia matsubarai Okada & Suzuki, 1952 1 1 BLENNIIDAE Xiphasia setifer Swainson, 1839 1 1 BOTHIDAE Arnoglossus capensis Boulenger, 1898 1 1 1 1 1 1 1 1 BOTHIDAE Arnoglossus imperialis (Rafinesque, 1810) 1 1 1 1 1 1 1 BOTHIDAE Arnoglossus laterna (Walbaum, 1792) 1 1 1 BOTHIDAE Arnoglossus thori Kyle, 1913 1 1 1 1 1 1 BOTHIDAE Bothus guibei Stauch, 1966 1 1 BOTHIDAE Bothus lunatus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 BOTHIDAE Bothus maculiferus (Poey, 1860) 1 1 1 1 1 1 BOTHIDAE Bothus mellissi Norman, 1931 1 1 BOTHIDAE Bothus ocellatus (Agassiz, 1831) 1 1 1 1 1 1 1 BOTHIDAE Bothus pantherinus (Rüppell, 1830) 1 1 BOTHIDAE Bothus podas_maderensis (Lowe, 1834) 1 1 1 1 BOTHIDAE Bothus podas_podas (Delaroche, 1809) 1 1 1 1 1 BOTHIDAE Bothus robinsi Topp & Hoff, 1972 1 1 1 1 1 1 BOTHIDAE Chascanopsetta lugubris Alcock, 1894 1 1 1 1 1 1 1 1 BOTHIDAE Chascanopsetta valderostratus (Alcock, 1890) 1 1 BOTHIDAE Engyophrys senta Ginsburg, 1933 1 1 1 1 BOTHIDAE Monolene antillarum Norman, 1933 1 1 1 1 1 1 BOTHIDAE Monolene atrimana Goode & Bean, 1886 1 1 1 BOTHIDAE Monolene megalepis Woods, 1961 1 1 BOTHIDAE Monolene mertensi (Poll, 1959) 1 1 BOTHIDAE Monolene microstoma Cadenat, 1937 1 BOTHIDAE Monolene sessilicauda Goode, 1880 1 1 BOTHIDAE Pseudorhombus arsius (Hamilton, 1822) 1 1 BOTHIDAE Pseudorhombus elevatus Ogilby, 1912 1 1 BOTHIDAE Trichopsetta caribbaea Anderson & Gutherz, 1967 1 1 BOTHIDAE Trichopsetta melasma Anderson & Gutherz, 1967 1 1 BOTHIDAE Trichopsetta orbisulcus Anderson & Gutherz, 1967 1 1 BOTHIDAE Trichopsetta ventralis (Goode & Bean, 1885) 1 1 BYTHITIDAE Bidenichthys capensis Barnard, 1934 1 1 BYTHITIDAE Calamopteryx goslinei Böhlke & Cohen, 1966 1 1 BYTHITIDAE Calamopteryx robinsorum Cohen, 1973 1 1 BYTHITIDAE Dermatopsoides kasougae (Smith, 1943) 1 BYTHITIDAE Dermatopsoides talboti Cohen, 1966 1 1 BYTHITIDAE Dinematichthys minyomma Sedor & Cohen, 1987 1 1 BYTHITIDAE Grammonoides opisthodon (Smith, 1934) 1 BYTHITIDAE Grammonus ater (Risso, 1810) 1 1 BYTHITIDAE Grammonus claudei (de la Torre y Huerta, 1930) 1 1 1 BYTHITIDAE Grammonus longhursti (Cohen, 1964) 1 1 1 BYTHITIDAE Ogilbia boehlkei Møller, Schwarzhans & Nielsen, 2005 1 1 BYTHITIDAE Ogilbia cayorum Møller, Schwarzhans & Nielsen, 20105 1 BYTHITIDAE Ogilbia jeffwilliamsi Møller, Schwarzhans & Nielsen, 2005 1 1 BYTHITIDAE Ogilbia mccoskeri Møller, Schwarzhans & Nielsen, 2005 1 BYTHITIDAE Ogilbia sabaji Møller, Schwarzhans & Nielsen, 2005 1 1 BYTHITIDAE Ogilbia sp.1(Abrolhos) 1 BYTHITIDAE Ogilbia sp.2 (Panama, Torro Pt.) 1 BYTHITIDAE Ogilbia suarezae Møller, Schwarzhans & Nielsen, 2005 1 1 BYTHITIDAE Ogilbia tyleri Møller, Schwarzhans & Nielsen, 2005 1 BYTHITIDAE Ogilbichthys ferocis Møller, Schwarzhans & Nielsen, 2004 1 BYTHITIDAE Ogilbichthys haitiensis Møller, Schwarzhans & Nielsen, 2004 1 BYTHITIDAE Ogilbichthys kakuki Møller, Schwarzhans & Nielsen, 2004 1 1 BYTHITIDAE Ogilbichthys longimanus Møller, Schwarzhans & Nielsen, 2004 1 1 BYTHITIDAE Ogilbichthys microphthalmus Møller, Schwarzhans & Nielsen, 2004 1 1 BYTHITIDAE Ogilbichthys puertoricoensis Møller, Schwarzhans & Nielsen, 2004 1 BYTHITIDAE Ogilbichthys tobagoensis Møller, Schwarzhans & Nielsen, 2004 1 BYTHITIDAE Petrotyx sanguineus (Meek & Hildebrand, 1928) 1 1 BYTHITIDAE Pseudogilbia sanblasensis Møller, Schwarzhans & Nielsen, 2004 1 BYTHITIDAE Stygnobrotula latebricola Böhlke, 1957 1 1 1 1 CAESIONIDAE Caesio caerulaurea Lacepède, 1801 1 1 CALLIONYMIDAE Callionymus lyra Linnaeus, 1758 1 1 1 CALLIONYMIDAE Callionymus maculatus Rafinesque, 1810 1 CALLIONYMIDAE Callionymus marleyi Regan, 1919 1 1 CALLIONYMIDAE Callionymus sanctaehelenae Fricke, 1983 1 CALLIONYMIDAE Diplogrammus pauciradiatus (Gill, 1865) 1 1 1 1 CALLIONYMIDAE Draculo celetus (Smith, 1963) 1 1 CALLIONYMIDAE Draculo shango (Davis & Robins, 1966) 1 CALLIONYMIDAE Foetorepus agassizii (Goode & Bean, 1888) 1 1 1 CALLIONYMIDAE Foetorepus dagmarae (Fricke, 1985) 1 1 CALLIONYMIDAE Foetorepus goodenbeani Nakabo & Hartel, 1999 1 1 CALLIONYMIDAE Paracallionymus costatus (Boulenger, 1898) 1 1 1 CALLIONYMIDAE Paradiplogrammus bairdi (Jordan, 1888) 1 1 1 1 1 1 1 1 1 CALLIONYMIDAE Synchiropus monacanthus Smith, 1935 1 1 CALLIONYMIDAE Synchiropus phaeton (Günther, 1861) 1 1 1 1 1 1 CARANGIDAE Alectis alexandrina (Geoffroy Saint-Hilaire, 1817) 1 1 1 1 1 CARANGIDAE Alectis ciliaris (Bloch, 1787) 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Campogramma glaycos (Lacepède, 1801) 1 1 1 1 CARANGIDAE Carangoides bartholomaei (Cuvier, 1833) 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Carangoides chrysophrys (Cuvier, 1833) 1 1 CARANGIDAE Carangoides equula (Temminck & Schlegel, 1844) 1 1 CARANGIDAE Carangoides ferdau (Forsskål, 1775) 1 1 CARANGIDAE Carangoides gymnostethus (Cuvier, 1833) 1 1 CARANGIDAE Carangoides ruber (Bloch, 1793) 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Caranx crysos (Mitchill, 1815) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Caranx hippos (Linnaeus, 1766) 1 1 1 1 1 1 1 1 1 CARANGIDAE Caranx latus Agassiz, 1831 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Caranx lugubris Poey, 1860 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Caranx senegallus Cuvier, 1833 1 1 1 CARANGIDAE Caranx fischeri Smith-Vaniz & Carpenter, 2007 1 1 1 1 1 CARANGIDAE Chloroscombrus chrysurus (Linnaeus, 1766) 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Hemicaranx amblyrhynchus (Cuvier, 1833) 1 1 1 1 CARANGIDAE Hemicaranx bicolor (Günther, 1860) 1 CARANGIDAE Lichia amia (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Oligoplites palometa (Cuvier, 1832) 1 1 1 CARANGIDAE Oligoplites saliens (Bloch, 1793) 1 1 1 1 CARANGIDAE Oligoplites saurus saurus (Bloch & Schneider, 1801) 1 1 1 1 1 CARANGIDAE Parona signata (Jenyns, 1841) 1 1 CARANGIDAE Pseudocaranx dentex (Bloch & Schneider, 1801) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Scomberoides commersonnianus Lacepède, 1801 1 1 CARANGIDAE Selene brownii (Cuvier, 1816) 1 1 1 1 CARANGIDAE Selene dorsalis (Gill, 1863) 1 1 CARANGIDAE Selene vomer (Linnaeus, 1758) 1 1 1 1 1 1 CARANGIDAE Selene setapinnis (Mitchill, 1815) 1 1 1 1 1 1 CARANGIDAE Seriola carpenteri Mather, 1971 1 1 1 1 1 1 1 CARANGIDAE Seriola dumerili (Risso, 1810) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Seriola fasciata (Bloch, 1793) 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Seriola lalandi Valenciennes, 1833 1 1 1 1 1 1 1 CARANGIDAE Seriola rivoliana Valenciennes, 1833 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Seriola zonata (Mitchill, 1815) 1 1 1 1 CARANGIDAE Trachinotus africanus Smith, 1967 1 1 CARANGIDAE Trachinotus botla (Shaw, 1803) 1 1 CARANGIDAE Trachinotus carolinus (Linnaeus, 1766) 1 1 1 1 1 CARANGIDAE Trachinotus cayennensis Cuvier, 1832 1 1 CARANGIDAE Trachinotus falcatus (Linnaeus, 1758) 1 1 1 1 1 1 1 CARANGIDAE Trachinotus goodei Jordan & Evermann, 1896 1 1 1 1 1 1 1 CARANGIDAE Trachinotus goreensis Cuvier, 1832 1 1 1 1 CARANGIDAE Trachinotus marginatus Cuvier, 1832 1 1 CARANGIDAE Trachinotus maxillosus Cuvier, 1832 1 1 1 1 CARANGIDAE Trachinotus ovatus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 CARANGIDAE Trachinotus teraia Cuvier, 1832 1 1 CARAPIDAE Carapus acus (Brünnich, 1768) 1 1 1 1 1 CARAPIDAE Carapus bermudensis (Jones, 1874) 1 1 1 1 1 1 CARAPIDAE Echiodon dawsoni Williams & Shipp, 1982 1 1 1 1 CARAPIDAE Echiodon dentatus (Cuvier, 1829) 1 CENTRACANTHIDAE Centracanthus cirrus Rafinesque, 1810 1 1 1 1 1 1 CENTRACANTHIDAE Spicara alta (Osório, 1917) 1 CENTRACANTHIDAE Spicara australis (Regan, 1921) 1 1 1 CENTRACANTHIDAE Spicara axillaris (Boulenger, 1900) 1 1 1 CENTRACANTHIDAE Spicara maena (Linnaeus, 1758) 1 1 CENTRACANTHIDAE Spicara melanurus (Valenciennes, 1830) 1 1 CENTRACANTHIDAE Spicara nigricauda (Norman, 1931) 1 1 CENTRACANTHIDAE Spicara smaris (Linnaeus, 1758) 1 1 1 CHAENOPSIDAE Acanthemblemaria aspera (Longley, 1927) 1 1 CHAENOPSIDAE Acanthemblemaria betinensis Smith-Vaniz & Palacio, 1974 1 CHAENOPSIDAE Acanthemblemaria chaplini Böhlke, 1957 1 1 CHAENOPSIDAE Acanthemblemaria greenfieldi Smith-Vaniz & Palacio, 1974 1 CHAENOPSIDAE Acanthemblemaria harpeza Williams, 2002 1 CHAENOPSIDAE Acanthemblemaria johnsoni Almany & Baldwin, 1996 1 1 CHAENOPSIDAE Acanthemblemaria maria Böhlke, 1961 1 1 CHAENOPSIDAE Acanthemblemaria medusa Smith-Vaniz & Palacio, 1974 1 1 CHAENOPSIDAE Acanthemblemaria paula Johnson & Brothers, 1989 1 CHAENOPSIDAE Acanthemblemaria rivasi Stephens, 1970 1 CHAENOPSIDAE Acanthemblemaria spinosa Metzelaar, 1919 1 1 CHAENOPSIDAE Chaenopsis cf._stephensi Robins & Randall, 1965 1 CHAENOPSIDAE Chaenopsis limbaughi Robins & Randall, 1965 1 1 1 CHAENOPSIDAE Chaenopsis megalops Smith-Vaniz, 2000 1 CHAENOPSIDAE Chaenopsis ocellata Gill, 1865 1 1 CHAENOPSIDAE Chaenopsis resh Robins & Randall, 1965 1 1 CHAENOPSIDAE Chaenopsis roseola Hastings & Shipp, 1981 1 1 CHAENOPSIDAE Chaenopsis stephensi Robins & Randall, 1965 1 1 CHAENOPSIDAE Coralliozetus cardonae Evermann & Marsh, 1899 1 1 CHAENOPSIDAE Ekemblemaria nigra (Meek & Hildebrand, 1928) 1 CHAENOPSIDAE Emblemaria atlantica Jordan & Evermann, 1898 1 1 1 1 CHAENOPSIDAE Emblemaria australis Ramos, Rocha & Rocha, 2003 1 1 CHAENOPSIDAE Emblemaria biocellata Stephens, 1970 1 1 CHAENOPSIDAE Emblemaria caldwelli Stephens, 1970 1 1 CHAENOPSIDAE Emblemaria caycedoi Acero P., 1984 1 CHAENOPSIDAE Emblemaria culmenis Stephens, 1970 1 1 CHAENOPSIDAE Emblemaria diphyodontis Stephens & Cervigón, 1970 1 1 CHAENOPSIDAE Emblemaria hyltoni Johnson & Greenfield, 1976 1 CHAENOPSIDAE Emblemaria pandionis Evermann & Marsh, 1900 1 1 1 CHAENOPSIDAE Emblemaria piratula Ginsburg & Reid, 1942 1 1 CHAENOPSIDAE Emblemaria sp.n. 1 1 CHAENOPSIDAE Emblemaria vitta Williams, 2002 1 1 CHAENOPSIDAE Emblemariopsis bahamensis Stephens, 1961 1 1 CHAENOPSIDAE Emblemariopsis bottomei Stephens, 1961 1 1 CHAENOPSIDAE Emblemariopsis dianae Tyler & Hastings, 2004 1 CHAENOPSIDAE Emblemariopsis diaphanus Longley, 1927 1 CHAENOPSIDAE Emblemariopsis leptocirris Stephens, 1970 1 1 CHAENOPSIDAE Emblemariopsis occidentalis Stephens, 1970 1 1 CHAENOPSIDAE Emblemariopsis pricei Greenfield, 1975 1 CHAENOPSIDAE Emblemariopsis ramirezi (Cervigón, 1999) 1 1 CHAENOPSIDAE Emblemariopsis randalli Cervigón, 1965 1 1 CHAENOPSIDAE Emblemariopsis ruetzleri Tyler & Tyler, 1997 1 1 CHAENOPSIDAE Emblemariopsis signifera (Ginsburg, 1942) 1 1 1 1 CHAENOPSIDAE Emblemariopsis sp.n.1 1 1 CHAENOPSIDAE Emblemariopsis sp.n.2 1 CHAENOPSIDAE Emblemariopsis sp.n.3 1 1 CHAENOPSIDAE Emblemariopsis tayrona (Acero P., 1987) 1 CHAENOPSIDAE Hemiemblemaria simulus Longley & Hildebrand, 1940 1 1 CHAENOPSIDAE Lucayablennius zingaro (Böhlke, 1957) 1 1 CHAENOPSIDAE Protemblemaria punctata Cervigón, 1966 1 1 CHAENOPSIDAE Stathmonotus gymnodermis Springer, 1955 1 1 CHAENOPSIDAE Stathmonotus hemphillii Bean, 1885 1 1 1 CHAENOPSIDAE Stathmonotus sp. 1 CHAENOPSIDAE Stathmonotus stahli_stahli (Evermann & Marsh, 1899) 1 1 CHAENOPSIDAE Stathmonotus stahli_tekla Nichols, 1910 1 1 CHAETODONTIDAE Chaetodon auriga Forsskål, 1775 1 1 CHAETODONTIDAE Chaetodon capistratus Linnaeus, 1758 1 1 1 1 CHAETODONTIDAE Chaetodon hoefleri Steindachner, 1881 1 1 1 1 1 1 CHAETODONTIDAE Chaetodon interruptus Ahl, 1923 1 1 CHAETODONTIDAE Chaetodon madagaskariensis Ahl, 1923 1 1 CHAETODONTIDAE Chaetodon marleyi Regan, 1921 1 1 1 CHAETODONTIDAE Chaetodon ocellatus Bloch, 1787 1 1 1 1 1 1 1 1 CHAETODONTIDAE Chaetodon robustus Günther, 1860 1 1 1 1 CHAETODONTIDAE Chaetodon sanctaehelenae Günther, 1868 1 1 1 CHAETODONTIDAE Chaetodon sedentarius Poey, 1860 1 1 1 1 1 1 1 1 CHAETODONTIDAE Chaetodon striatus Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 CHAETODONTIDAE Chaetodon unimaculatus Bloch, 1787 1 1 CHAETODONTIDAE Chaetodon vagabundus Linnaeus, 1758 1 1 CHAETODONTIDAE Prognathodes aculeatus (Poey, 1860) 1 1 1 1 CHAETODONTIDAE Prognathodes aya (Jordan, 1886) 1 1 CHAETODONTIDAE Prognathodes brasiliensis Burgess, 2001 1 1 1 CHAETODONTIDAE Prognathodes dichrous (Günther, 1869) 1 1 CHAETODONTIDAE Prognathodes guyanensis (Durand, 1960) 1 1 1 1 1 CHAETODONTIDAE Prognathodes marcellae (Poll, 1950) 1 1 1 1 1 CHAETODONTIDAE Prognathodes obliquus (Lubbock & Edwards, 1980) 1 CHANIDAE Chanos chanos (Forsskål, 1775) 1 1 CHEILODACTYLIDAE Cheilodactylus fasciatus Lacepède, 1803 1 1 1 CHEILODACTYLIDAE Cheilodactylus pixi Smith, 1980 1 1 CHEILODACTYLIDAE Chirodactylus brachydactylus (Cuvier, 1830) 1 1 1 CHEILODACTYLIDAE Chirodactylus grandis (Günther, 1860) 1 1 1 CHEILODACTYLIDAE Nemadactylus bergi (Norman, 1937) 1 1 CHLOPSIDAE Catesbya pseudomuraena Böhlke & Smith, 1968 1 CHLOPSIDAE Chilorhinus suensonii Lütken, 1852 1 1 1 1 CHLOPSIDAE Chlopsis bicolor Rafinesque, 1810 1 1 1 1 1 1 1 1 CHLOPSIDAE Chlopsis dentatus (Seale, 1917) 1 1 1 1 1 1 CHLOPSIDAE Kaupichthys hyoproroides (Strömman, 1896) 1 1 1 1 1 CHLOPSIDAE Kaupichthys nuchalis Böhlke, 1967 1 1 1 CIRRHITIDAE Amblycirrhitus earnshawi Lubbock, 1978 1 CIRRHITIDAE Amblycirrhitus pinos (Mowbray, 1927) 1 1 1 1 1 1 1 1 CIRRHITIDAE Cirrhitus atlanticus Osório, 1893 1 1 1 CIRRHITIDAE Cirrhitus pinnulatus (Forster, 1801) 1 1 CIRRHITIDAE Cyprinocirrhites polyactis (Bleeker, 1874) 1 1 CLINIDAE Blennioclinus brachycephalus (Valenciennes, 1836) 1 1 1 CLINIDAE Blennioclinus stella Smith, 1946 1 1 CLINIDAE Blennophis anguillaris (Valenciennes, 1836) 1 1 1 CLINIDAE Blennophis striatus (Gilchrist & Thompson, 1908) 1 1 1 CLINIDAE Calliclinus geniguttatus (Valenciennes, 1836) 1 CLINIDAE Cancelloxus burrelli Smith, 1961 1 1 CLINIDAE Cancelloxus elongatus Heemstra & Wright, 1986 1 CLINIDAE Cirrhibarbis capensis Valenciennes, 1836 1 1 1 CLINIDAE Climacoporus navalis Barnard, 1935 1 1 CLINIDAE Clinitrachus argentatus (Risso, 1810) 1 1 CLINIDAE Clinoporus biporosus (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Clinus acuminatus (Bloch & Schneider, 1801) 1 1 CLINIDAE Clinus agilis Smith, 1931 1 1 CLINIDAE Clinus berrisfordi Penrith, 1967 1 1 CLINIDAE Clinus brevicristatus Gilchrist & Thompson, 1908 1 1 CLINIDAE Clinus cottoides Valenciennes, 1836 1 1 1 CLINIDAE Clinus helenae (Smith, 1946) 1 1 CLINIDAE Clinus heterodon Valenciennes, 1836 1 1 CLINIDAE Clinus latipennis Valenciennes, 1836 1 1 CLINIDAE Clinus navalis (Barnard, 1935) 1 1 1 CLINIDAE Clinus nematopterus Günther, 1861 1 CLINIDAE Clinus robustus Gilchrist & Thompson, 1908 1 1 CLINIDAE Clinus rotundifrons Barnard, 1937 1 1 CLINIDAE Clinus spatulatus Bennett, 1983 1 CLINIDAE Clinus superciliosus (Linnaeus, 1758) 1 1 1 CLINIDAE Clinus taurus Gilchrist & Thompson, 1908 1 1 CLINIDAE Clinus venustris Gilchrist & Thompson, 1908 1 1 CLINIDAE Clinus woodi (Smith, 1946) 1 1 CLINIDAE Fucomimus mus (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Muraenoclinus dorsalis (Bleeker, 1860) 1 1 1 CLINIDAE Pavoclinus graminis (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Pavoclinus laurentii (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Pavoclinus litorafontis Penrith, 1965 1 CLINIDAE Pavoclinus mentalis (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Pavoclinus myae Christensen, 1978 1 1 CLINIDAE Pavoclinus pavo (Gilchrist & Thompson, 1908) 1 1 1 CLINIDAE Pavoclinus profundus Smith, 1961 1 CLINIDAE Pavoclinus smalei Heemstra & Wright, 1986 1 CLINIDAE Ribeiroclinus eigenmanni (Jordan, 1888) 1 1 1 CLINIDAE Smithichthys fucorum (Gilchrist & Thompson, 1908) 1 1 CLINIDAE Xenopoclinus kochi Smith, 1948 1 1 CLINIDAE Xenopoclinus leprosus Smith, 1961 1 1 CONGRIDAE Ariosoma anale (Poey, 1860) 1 1 1 1 1 CONGRIDAE Ariosoma balearicum (Delaroche, 1809) 1 1 1 1 1 1 1 1 1 1 1 1 1 CONGRIDAE Bassanago albescens (Barnard, 1923) 1 CONGRIDAE Conger cinereus Rüppell, 1830 1 1 CONGRIDAE Conger conger (Linnaeus, 1758) 1 1 1 1 1 1 1 CONGRIDAE Conger orbignianus Valenciennes, 1842 1 1 1 1 CONGRIDAE Conger triporiceps Kanazawa, 1958 1 1 1 1 CONGRIDAE Conger wilsoni (Bloch & Schneider, 1801) 1 1 CONGRIDAE Gorgasia inferomaculata (Blache, 1977) 1 1 CONGRIDAE Heteroconger camelopardalis (Lubbock, 1980) 1 1 1 1 1 1 CONGRIDAE Heteroconger longissimus Günther, 1870 1 1 1 1 1 1 1 1 1 1 1 CONGRIDAE Heteroconger luteolus Smith, 1989 1 1 CONGRIDAE Paraconger caudilimbatus (Poey, 1867) 1 1 1 CONGRIDAE Paraconger guianensis Kanazawa, 1961 1 CONGRIDAE Paraconger macrops (Günther, 1870) 1 1 CONGRIDAE Paraconger notialis Kanazawa, 1961 1 1 1 1 DACTYLOPTERIDAE Dactyloptena orientalis (Cuvier, 1829) 1 1 DACTYLOPTERIDAE Dactylopterus volitans (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 DACTYLOSCOPIDAE Dactylagnus peratikos Böhlke & Caldwell, 1961 1 DACTYLOSCOPIDAE Dactyloscopus boehlkei Dawson, 1982 1 DACTYLOSCOPIDAE Dactyloscopus comptus Dawson, 1982 1 DACTYLOSCOPIDAE Dactyloscopus crossotus Starks, 1913 1 1 1 1 1 DACTYLOSCOPIDAE Dactyloscopus foraminosus Dawson, 1982 1 1 1 DACTYLOSCOPIDAE Dactyloscopus moorei (Fowler, 1906) 1 1 1 DACTYLOSCOPIDAE Dactyloscopus poeyi Gill, 1861 1 1 DACTYLOSCOPIDAE Dactyloscopus tridigitatus Gill, 1859 1 1 1 1 1 1 1 1 DACTYLOSCOPIDAE Gillellus greyae Kanazawa, 1952 1 1 1 1 1 1 DACTYLOSCOPIDAE Gillellus healae Dawson, 1982 1 1 1 DACTYLOSCOPIDAE Gillellus inescatus Williams, 2002 1 DACTYLOSCOPIDAE Gillellus jacksoni Dawson, 1982 1 DACTYLOSCOPIDAE Gillellus uranidea Böhlke, 1968 1 1 1 DACTYLOSCOPIDAE Gillelus sp.n. 1 1 DACTYLOSCOPIDAE Leurochilus acon Böhlke, 1968 1 DACTYLOSCOPIDAE Myxodagnus belone Böhlke, 1968 1 DACTYLOSCOPIDAE Platygillellus brasiliensis Feitoza, 2002 1 DACTYLOSCOPIDAE Platygillellus rubrocinctus (Longley, 1934) 1 1 1 DACTYLOSCOPIDAE Platygillellus smithi Dawson, 1982 1 DACTYLOSCOPIDAE Storrsia olsoni Dawson, 1982 1 1 DIODONTIDAE Chilomycterus antennatus (Cuvier, 1816) 1 1 1 1 DIODONTIDAE Chilomycterus antillarum Jordan & Rutter, 1897 1 1 1 1 1 1 DIODONTIDAE Chilomycterus reticulatus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 DIODONTIDAE Chilomycterus schoepfii (Walbaum, 1792) 1 1 1 DIODONTIDAE Chilomycterus spinosus_mauretanic(uLse Danois, 1954) 1 1 1 1 DIODONTIDAE Chilomycterus spinosus_spinosus (Linnaeus, 1758) 1 1 1 DIODONTIDAE Cyclichthys orbicularis (Bloch, 1785) 1 1 DIODONTIDAE Cyclichthys spilostylus (Leis & Randall, 1982) 1 1 DIODONTIDAE Diodon eydouxii Brisout de Barneville, 1846 1 1 1 1 1 1 1 1 DIODONTIDAE Diodon holocanthus Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 DIODONTIDAE Diodon hystrix Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 DIODONTIDAE Diodon liturosus Shaw, 1804 1 1 DIODONTIDAE Lophodiodon calori (Bianconi, 1854) 1 1 FISTULARIIDAE Fistularia commersonii Rüppell, 1838 1 1 FISTULARIIDAE Fistularia petimba Lacepède, 1803 1 1 1 1 1 1 1 1 1 1 1 1 1 FISTULARIIDAE Fistularia tabacaria Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 GOBIESOCIDAE Acyrtops amplicirrus Briggs, 1955 1 GOBIESOCIDAE Acyrtops beryllinus (Hildebrand & Ginsburg, 1926) 1 1 1 1 GOBIESOCIDAE Acyrtops sp. 1 1 1 GOBIESOCIDAE Acyrtus artius Briggs, 1955 1 1 1 1 GOBIESOCIDAE Acyrtus pauciradiatus Sampaio, de Anchieta, Nunes & Mendes, 2004 1 GOBIESOCIDAE Acyrtus rubiginosus (Poey, 1868) 1 1 GOBIESOCIDAE Apletodon dentatus (Facciolà, 1887) 1 1 1 1 GOBIESOCIDAE Apletodon incognitus Hofrichter & Patzner, 1997 1 1 1 1 1 GOBIESOCIDAE Apletodon pellegrini (Chabanaud, 1925) 1 1 1 1 1 GOBIESOCIDAE Apletodon wirtzi Fricke, 2007 1 GOBIESOCIDAE Arcos macrophthalmus (Günther, 1861) 1 1 1 GOBIESOCIDAE Chorisochismus dentex (Pallas, 1769) 1 1 1 GOBIESOCIDAE Derilissus altifrons Smith-Vaniz, 1971 1 1 GOBIESOCIDAE Derilissus kremnobates Fraser, 1970 1 GOBIESOCIDAE Derilissus nanus Briggs, 1969 1 GOBIESOCIDAE Derilissus bviitmtigaecrulata Fraser, 1970 1 GOBIESOCIDAE Diplecogaster bimaculata (Bonnaterre, 1788) 1 1 1 GOBIESOCIDAE Diplecogaster pectoralis Briggs, 1955 1 1 1 1 GOBIESOCIDAE Diplecogaster ctenocrypta Briggs, 1955 1 GOBIESOCIDAE Diplecogaster megalops Briggs, 1955 1 1 1 GOBIESOCIDAE Eckloniaichthys scylliorhiniceps Smith, 1943 1 1 1 GOBIESOCIDAE Gobiesox barbatulus Starks, 1913 1 1 1 1 GOBIESOCIDAE Gobiesox lucayanus Briggs, 1963 1 GOBIESOCIDAE Gobiesox marmoratus Jenyns, 1842 1 GOBIESOCIDAE Gobiesox nigripinnis (Peters, 1860) 1 1 GOBIESOCIDAE Gobiesox punctulatus (Poey, 1876) 1 1 1 GOBIESOCIDAE Gobiesox strumosus Cope, 1870 1 1 1 1 1 1 GOBIESOCIDAE Gouania willdenowi (Risso, 1810) 1 1 GOBIESOCIDAE Lecanogaster chrysea Briggs, 1957 1 GOBIESOCIDAE Lepadogaster candolii Risso, 1810 1 1 1 1 1 GOBIESOCIDAE Lepadogaster lepadogaster (Bonnaterre, 1788) 1 1 1 1 1 1 GOBIESOCIDAE Lepadogaster purpurea (Bonnaterre, 1788) 1 1 1 1 GOBIESOCIDAE Opeatogenys cadenati Briggs, 1957 1 1 1 1 GOBIESOCIDAE Opeatogenys gracilis (Canestrini, 1864) 1 1 GOBIESOCIDAE Tomicodon australis Briggs, 1955 1 1 1 GOBIESOCIDAE Tomicodon briggsi Williams & Tyler, 2003 1 1 GOBIESOCIDAE Tomicodon clarkei Williams & Tyler, 2003 1 GOBIESOCIDAE Tomicodon cryptus Williams & Tyler, 2003 1 1 GOBIESOCIDAE Tomicodon fasciatus (Peters, 1859) 1 1 GOBIESOCIDAE Tomicodon lavettsmithi Williams & Tyler, 2003 1 GOBIESOCIDAE Tomicodon leurodiscus Williams & Tyler, 2003 1 1 GOBIESOCIDAE Tomicodon reitzae Briggs, 2001 1 1 GOBIESOCIDAE Tomicodon rhabdotus Smith-Vaniz, 1969 1 GOBIESOCIDAE Tomicodon rupestris (Poey, 1860) 1 1 GOBIIDAE Barbulifer antennatus Böhlke & Robins, 1968 1 GOBIIDAE Barbulifer ceuthoecus (Jordan & Gilbert, 1884) 1 1 1 1 GOBIIDAE Bathygobius burtoni (O'Shaughnessy, 1875) 1 1 1 GOBIIDAE Bathygobius casamancus (Rochebrune, 1880) 1 1 1 1 GOBIIDAE Bathygobius curacao (Metzelaar, 1919) 1 1 1 1 GOBIIDAE Bathygobius mystacium Ginsburg, 1947 1 1 1 1 GOBIIDAE Bathygobius soporator (Valenciennes, 1837) 1 1 1 1 1 1 1 1 1 1 1 1 1 GOBIIDAE Bollmannia boqueronensis Evermann & Marsh, 1899 1 1 GOBIIDAE Bollmannia communis Ginsburg, 1942 1 1 GOBIIDAE Bollmannia eigenmanni (Garman, 1896) 1 GOBIIDAE Bollmannia litura Ginsburg, 1935 1 GOBIIDAE Bollmannia sp. n. 1 1 GOBIIDAE Buenia affinis Iljin, 1930 1 1 GOBIIDAE Buenia jeffreysii (Günther, 1867) 1 GOBIIDAE Buenia sp.n.(Canaries) 1 1 GOBIIDAE Caffrogobius agulhensis (Barnard, 1927) 1 1 GOBIIDAE Caffrogobius caffer (Günther, 1874) 1 1 GOBIIDAE Caffrogobius multifasciatus (Smith, 1959) 1 1 GOBIIDAE Caffrogobius natalensis (Günther, 1874) 1 1 GOBIIDAE Caffrogobius nudiceps (Valenciennes, 1837) 1 1 1 GOBIIDAE Caffrogobius saldanha (Barnard, 1927) 1 1 1 GOBIIDAE Chriolepis benthonis Ginsburg, 1953 1 GOBIIDAE Chriolepis fisheri Herre, 1942 1 1 1 GOBIIDAE Chriolepis vespa Hastings & Bortone, 1981 1 1 GOBIIDAE Chriolepsis sp.n. 1 1 GOBIIDAE Chromogobius britoi Van Tassell, 2001 1 1 GOBIIDAE Chromogobius quadrivittatus (Steindachner, 1863) 1 1 GOBIIDAE Chromogobius zebratus (Kolombatovic, 1891) 1 1 GOBIIDAE Corcyrogobius liechtensteini (Kolombatovic, 1891) 1 1 GOBIIDAE Corcyrogobius lubbocki Miller, 1988 1 1 GOBIIDAE Coryphopterus alloides Böhlke & Robins, 1960 1 1 GOBIIDAE Coryphopterus dicrus Böhlke & Robins, 1960 1 1 1 1 1 GOBIIDAE Coryphopterus eidolon Böhlke & Robins, 1960 1 1 1 GOBIIDAE Coryphopterus glaucofraenum Gill, 1863 1 1 1 1 1 1 1 1 GOBIIDAE Coryphopterus hyalinus Böhlke & Robins, 1962 1 1 1 GOBIIDAE Coryphopterus lipernes Böhlke & Robins, 1962 1 1 GOBIIDAE Coryphopterus personatus (Jordan & Thompson, 1905) 1 1 1 1 GOBIIDAE Coryphopterus punctipectophorus Springer, 1960 1 1 GOBIIDAE Coryphopterus thrix Böhlke & Robins, 1960 1 1 1 1 GOBIIDAE Coryphopterus venezuelae Cervigón, 1966 1 1 GOBIIDAE Ctenogobius saepepallens (Gilbert & Randall, 1968) 1 1 1 1 1 GOBIIDAE Deltentosteus collonianus (Risso, 1820) 1 1 GOBIIDAE Deltentosteus quadrimaculatus (Valenciennes, 1837) 1 1 GOBIIDAE Didogobius bentuvii Miller, 1966 1 GOBIIDAE Didogobius kochi Van Tassell, 1988 1 1 GOBIIDAE Didogobius schlieweni Miller, 1993 1 1 GOBIIDAE Didogobius sp.n. 1 GOBIIDAE Didogobius splechtnai Ahnelt & Patzner, 1995 1 1 GOBIIDAE Elacatinus atronasus (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus chancei (Beebe & Hollister, 1933) 1 1 GOBIIDAE Elacatinus dilepis (Robins & Böhlke, 1964) 1 1 GOBIIDAE Elacatinus evelynae (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus figaro Sazima, Moura & Rosa, 1997 1 1 GOBIIDAE Elacatinus gemmatus (Ginsburg, 1939) 1 1 GOBIIDAE Elacatinus genie (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus horsti (Metzelaar, 1922) 1 1 GOBIIDAE Elacatinus illecebrosus (Böhlke & Robins, 1968) 1 GOBIIDAE Elacatinus lori Colin, 2002 1 GOBIIDAE Elacatinus louisae (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus macrodon (Beebe & Tee-Van, 1928) 1 1 1 GOBIIDAE Elacatinus multifasciatus (Steindachner, 1876) 1 1 GOBIIDAE Elacatinus oceanops Jordan, 1904 1 1 1 GOBIIDAE Elacatinus pallens (Ginsburg, 1939) 1 1 GOBIIDAE Elacatinus pridisi Guimarães, Gasparini & Rocha, 2004 1 GOBIIDAE Elacatinus prochilos (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus randalli (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus saucrus (Robins, 1960) 1 1 GOBIIDAE Elacatinus sp.n.(Noronha) 1 1 GOBIIDAE Elacatinus tenox (Böhlke & Robins, 1968) 1 1 GOBIIDAE Elacatinus xanthiprora (Böhlke & Robins, 1968) 1 1 1 GOBIIDAE Elacatinus zebrellus (Robins, 1958) 1 1 GOBIIDAE Evermannichthys bicolor Thacker, 2001 1 GOBIIDAE Evermannichthys convictor Böhlke & Robins, 1969 1 GOBIIDAE Evermannichthys metzelaari Hubbs, 1923 1 1 GOBIIDAE Evermannichthys spilus Böhlke & Robins, 1969 1 GOBIIDAE Evermannichthys spongicola (Radcliffe, 1917) 1 1 1 GOBIIDAE Gammogobius steinitzi Bath, 1971 1 1 GOBIIDAE Ginsburgellus novemlineatus (Fowler, 1950) 1 1 GOBIIDAE Glossogobius callidus (Smith, 1937) 1 1 GOBIIDAE Gnatholepis thompsoni Jordan, 1904 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 GOBIIDAE Gobiosoma grosvenori (Robins, 1964) 1 1 GOBIIDAE Gobiosoma hemigymnum (Eigenmann & Eigenmann, 1888) 1 1 1 GOBIIDAE Gobiosoma hildebrandi (Ginsburg, 1939) 1 GOBIIDAE Gobiosoma longipala Ginsburg, 1933 1 1 GOBIIDAE Gobiosoma parri Ginsburg, 1933 1 GOBIIDAE Gobiosoma robustum Ginsburg, 1933 1 1 1 GOBIIDAE Gobiosoma spes (Ginsburg, 1939) 1 1 GOBIIDAE Gobiosoma spilotum (Ginsburg, 1939) 1 1 GOBIIDAE Gobiosoma yucatanum Dawson, 1971 1 GOBIIDAE Gobius ater Bellotti, 1888 1 1 GOBIIDAE Gobius ateriformis Brito & Miller, 2001 1 GOBIIDAE Gobius auratus Risso, 1810 1 1 GOBIIDAE Gobius bucchichi Steindachner, 1870 1 1 1 GOBIIDAE Gobius cobitis Pallas, 1814 1 1 1 GOBIIDAE Gobius cruentatus Gmelin, 1789 1 1 1 1 GOBIIDAE Gobius fallax Sarato, 1889 1 1 GOBIIDAE Gobius gasteveni Miller, 1974 1 1 1 GOBIIDAE Gobius geniporus Valenciennes, 1837 1 1 GOBIIDAE Gobius kolombatovici Kovacic & Miller, 2000 1 GOBIIDAE Gobius niger Linnaeus, 1758 1 1 1 1 1 GOBIIDAE Gobius paganellus Linnaeus, 1758 1 1 1 1 1 1 1 GOBIIDAE Gobius roulei de Buen, 1928 1 1 1 GOBIIDAE Gobius rubropunctatus Delais, 1951 1 1 GOBIIDAE Gobius senegambiensis Metzelaar, 1919 1 1 1 GOBIIDAE Gobius strictus Fage, 1907 1 1 GOBIIDAE Gobius tetrophthalmus Brito & Miller, 2001 1 GOBIIDAE Gobius vittatus Vinciguerra, 1883 1 1 GOBIIDAE Gobius xanthocephalus Heymer & Zander, 1992 1 1 1 GOBIIDAE Gobulus myersi Ginsburg, 1939 1 1 1 1 GOBIIDAE Gorogobius nigricinctus (Delais, 1951) 1 1 1 GOBIIDAE Hetereleotris zonata (Fowler, 1934) 1 1 GOBIIDAE Knipowitschia caucasica (Berg, 1916) 1 GOBIIDAE Knipowitschia panizzae (Verga, 1841) 1 GOBIIDAE Lebetus guilleti (Le Danois, 1913) 1 1 GOBIIDAE Lebetus scorpioides (Collett, 1874) 1 1 GOBIIDAE Lesueurigobius friesii (Malm, 1874) 1 1 GOBIIDAE Lesueurigobius heterofasciatus Maul, 1971 1 1 1 GOBIIDAE Lesueurigobius koumansi (Norman, 1935) 1 1 GOBIIDAE Lesueurigobius sanzi (de Buen, 1918) 1 GOBIIDAE Lesueurigobius suerii (Risso, 1810) 1 1 1 1 1 GOBIIDAE Lythrypnus brasiliensis Greenfield, 1988 1 1 GOBIIDAE Lythrypnus crocodilus (Beebe & Tee-Van, 1928) 1 1 1 GOBIIDAE Lythrypnus elasson Böhlke & Robins, 1960 1 1 GOBIIDAE Lythrypnus heterochroma Ginsburg, 1939 1 1 GOBIIDAE Lythrypnus minimus Garzón & Acero P., 1988 1 1 GOBIIDAE Lythrypnus mowbrayi (Bean, 1906) 1 GOBIIDAE Lythrypnus nesiotes Böhlke & Robins, 1960 1 1 GOBIIDAE Lythrypnus okapia Robins & Böhlke, 1964 1 1 1 GOBIIDAE Lythrypnus phorellus Böhlke & Robins, 1960 1 1 1 1 GOBIIDAE Lythrypnus sp.1(Noronha) 1 1 GOBIIDAE Lythrypnus sp.2(Trindade) 1 GOBIIDAE Lythrypnus spilus Böhlke & Robins, 1960 1 1 1 1 GOBIIDAE Mauligobius maderensis (Valenciennes, 1837) 1 1 GOBIIDAE Mauligobius nigri (Günther, 1861) 1 GOBIIDAE Microgobius carri Fowler, 1945 1 1 1 1 1 GOBIIDAE Microgobius microlepis Longley & Hildebrand, 1940 1 1 1 GOBIIDAE Microgobius signatus Poey, 1876 1 1 GOBIIDAE Millerigobius macrocephalus (Smith, 1948) 1 1 GOBIIDAE Monishia william (Smith, 1948) 1 1 GOBIIDAE Nematogobius brachynemus Pfaff, 1933 1 1 1 GOBIIDAE Nematogobius maindroni (Sauvage, 1880) 1 1 1 1 GOBIIDAE Nes longus (Nichols, 1914) 1 1 1 1 GOBIIDAE Odondebuenia balearica (Pellegrin & Fage, 1907) 1 1 GOBIIDAE Oligolepis acutipennis (Valenciennes, 1837) 1 1 GOBIIDAE Oligolepis keiensis (Smith, 1938) 1 1 GOBIIDAE Ophiogobius ophicephalus (Jenyns, 1842) 1 GOBIIDAE Palatogobius paradoxus Gilbert, 1971 1 1 GOBIIDAE Pariah scotius Böhlke, 1969 1 GOBIIDAE Parrella macropteryx Ginsburg, 1939 1 1 1 GOBIIDAE Periophthalmus barbarus (Linnaeus, 1766) 1 1 GOBIIDAE Pomatoschistus bathi Miller, 1982 1 1 GOBIIDAE Pomatoschistus canestrinii (Ninni, 1883) 1 GOBIIDAE Pomatoschistus knerii (Steindachner, 1861) 1 1 GOBIIDAE Pomatoschistus marmoratus (Risso, 1810) 1 1 GOBIIDAE Pomatoschistus microps (Krøyer, 1838) 1 1 1 GOBIIDAE Pomatoschistus minutus (Pallas, 1770) 1 1 1 GOBIIDAE Pomatoschistus norvegicus (Collett, 1902) 1 1 GOBIIDAE Pomatoschistus pictus (Malm, 1865) 1 1 1 1 GOBIIDAE Pomatoschistus quagga (Heckel, 1837) 1 1 GOBIIDAE Pomatoschistus tortonesei Miller, 1969 1 GOBIIDAE Porogobius schlegelii (Günther, 1861) 1 1 1 1 GOBIIDAE Priolepis ascensionis (Dawson & Edwards, 1987) 1 1 GOBIIDAE Priolepis dawsoni Greenfield, 1989 1 1 1 1 1 GOBIIDAE Priolepis hipoliti (Metzelaar, 1922) 1 1 1 1 GOBIIDAE Priolepis robinsi Garzón-Ferreira & Acero P., 1991 1 GOBIIDAE Priolepis sp.n. 1 1 GOBIIDAE Psammogobius knysnaensis Smith, 1935 1 1 1 GOBIIDAE Pseudaphya ferreri (de Buen & Fage, 1908) 1 1 GOBIIDAE Psilotris alepis Ginsburg, 1953 1 1 GOBIIDAE Psilotris amblyrhynchus Smith & Baldwin, 1999 1 GOBIIDAE Psilotris batrachodes Böhlke, 1963 1 1 GOBIIDAE Psilotris boehlkei Greenfield, 1993 1 GOBIIDAE Psilotris celsus Böhlke, 1963 1 1 1 1 1 GOBIIDAE Psilotris kaufmani Greenfield, Findley & Johnson, 1993 1 1 GOBIIDAE Pycnomma roosevelti Ginsburg, 1939 1 1 GOBIIDAE Redigobius dewaali (Weber, 1897) 1 1 GOBIIDAE Risor ruber (Rosén, 1911) 1 1 1 1 1 GOBIIDAE Speleogobius trigloides Zander & Jelinek, 1976 1 GOBIIDAE Thorogobius angolensis (Norman, 1935) 1 1 GOBIIDAE Thorogobius ephippiatus (Lowe, 1839) 1 1 1 1 1 GOBIIDAE Thorogobius macrolepis (Kolombatovic, 1891) 1 1 GOBIIDAE Thorogobius rofeni Miller, 1988 1 GOBIIDAE Vanneaugobius canariensis Van Tassell, Miller & Brito, 1988 1 1 1 1 1 GOBIIDAE Vanneaugobius dollfusi Brownell, 1978 1 1 GOBIIDAE Vanneaugobius pruvoti (Fage, 1907) 1 1 GOBIIDAE Varicus bucca Robins & Böhlke, 1961 1 GOBIIDAE Varicus imswe Greenfield, 1981 1 GOBIIDAE Varicus marilynae Gilmore, 1979 1 1 GOBIIDAE Vomerogobius flavus Gilbert, 1971 1 1 GOBIIDAE Wheelerigobius maltzani (Steindachner, 1881) 1 1 1 GOBIIDAE Wheelerigobius wirtzi Miller, 1988 1 1 GOBIIDAE Zebrus zebrus (Risso, 1827) 1 1 GOBIIDAE Zosterisessor ophiocephalus (Pallas, 1814) 1 1 GRAMMATIDAE Gramma brasiliensis Sazima, Gasparini & Moura, 1998 1 1 1 GRAMMATIDAE Gramma linki Starck & Colin, 1978 1 1 GRAMMATIDAE Gramma loreto Poey, 1868 1 1 GRAMMATIDAE Gramma melacara Böhlke & Randall, 1963 1 1 GRAMMATIDAE Lipogramma anabantoides Böhlke, 1960 1 1 GRAMMATIDAE Lipogramma evides Robins & Colin, 1979 1 1 GRAMMATIDAE Lipogramma flavescens Gilmore & Jones, 1988 1 1 GRAMMATIDAE Lipogramma klayi Randall, 1963 1 1 GRAMMATIDAE Lipogramma regium Robins & Colin, 1979 1 1 1 GRAMMATIDAE Lipogramma robinsi Gilmore, 1997 1 1 GRAMMATIDAE Lipogramma roseum Gilbert, 1979 1 1 GRAMMATIDAE Lipogramma trilineatum Randall, 1963 1 1 HAEMULIDAE Anisotremus moricandi (Ranzani, 1842) 1 1 1 1 HAEMULIDAE Anisotremus surinamensis (Bloch, 1791) 1 1 1 1 1 1 1 1 1 HAEMULIDAE Anisotremus virginicus (Linnaeus, 1758) 1 1 1 1 1 HAEMULIDAE Haemulon album Cuvier, 1830 1 1 1 1 1 HAEMULIDAE Haemulon aurolineatum Cuvier, 1830 1 1 1 1 1 1 HAEMULIDAE Haemulon steindachneri (Jordan & Gilbert, 1882) 1 1 1 1 1 HAEMULIDAE Haemulon boschmae (Metzelaar, 1919) 1 1 HAEMULIDAE Haemulon bonariense Cuvier, 1830 1 1 HAEMULIDAE Haemulon carbonarium Poey, 1860 1 1 1 1 HAEMULIDAE Haemulon chrysargyreum Günther, 1859 1 1 1 1 1 HAEMULIDAE Haemulon flavolineatum (Desmarest, 1823) 1 1 1 1 HAEMULIDAE Haemulon macrostomum Günther, 1859 1 1 1 HAEMULIDAE Haemulon melanurum (Linnaeus, 1758) 1 1 1 1 1 HAEMULIDAE Haemulon parra (Desmarest, 1823) 1 1 1 1 1 1 1 HAEMULIDAE Haemulon plumieri (Lacepède, 1801) 1 1 1 1 1 HAEMULIDAE Haemulon sciurus (Lacepède, 1801) 1 1 1 1 HAEMULIDAE Haemulon squamipinna Rocha & Rosa, 1999 1 1 HAEMULIDAE Haemulon striatum (Linnaeus, 1758) 1 1 1 1 1 HAEMULIDAE Orthopristis chrysoptera (Linnaeus, 1766) 1 1 1 1 HAEMULIDAE Orthopristis ruber (Cuvier, 1830) 1 1 1 1 HAEMULIDAE Parapristipoma humile (Bowdich, 1825) 1 1 1 HAEMULIDAE Parapristipoma octolineatum (Valenciennes, 1833) 1 1 1 1 1 1 1 HAEMULIDAE Plectorhinchus chubbi (Regan, 1919) 1 1 HAEMULIDAE Plectorhinchus macrolepis (Boulenger, 1899) 1 1 1 HAEMULIDAE Plectorhinchus mediterraneus (Guichenot, 1850) 1 1 1 1 1 1 1 HAEMULIDAE Pomadasys commersonnii (Lacepède, 1801) 1 1 HAEMULIDAE Pomadasys corvinaeformis (Steindachner, 1868) 1 1 1 HAEMULIDAE Pomadasys crocro (Cuvier, 1830) 1 1 HAEMULIDAE Pomadasys incisus (Bowdich, 1825) 1 1 1 1 1 1 1 1 HAEMULIDAE Pomadasys jubelini (Cuvier, 1830) 1 1 1 1 HAEMULIDAE Pomadasys multimaculatum (Playfair, 1867) 1 1 HAEMULIDAE Pomadasys olivaceus (Day, 1875) 1 1 1 HAEMULIDAE Pomadasys perotaei (Cuvier, 1830) 1 1 1 1 HAEMULIDAE Pomadasys rogerii (Cuvier, 1830) 1 1 1 1 HAEMULIDAE Pomadasys striatus (Gilchrist & Thompson, 1908) 1 1 HAEMULIDAE Pomadasys stridens (Forsskål, 1775) 1 HAEMULIDAE Pomadasys suillus (Valenciennes, 1833) 1 1 HOLOCENTRIDAE Corniger spinosus Agassiz, 1831 1 1 1 1 1 1 1 HOLOCENTRIDAE Holocentrus adscensionis (Osbeck, 1765) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 HOLOCENTRIDAE Holocentrus rufus (Walbaum, 1792) 1 1 1 1 1 1 HOLOCENTRIDAE Myripristis jacobus Cuvier, 1829 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 HOLOCENTRIDAE Neoniphon marianus (Cuvier, 1829) 1 1 1 HOLOCENTRIDAE Ostichthys trachypoma (Günther, 1859) 1 1 1 HOLOCENTRIDAE Plectrypops retrospinis (Guichenot, 1853) 1 1 1 1 1 1 1 1 HOLOCENTRIDAE Sargocentron bullisi (Cuvier, 1829) 1 1 1 1 1 1 HOLOCENTRIDAE Sargocentron coruscum (Poey, 1860) 1 1 1 HOLOCENTRIDAE Sargocentron hastatum (Cuvier, 1829) 1 1 1 1 1 1 HOLOCENTRIDAE Sargocentron poco (Woods, 1965) 1 1 1 HOLOCENTRIDAE Sargocentron punctatissimum (Cuvier, 1829) 1 1 HOLOCENTRIDAE Sargocentron vexillarium (Poey, 1860) 1 1 1 1 KUHLIIDAE Kuhlia mugil (Forster, 1801) 1 1 KUHLIIDAE Parakuhlia macrophthalmus (Osório, 1893) 1 KYPHOSIDAE Girella stuebeli Troschel, 1866 1 KYPHOSIDAE Kyphosus bigibbus Lacepède, 1801 1 1 KYPHOSIDAE Kyphosus incisor (Cuvier, 1831) 1 1 1 1 1 1 1 1 1 1 1 1 1 KYPHOSIDAE Kyphosus sectatrix (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 KYPHOSIDAE Kyphosus vaigiensis (Quoy & Gaimard, 1825) 1 1 LABRIDAE Acantholabrus palloni (Risso, 1810) 1 1 1 1 1 1 1 1 1 LABRIDAE Anampses caeruleopunctatus Rüppell, 1829 1 1 LABRIDAE Bodianus insularis Gomon & Lubbock, 1980 1 1 1 LABRIDAE Bodianus pulchellus (Poey, 1860) 1 1 1 1 1 1 1 1 1 1 LABRIDAE Bodianus rufus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 LABRIDAE Bodianus scrofa (Valenciennes, 1839) 1 1 1 1 LABRIDAE Bodianus speciosus (Bowdich, 1825) 1 1 1 1 LABRIDAE Clepticus africanus Heiser, Moura & Robertson, 2000 1 LABRIDAE Clepticus brasiliensis Heiser, Moura & Robertson, 2000 1 1 1 1 1 LABRIDAE Clepticus parrae (Bloch & Schneider, 1801) 1 1 1 1 LABRIDAE Coris atlantica Günther, 1862 1 1 1 1 LABRIDAE Coris julis (Linnaeus, 1758) 1 1 1 1 1 1 LABRIDAE Ctenolabrus rupestris (Linnaeus, 1758) 1 1 1 LABRIDAE Decodon puellaris (Poey, 1860) 1 1 1 1 1 1 LABRIDAE Doratonotus megalepis Günther, 1862 1 1 1 1 1 1 1 1 1 1 LABRIDAE Halichoeres bathyphilus (Beebe & Tee-Van, 1932) 1 1 1 1 1 LABRIDAE Halichoeres bivittatus (Bloch, 1791) 1 1 1 1 1 LABRIDAE Halichoeres brasiliensis (Bloch, 1791) 1 1 1 LABRIDAE Halichoeres caudalis (Poey, 1860) 1 1 1 LABRIDAE Halichoeres cyanocephalus (Bloch, 1791) 1 1 1 LABRIDAE Halichoeres dimidiatus (Agassiz, 1831) 1 1 1 1 LABRIDAE Halichoeres garnoti (Valenciennes, 1839) 1 1 1 1 LABRIDAE Halichoeres maculipinna (Müller & Troschel, 1848) 1 1 1 1 LABRIDAE Halichoeres nebulosus (Valenciennes, 1839) 1 1 LABRIDAE Halichoeres penrosei Starks, 1913 1 1 1 LABRIDAE Halichoeres pictus (Poey, 1860) 1 1 1 1 LABRIDAE Halichoeres poeyi (Steindachner, 1867) 1 1 1 1 1 1 LABRIDAE Halichoeres radiatus (Linnaeus, 1758) 1 1 1 1 1 1 1 LABRIDAE Halichoeres socialis Randall & Lobel, 2003 1 LABRIDAE Halichoeres sp.n. (G.of Mexico) 1 LABRIDAE Labroides dimidiatus (Valenciennes, 1839) 1 1 LABRIDAE Labrus bergylta Ascanius, 1767 1 1 1 1 1 LABRIDAE Labrus bimaculatus Linnaeus, 1758 1 1 1 1 1 1 LABRIDAE Labrus merula Linnaeus, 1758 1 1 1 LABRIDAE Labrus viridis Linnaeus, 1758 1 1 LABRIDAE Lachnolaimus maximus (Walbaum, 1792) 1 1 1 1 1 LABRIDAE Lappanella fasciata (Cocco, 1833) 1 1 1 1 LABRIDAE Lappanella guineensis Bauchot, 1969 1 LABRIDAE Stethojulis interrupta (Bleeker, 1851) 1 1 LABRIDAE Stethojulis strigiventer (Bennett, 1833) 1 1 LABRIDAE Symphodus bailloni (Valenciennes, 1839) 1 1 LABRIDAE Symphodus cinereus (Bonnaterre, 1788) 1 1 LABRIDAE Symphodus caeruleus (Azevedo, 1999) 1 LABRIDAE Symphodus doderleini Jordan, 1890 1 1 LABRIDAE Symphodus mediterraneus (Linnaeus, 1758) 1 1 1 1 1 1 LABRIDAE Symphodus melanocercus (Risso, 1810) 1 1 LABRIDAE Symphodus melops (Linnaeus, 1758) 1 1 1 LABRIDAE Symphodus ocellatus (Forsskål, 1775) 1 1 LABRIDAE Symphodus roissali (Risso, 1810) 1 1 LABRIDAE Symphodus rostratus (Bloch, 1791) 1 1 LABRIDAE Symphodus staitii (Nordmann, 1840) 1 LABRIDAE Symphodus tinca (Linnaeus, 1758) 1 1 1 LABRIDAE Symphodus trutta (Lowe, 1834) 1 1 1 LABRIDAE Thalassoma ascensionis (Quoy & Gaimard, 1834) 1 LABRIDAE Thalassoma bifasciatum (Bloch, 1791) 1 1 1 1 LABRIDAE Thalassoma hebraicum (Lacepède, 1801) 1 1 LABRIDAE Thalassoma lunare (Linnaeus, 1758) 1 1 LABRIDAE Thalassoma newtoni (Osório, 1891) 1 LABRIDAE Thalassoma noronhanum (Boulenger, 1890) 1 1 1 1 1 1 LABRIDAE Thalassoma pavo (Linnaeus, 1758) 1 1 1 1 1 1 1 1 LABRIDAE Thalassoma purpureum (Forsskål, 1775) 1 1 LABRIDAE Thalassoma sanctaehelenae (Valenciennes, 1839) 1 LABRIDAE Xyrichtys blanchardi (Cadenat & Marchal, 1963) 1 1 LABRIDAE Xyrichtys incandescens Edwards & Lubbock, 1981 1 1 1 LABRIDAE Xyrichtys martinicensis Valenciennes, 1840 1 1 1 1 1 LABRIDAE Xyrichtys novacula (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 LABRIDAE Xyrichtys sanctaehelenae (Günther, 1868) 1 1 1 LABRIDAE Xyrichtys splendens Castelnau, 1855 1 1 1 1 1 1 LABRISOMIDAE Labrisomus albigenys Beebe & Tee-Van, 1928 1 1 LABRISOMIDAE Labrisomus bucciferus (Poey, 1868) 1 1 LABRISOMIDAE Labrisomus cricota Sazima, Gasparini & Moura, 2002 1 1 LABRISOMIDAE Labrisomus filamentosus Springer, 1960 1 1 1 LABRISOMIDAE Labrisomus gobio (Valenciennes, 1836) 1 1 LABRISOMIDAE Labrisomus guppyi (Norman, 1922) 1 1 LABRISOMIDAE Labrisomus haitiensis Beebe & Tee-Van, 1928 1 1 1 LABRISOMIDAE Labrisomus kalisherae (Jordan, 1904) 1 1 1 1 1 1 LABRISOMIDAE Labrisomus nigricinctus Howell Rivero, 1936 1 1 LABRISOMIDAE Labrisomus nuchipinnis (Quoy & Gaimard, 1824) 1 1 1 1 1 1 1 1 1 1 1 1 1 LABRISOMIDAE Labrisomus sp.n. 1 1 LABRISOMIDAE Malacoctenus africanus Cadenat, 1951 1 1 LABRISOMIDAE Malacoctenus aurolineatus Smith, 1957 1 1 LABRISOMIDAE Malacoctenus boehlkei Springer, 1959 1 1 LABRISOMIDAE Malacoctenus delalandii (Valenciennes, 1836) 1 1 1 1 LABRISOMIDAE Malacoctenus erdmani Smith, 1957 1 1 LABRISOMIDAE Malacoctenus gilli (Steindachner, 1867) 1 1 LABRISOMIDAE Malacoctenus macropus (Poey, 1868) 1 1 1 LABRISOMIDAE Malacoctenus sp.1(aff.triangulatus) 1 1 LABRISOMIDAE Malacoctenus sp.2(Trindade) 1 LABRISOMIDAE Malacoctenus sp.3(St_Pauls) 1 LABRISOMIDAE Malacoctenus sp.4(Noronha) 1 1 LABRISOMIDAE Malacoctenus triangulatus Springer, 1959 1 1 1 LABRISOMIDAE Malacoctenus versicolor (Poey, 1876) 1 1 LABRISOMIDAE Nemaclinus atelestos Böhlke & Springer, 1975 1 1 1 LABRISOMIDAE Paraclinus arcanus Guimarães & Bacellar, 2002 1 1 LABRISOMIDAE Paraclinus barbatus Springer, 1955 1 1 LABRISOMIDAE Paraclinus cingulatus (Evermann & Marsh, 1899) 1 1 LABRISOMIDAE Paraclinus fasciatus (Steindachner, 1876) 1 1 1 LABRISOMIDAE Paraclinus grandicomis (Rosén, 1911) 1 1 LABRISOMIDAE Paraclinus infrons Böhlke, 1960 1 1 LABRISOMIDAE Paraclinus marmoratus (Steindachner, 1876) 1 1 LABRISOMIDAE Paraclinus naeorhegmis Böhlke, 1960 1 1 LABRISOMIDAE Paraclinus nigripinnis (Steindachner, 1867) 1 1 1 LABRISOMIDAE Paraclinus rubicundus (Starks, 1913) 1 1 LABRISOMIDAE Paraclinus spectator Guimarães & Bacellar, 2002 1 LABRISOMIDAE Starksia atlantica Longley, 1934 1 1 LABRISOMIDAE Starksia brasiliensis (Gilbert, 1900) 1 1 LABRISOMIDAE Starksia culebrae (Evermann & Marsh, 1899) 1 LABRISOMIDAE Starksia elongata Gilbert, 1971 1 1 LABRISOMIDAE Starksia fasciata (Longley, 1934) 1 LABRISOMIDAE Starksia guttata (Fowler, 1931) 1 1 LABRISOMIDAE Starksia hassi Klausewitz, 1958 1 1 LABRISOMIDAE Starksia lepicoelia Böhlke & Springer, 1961 1 1 LABRISOMIDAE Starksia leucovitta Williams & Mounts, 2003 1 LABRISOMIDAE Starksia melasma Williams & Mounts, 2003 1 LABRISOMIDAE Starksia multilepis Williams & Mounts, 2003 1 1 LABRISOMIDAE Starksia nanodes Böhlke & Springer, 1961 1 1 LABRISOMIDAE Starksia occidentalis Greenfield, 1979 1 LABRISOMIDAE Starksia ocellata (Steindachner, 1876) 1 1 1 LABRISOMIDAE Starksia rava Williams & Mounts, 2003 1 1 LABRISOMIDAE Starksia sella Williams & Mounts, 2003 1 1 LABRISOMIDAE Starksia sluiteri (Metzelaar, 1919) 1 1 LABRISOMIDAE Starksia smithvanizi Williams & Mounts, 2003 1 LABRISOMIDAE Starksia sp.n.5(Trindade) 1 LABRISOMIDAE Starksia sp.n.6(LA_Rocha) 1 LABRISOMIDAE Starksia starcki Gilbert, 1971 1 1 LABRISOMIDAE Starksia variabilis Greenfield, 1979 1 LABRISOMIDAE Starksia y-lineata Gilbert, 1965 1 LETHRINIDAE Lethrinus atlanticus Goode & Bean, 1879 1 1 1 LETHRINIDAE Lethrinus nebulosus Linnaeus, 1758 1 1 LUTJANIDAE Aphareus rutilans (Peters, 1877) 1 1 LUTJANIDAE Apsilus dentatus (Kaup, 1856) 1 1 1 LUTJANIDAE Apsilus fuscus Valenciennes, 1830 1 1 1 1 LUTJANIDAE Etelis oculatus (Valenciennes, 1828) 1 1 1 1 1 1 LUTJANIDAE Lutjanus agennes Bleeker, 1863 1 1 1 LUTJANIDAE Lutjanus alexandrei Moura & Lindeman, 2007 1 1 LUTJANIDAE Lutjanus analis (Cuvier, 1828) 1 1 1 1 1 1 LUTJANIDAE Lutjanus apodus (Walbaum, 1792) 1 1 1 1 1 LUTJANIDAE Lutjanus buccanella (Cuvier, 1828) 1 1 1 1 1 1 LUTJANIDAE Lutjanus campechanus (Poey, 1860) 1 1 1 LUTJANIDAE Lutjanus cyanopterus (Cuvier, 1828) 1 1 1 1 1 1 LUTJANIDAE Lutjanus dentatus (Duméril, 1861) 1 1 1 LUTJANIDAE Lutjanus endecacanthus Bleeker, 1863 1 1 LUTJANIDAE Lutjanus fulgens (Valenciennes, 1830) 1 1 1 1 LUTJANIDAE Lutjanus goreensis (Valenciennes, 1830) 1 1 1 1 1 1 LUTJANIDAE Lutjanus griseus (Linnaeus, 1758) 1 1 1 1 1 1 1 LUTJANIDAE Lutjanus jocu (Bloch & Schneider, 1801) 1 1 1 1 1 1 1 1 1 LUTJANIDAE Lutjanus mahogoni (Cuvier, 1828) 1 1 LUTJANIDAE Lutjanus purpureus Poey, 1876 1 1 1 1 1 1 LUTJANIDAE Lutjanus sanguineus (Cuvier, 1828) 1 1 LUTJANIDAE Lutjanus synagris (Linnaeus, 1758) 1 1 1 1 1 1 LUTJANIDAE Lutjanus vivanus (Cuvier, 1828) 1 1 1 1 1 1 1 LUTJANIDAE Ocyurus chrysurus (Bloch, 1791) 1 1 1 1 1 1 1 1 LUTJANIDAE Pristipomoides aquilonaris (Goode & Bean, 1896) 1 1 1 1 1 LUTJANIDAE Pristipomoides freemani Anderson, 1966 1 1 1 1 1 1 1 LUTJANIDAE Pristipomoides macrophthalmus (Müller & Troschel, 1848) 1 1 1 LUTJANIDAE Rhomboplites aurorubens (Cuvier, 1829) 1 1 1 1 1 1 1 MALACANTHIDAE Branchiostegus semifasciatus (Norman, 1931) 1 1 1 MALACANTHIDAE Caulolatilus bermudensis Dooley, 1981 1 MALACANTHIDAE Caulolatilus chrysops (Valenciennes, 1833) 1 1 1 1 1 MALACANTHIDAE Caulolatilus cyanops Poey, 1866 1 1 1 MALACANTHIDAE Caulolatilus dooleyi Berry, 1978 1 MALACANTHIDAE Caulolatilus guppyi Beebe & Tee-Van, 1937 1 MALACANTHIDAE Caulolatilus intermedius Howell Rivero, 1936 1 1 MALACANTHIDAE Caulolatilus microps Goode & Bean, 1878 1 1 1 MALACANTHIDAE Caulolatilus williamsi Dooley & Berry, 1977 1 MALACANTHIDAE Lopholatilus chamaeleonticeps Goode & Bean, 1879 1 1 1 MALACANTHIDAE Lopholatilus villarii Miranda-Ribeiro, 1915 1 1 MALACANTHIDAE Malacanthus plumieri (Bloch, 1786) 1 1 1 1 1 1 1 1 1 1 MICRODESMIDAE Cerdale fasciata Dawson, 1974 1 MICRODESMIDAE Cerdale floridana Longley, 1934 1 1 1 1 MICRODESMIDAE Microdesmus aethiopicus (Chabanaud, 1927) 1 1 MICRODESMIDAE Microdesmus africanus Dawson, 1979 1 MICRODESMIDAE Microdesmus bahianus Dawson, 1973 1 1 1 MICRODESMIDAE Microdesmus carri Gilbert, 1966 1 MICRODESMIDAE Microdesmus lanceolatus Dawson, 1962 1 MICRODESMIDAE Microdesmus longipinnis (Weymouth, 1910) 1 1 1 1 1 1 1 MICRODESMIDAE Microdesmus luscus Dawson, 1977 1 1 MONACANTHIDAE Aluterus heudelotii Hollard, 1855 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Aluterus monoceros (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Aluterus schoepfii (Walbaum, 1792) 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Aluterus scriptus (Osbeck, 1765) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Cantherhines dumerilii (Hollard, 1854) 1 1 MONACANTHIDAE Cantherhines fronticinctus (Günther, 1867) 1 1 MONACANTHIDAE Cantherhines macrocerus (Hollard, 1853) 1 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Cantherhines pardalis (Rüppell, 1837) 1 1 1 MONACANTHIDAE Cantherhines pullus (Ranzani, 1842) 1 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Monacanthus ciliatus (Mitchill, 1818) 1 1 1 1 1 1 1 1 MONACANTHIDAE Monacanthus tuckeri Bean, 1906 1 1 1 1 MONACANTHIDAE Pervagor janthinosoma (Bleeker, 1854) 1 1 MONACANTHIDAE Pseudalutarius nasicornis (Temminck & Schlegel, 1850) 1 1 MONACANTHIDAE Stephanolepis auratus (Castelnau, 1861) 1 1 MONACANTHIDAE Stephanolepis hispidus (Linnaeus, 1766) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MONACANTHIDAE Stephanolepis setifer (Bennett, 1831) 1 1 1 1 1 1 MONACANTHIDAE Thamnaconus modestoides (Barnard, 1927) 1 1 MORINGUIDAE Moringua edwardsi (Jordan & Bollman, 1889) 1 1 1 1 1 1 MORINGUIDAE Neoconger mucronatus Girard, 1858 1 1 1 1 MULLIDAE Mulloidichthys flavolineatus (Lacepède, 1801) 1 1 MULLIDAE Mulloidichthys martinicus (Cuvier, 1829) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MULLIDAE Mullus argentinae Hubbs & Marini, 1933 1 1 MULLIDAE Mullus auratus Jordan & Gilbert, 1882 1 1 1 MULLIDAE Mullus barbatus Linnaeus, 1758 1 1 1 MULLIDAE Mullus surmuletus (Peters, 1877) 1 1 1 1 1 1 1 MULLIDAE Parupeneus barberinus (Kaup, 1856) 1 1 MULLIDAE Parupeneus indicus (Shaw, 1803) 1 1 MULLIDAE Parupeneus rubescens (Lacepède, 1801) 1 1 MULLIDAE Pseudupeneus maculatus (Bloch, 1793) 1 1 1 1 1 1 1 1 1 MULLIDAE Pseudupeneus prayensis (Cuvier, 1829) 1 1 1 1 1 1 MULLIDAE Upeneus parvus Poey, 1852 1 1 1 1 1 MURAENIDAE Anarchias longicaudis (Peters, 1877) 1 1 1 1 1 MURAENIDAE Anarchias similis (Lea, 1913) 1 1 1 1 1 1 1 MURAENIDAE Channomuraena vittata (Richardson, 1845) 1 1 1 1 1 1 1 1 1 MURAENIDAE Echidna catenata (Bloch, 1795) 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Echidna peli (Kaup, 1856) 1 1 1 1 MURAENIDAE Enchelycore anatina (Lowe, 1838) 1 1 1 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Enchelycore carychroa Böhlke & Böhlke, 1976 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Enchelycore nigricans (Bonnaterre, 1788) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax afer Bloch, 1795 1 1 1 1 MURAENIDAE Gymnothorax bacalladoi Böhlke & Brito, 1987 1 1 1 MURAENIDAE Gymnothorax conspersus Poey, 1867 1 1 1 1 1 MURAENIDAE Gymnothorax favagineus Bloch & Schneider, 1801 1 1 MURAENIDAE Gymnothorax funebris Ranzani, 1840 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax hubbsi Böhlke & Böhlke, 1977 1 1 1 MURAENIDAE Gymnothorax johnsoni (Smith, 1962) 1 1 MURAENIDAE Gymnothorax kolpos Böhlke & Böhlke, 1980 1 1 1 MURAENIDAE Gymnothorax maderensis (Johnson, 1862) 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax mareei Poll, 1953 1 1 1 MURAENIDAE Gymnothorax meleagris (Shaw, 1795) 1 1 MURAENIDAE Gymnothorax miliaris (Kaup, 1856) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax moringa (Cuvier, 1829) 1 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax nigromarginatus (Girard, 1858) 1 1 MURAENIDAE Gymnothorax ocellatus Agassiz, 1831 1 1 1 1 1 MURAENIDAE Gymnothorax polygonius Poey, 1875 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax saxicola Jordan & Davis, 1891 1 1 1 1 MURAENIDAE Gymnothorax undulatus (Lacepède, 1803) 1 1 MURAENIDAE Gymnothorax unicolor (Delaroche, 1809) 1 1 1 1 1 1 1 1 MURAENIDAE Gymnothorax vicinus (Castelnau, 1855) 1 1 1 1 1 1 1 1 1 1 1 1 1 MURAENIDAE Monopenchelys acuta (Parr, 1930) 1 1 1 1 MURAENIDAE Muraena augusti (Kaup, 1856) 1 1 1 1 MURAENIDAE Muraena helena Linnaeus, 1758 1 1 1 1 1 1 1 1 MURAENIDAE Muraena melanotis (Kaup, 1860) 1 1 1 1 1 MURAENIDAE Muraena pavonina Richardson, 1845 1 1 1 1 1 MURAENIDAE Muraena retifera Goode & Bean, 1882 1 1 1 1 MURAENIDAE Muraena robusta Osório, 1911 1 1 1 1 1 1 MURAENIDAE Uropterygius macularius (Lesueur, 1825) 1 1 1 1 1 1 1 1 MURAENIDAE Uropterygius wheeleri Blache, 1967 1 1 1 1 MYROCONGRIDAE Myroconger compressus Günther, 1870 1 1 1 1 OGCOCEPHALIDAE Halieutaea fitzsimonsi (Gilchrist & Thompson, 1916) 1 1 OGCOCEPHALIDAE Halieutichthys aculeatus (Mitchill, 1818) 1 1 1 1 1 OGCOCEPHALIDAE Ogcocephalus corniger Bradbury, 1980 1 1 1 OGCOCEPHALIDAE Ogcocephalus cubifrons (Richardson, 1836) 1 OGCOCEPHALIDAE Ogcocephalus declivirostris Bradbury, 1980 1 1 1 OGCOCEPHALIDAE Ogcocephalus nasutus (Cuvier, 1829) 1 1 1 OGCOCEPHALIDAE Ogcocephalus notatus (Valenciennes, 1837) 1 1 1 1 OGCOCEPHALIDAE Ogcocephalus pantostictus Bradbury, 1980 1 1 OGCOCEPHALIDAE Ogcocephalus parvus Longley & Hildebrand, 1940 1 1 1 1 OGCOCEPHALIDAE Ogcocephalus pumilus Bradbury, 1980 1 1 OGCOCEPHALIDAE Ogcocephalus radiatus (Mitchill, 1818) 1 1 1 OGCOCEPHALIDAE Ogcocephalus rostellum Bradbury, 1980 1 1 1 OGCOCEPHALIDAE Ogcocephalus vespertilio (Linnaeus, 1758) 1 1 1 1 OPHICHTHIDAE Ahlia egmontis (Jordan, 1884) 1 1 1 1 1 1 OPHICHTHIDAE Aplatophis chauliodus Böhlke, 1956 1 1 OPHICHTHIDAE Aprognathodon platyventris Böhlke, 1967 1 1 OPHICHTHIDAE Apterichtus anguiformis (Peters, 1877) 1 1 1 1 OPHICHTHIDAE Apterichtus ansp (Böhlke, 1968) 1 1 1 1 1 OPHICHTHIDAE Apterichtus caecus (Linnaeus, 1758) 1 1 1 1 1 OPHICHTHIDAE Apterichtus gracilis (Kaup, 1856) 1 OPHICHTHIDAE Apterichtus kendalli (Gilbert, 1891) 1 1 1 OPHICHTHIDAE Bascanichthys bascanium (Jordan, 1884) 1 1 1 OPHICHTHIDAE Bascanichthys ceciliae Blache & Cadenat, 1971 1 OPHICHTHIDAE Bascanichthys inopinatus McCosker, Böhlke & Böhlke, 1989 1 OPHICHTHIDAE Bascanichthys paulensis Storey, 1939 1 1 1 OPHICHTHIDAE Bascanichthys scuticaris (Goode & Bean, 1880) 1 1 1 OPHICHTHIDAE Callechelys bilinearis Kanazawa, 1952 1 1 1 1 OPHICHTHIDAE Callechelys guineensis (Osório, 1893) 1 1 1 1 1 OPHICHTHIDAE Callechelys leucoptera (Cadenat, 1954) 1 OPHICHTHIDAE Callechelys muraena Jordan & Evermann, 1887 1 1 OPHICHTHIDAE Callechelys springeri (Ginsburg, 1951) 1 1 OPHICHTHIDAE Caralophia loxochila Böhlke, 1955 1 1 1 OPHICHTHIDAE Echelus myrus (Linnaeus, 1758) 1 1 1 1 1 OPHICHTHIDAE Echiophis foresti (Cadenat & Roux, 1964) 1 OPHICHTHIDAE Echiophis intertinctus (Richardson, 1848) 1 1 1 1 OPHICHTHIDAE Echiophis punctifer (Kaup, 1860) 1 1 1 1 1 OPHICHTHIDAE Ethadophis akkistikos McCosker & Böhlke, 1984 1 OPHICHTHIDAE Gordiichthys ergodes McCosker, Böhlke & Böhlke, 1989 1 OPHICHTHIDAE Gordiichthys irretitus Jordan & Davis, 1891 1 1 OPHICHTHIDAE Gordiichthys randalli McCosker & Böhlke, 1984 1 OPHICHTHIDAE Hemerorhinus opici Blache & Bauchot, 1972 1 OPHICHTHIDAE Herpetiochthys regius (Richardson, 1848) 1 1 OPHICHTHIDAE Ichthyapus insularis McCosker, 2004 1 OPHICHTHIDAE Ichthyapus ophioneus (Bleeker, 1857) 1 1 1 1 1 1 1 1 OPHICHTHIDAE Leiuranus semicinctus (Lay & Bennett, 1839) 1 1 OPHICHTHIDAE Letharchus aliculatus McCosker, 1974 1 OPHICHTHIDAE Letharchus velifer Goode & Bean, 1882 1 1 OPHICHTHIDAE Myrichthys breviceps (Richardson, 1848) 1 1 1 1 1 1 1 1 OPHICHTHIDAE Myrichthys ocellatus (Lesueur, 1825) 1 1 1 1 1 1 1 1 OPHICHTHIDAE Myrichthys pardalis (Valenciennes, 1839) 1 1 1 1 OPHICHTHIDAE Myrophis anterodorsalis McCosker, Böhlke & Böhlke, 1989 1 OPHICHTHIDAE Myrophis platyrhynchus Breder, 1927 1 1 1 1 OPHICHTHIDAE Mystriophis crosnieri Blache, 1971 1 1 1 1 OPHICHTHIDAE Mystriophis rostellatus (Richardson, 1848) 1 1 1 OPHICHTHIDAE Ophichthus apicalis (Anonymous [Bennett], 1830) 1 1 OPHICHTHIDAE Ophichthus brasiliensis (Kaup, 1856) 1 OPHICHTHIDAE Ophichthus cruentifer (Goode & Bean, 1896) 1 1 OPHICHTHIDAE Ophichthus cylindroideus (Ranzani, 1840) 1 1 1 1 OPHICHTHIDAE Ophichthus gomesii (Castelnau, 1855) 1 1 1 1 1 1 OPHICHTHIDAE Ophichthus hyposagmatus McCosker & Böhlke, 1984 1 1 OPHICHTHIDAE Ophichthus melanoporus Kanazawa, 1963 1 1 OPHICHTHIDAE Ophichthus menezesi McCosker & Böhlke, 1984 1 1 OPHICHTHIDAE Ophichthus ophis (Linnaeus, 1758) 1 1 1 1 1 1 1 1 OPHICHTHIDAE Ophichthus puncticeps (Kaup, 1860) 1 1 1 OPHICHTHIDAE Ophichthus regius (Richardson, 1848) 1 1 OPHICHTHIDAE Ophichthus rex Böhlke & Caruso, 1980 1 OPHICHTHIDAE Ophichthus rufus (Rafinesque, 1810) 1 1 1 OPHICHTHIDAE Ophichthus unicolor Regan, 1908 1 1 OPHICHTHIDAE Ophisurus serpens (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 OPHICHTHIDAE Phaenomonas longissima (Cadenat & Marchal, 1963) 1 1 1 OPHICHTHIDAE Pisodonophis boro (Hamilton, 1822) 1 1 OPHICHTHIDAE Pisodonophis cancrivorus (Richardson, 1848) 1 1 OPHICHTHIDAE Pisodonophis semicinctus (Richardson, 1848) 1 1 1 1 1 OPHICHTHIDAE Pseudomyrophis atlanticus Blache, 1975 1 OPHICHTHIDAE Quassiremus ascensionis (Studer, 1889) 1 1 1 1 1 OPHICHTHIDAE Scolecenchelys gymnotus (Bleeker, 1857) 1 1 OPHICHTHIDAE Scolecenchelys laticaudata (Ogilby, 1897) 1 1 OPHICHTHIDAE Asarcenchelys longimanus McCosker, 1985 1 OPHICHTHIDAE Brachysomophis atlanticus Blache & Saldanha, 1972 1 OPHIDIIDAE Brotula barbata (Bloch & Schneider, 1801) 1 1 1 1 1 1 1 1 OPHIDIIDAE Brotula sp.n. 1 OPHIDIIDAE Genypterus blacodes (Forster, 1801) 1 1 OPHIDIIDAE Genypterus brasiliensis Regan, 1903 1 1 OPHIDIIDAE Ophidion barbatum Linnaeus, 1758 1 1 1 OPHIDIIDAE Ophidion beani Jordan & Gilbert, 1883 1 1 OPHIDIIDAE Ophidion grayi (Fowler, 1948) 1 1 OPHIDIIDAE Ophidion holbrookii (Putnam, 1874) 1 1 1 1 OPHIDIIDAE Ophidion lagochila (Böhlke & Robins, 1959) 1 1 OPHIDIIDAE Ophidion lozanoi Matallanas, 1990 1 1 OPHIDIIDAE Ophidion marginatum (DeKay, 1842) 1 1 OPHIDIIDAE Ophidion nocomis Robins & Böhlke, 1959 1 OPHIDIIDAE Ophidion robinsi Fahay, 1992 1 OPHIDIIDAE Ophidion saldanhai Matallanas & Brito, 1999 1 1 OPHIDIIDAE Ophidion selenops Robins & Böhlke, 1959 1 1 OPHIDIIDAE Ophidion sp. 1 OPHIDIIDAE Ophidion welshi (Nichols & Breder, 1922) 1 1 OPHIDIIDAE Otophidium chickcharney Böhlke & Robins, 1959 1 1 1 OPHIDIIDAE Otophidium dormitator Böhlke & Robins, 1959 1 1 1 OPHIDIIDAE Otophidium omostigmum (Jordan & Gilbert, 1882) 1 1 1 OPHIDIIDAE Parophidion schmidti (Miranda Ribeiro, 1903) 1 1 1 1 OPHIDIIDAE Parophidion vassali (Risso, 1810) 1 1 1 1 OPHIDIIDAE Petrotyx sanguineus (Meek & Hildebrand, 1928) 1 1 OPHIDIIDAE Raneya fluminensis (Miranda Ribeiro, 1903) 1 1 OPISTOGNATHIDAE Lonchopisthus higmani Mead, 1959 1 1 OPISTOGNATHIDAE Lonchopisthus lemur (Myers, 1935) 1 1 1 OPISTOGNATHIDAE Lonchopisthus micrognathus (Poey, 1860) 1 1 OPISTOGNATHIDAE Lonchopisthus sp.n. 1 1 OPISTOGNATHIDAE Opistognathus aurifrons (Jordan & Thompson, 1905) 1 1 1 1 OPISTOGNATHIDAE Opistognathus brasiliensis Smith-Vaniz, 1997 1 OPISTOGNATHIDAE Opistognathus cuvieri Valenciennes, 1836 1 1 OPISTOGNATHIDAE Opistognathus gilberti Böhlke, 1967 1 1 OPISTOGNATHIDAE Opistognathus leprocarus Smith-Vaniz, 1997 1 OPISTOGNATHIDAE Opistognathus lonchurus Jordan & Gilbert, 1882 1 1 1 1 OPISTOGNATHIDAE Opistognathus macrognathus Poey, 1860 1 1 1 1 OPISTOGNATHIDAE Opistognathus maxillosus Poey, 1860 1 1 OPISTOGNATHIDAE Opistognathus megalepis Smith-Vaniz, 1972 1 OPISTOGNATHIDAE Opistognathus melachasme Smith-Vaniz, 1972 1 OPISTOGNATHIDAE Opistognathus nothus Smith-Vaniz, 1997 1 1 1 OPISTOGNATHIDAE Opistognathus robinsi Smith-Vaniz, 1997 1 1 1 OPISTOGNATHIDAE Opistognathus signatus Smith-Vaniz, 1997 1 1 OPISTOGNATHIDAE Opistognathus sp.n.(aff.aurifrons) 1 1 1 1 OPISTOGNATHIDAE Opistognathus whitehursti (Longley, 1927) 1 1 1 1 OSTRACIIDAE Acanthostracion guineensis (Bleeker, 1865) 1 1 OSTRACIIDAE Acanthostracion notacanthus (Bleeker, 1863) 1 1 1 1 1 OSTRACIIDAE Acanthostracion polygonius Poey, 1876 1 1 1 1 1 1 1 1 1 OSTRACIIDAE Acanthostracion quadricornis (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 OSTRACIIDAE Anoplocapros robustus (Fraser-Brunner, 1941) 1 OSTRACIIDAE Lactophrys bicaudalis Linnaeus, 1758 1 1 1 1 OSTRACIIDAE Lactophrys trigonus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 OSTRACIIDAE Lactophrys triqueter (Linnaeus, 1758) 1 1 1 1 1 1 OSTRACIIDAE Lactoria cornuta (Linnaeus, 1758) 1 1 OSTRACIIDAE Lactoria diaphana (Bloch & Schneider, 1801) 1 1 OSTRACIIDAE Lactoria fornasini (Bianconi, 1846) 1 1 OSTRACIIDAE Ostracion cubicus Linnaeus, 1758 1 1 OSTRACIIDAE Ostracion meleagris Shaw, 1796 1 1 OSTRACIIDAE Tetrosomus concatenatus (Bloch, 1785) 1 1 PARALICHTHYIDAE Ancylopsetta cycloidea Tyler, 1959 1 1 PARALICHTHYIDAE Ancylopsetta dilecta (Goode & Bean, 1883) 1 1 PARALICHTHYIDAE Ancylopsetta kumperae Tyler, 1959 1 1 PARALICHTHYIDAE Ancylopsetta microctenus Gutherz, 1966 1 PARALICHTHYIDAE Ancylopsetta quadrocellata Gill, 1864 1 1 PARALICHTHYIDAE Citharichthys arctifrons (Mitchill, 1815) 1 1 PARALICHTHYIDAE Citharichthys arenaceus Evermann & Marsh, 1900 1 1 1 1 1 PARALICHTHYIDAE Citharichthys cornutus (Günther, 1880) 1 1 1 1 1 1 PARALICHTHYIDAE Citharichthys dinoceros Goode & Bean, 1886 1 1 1 1 PARALICHTHYIDAE Citharichthys gymnorhinus Gutherz & Blackman, 1970 1 1 1 PARALICHTHYIDAE Citharichthys macrops Dresel, 1885 1 1 1 1 1 PARALICHTHYIDAE Citharichthys minutus Cervigón, 1982 1 1 PARALICHTHYIDAE Citharichthys spilopterus Günther, 1862 1 1 1 1 1 PARALICHTHYIDAE Citharichthys stampflii (Steindachner, 1894) 1 1 1 PARALICHTHYIDAE Citharichthys valdezi Cervigón, 1986 1 1 PARALICHTHYIDAE Cyclopsetta chittendeni Bean, 1895 1 1 1 1 PARALICHTHYIDAE Cyclopsetta decussata Gunter, 1946 1 PARALICHTHYIDAE Cyclopsetta fimbriata (Goode & Bean, 1885) 1 1 1 1 1 PARALICHTHYIDAE Etropus crossotus Jordan & Gilbert, 1882 1 1 1 1 1 PARALICHTHYIDAE Etropus cyclosquamus Leslie & Stewart, 1986 1 PARALICHTHYIDAE Etropus intermedius Norman, 1933 1 1 1 PARALICHTHYIDAE Etropus longimanus Norman, 1933 1 1 PARALICHTHYIDAE Etropus microstomus (Gill, 1864) 1 1 PARALICHTHYIDAE Etropus rimosus Goode & Bean, 1885 1 1 PARALICHTHYIDAE Gastropsetta frontalis Bean, 1895 1 1 1 PARALICHTHYIDAE Paralichthys albigutta Jordan & Gilbert, 1882 1 1 1 PARALICHTHYIDAE Paralichthys brasiliensis (Ranzani, 1842) 1 1 1 PARALICHTHYIDAE Paralichthys dentatus (Linnaeus, 1766) 1 PARALICHTHYIDAE Paralichthys isosceles Jordan, 1891 1 1 PARALICHTHYIDAE Paralichthys lethostigma Jordan & Gilbert, 1884 1 1 PARALICHTHYIDAE Paralichthys oblongus (Mitchill, 1815) 1 1 PARALICHTHYIDAE Paralichthys orbignyanus (Valenciennes, 1842) 1 1 PARALICHTHYIDAE Paralichthys patagonicus Jordan, 1889 1 1 PARALICHTHYIDAE Paralichthys squamilentus Jordan & Gilbert, 1882 1 1 PARALICHTHYIDAE Paralichthys triocellatus Miranda-Ribeiro, 1903 1 1 PARALICHTHYIDAE Paralichthys tropicus Ginsburg, 1933 1 1 PARALICHTHYIDAE Pseudorhombus elevatus Ogilby, 1912 1 1 PARALICHTHYIDAE Syacium guineensis (Bleeker, 1862) 1 1 1 1 1 PARALICHTHYIDAE Syacium gunteri Ginsburg, 1933 1 1 1 PARALICHTHYIDAE Syacium micrurum Ranzani, 1842 1 1 1 1 PARALICHTHYIDAE Syacium papillosum (Linnaeus, 1758) 1 1 1 1 1 1 1 PARALICHTHYIDAE Xystreurys rasile (Jordan, 1891) 1 1 PEMPHERIDAE Pempheris poeyi Bean, 1885 1 1 1 PEMPHERIDAE Pempheris schomburgki Müller & Troschel, 1848 1 1 1 1 1 1 1 PINGUIPEDIDAE Parapercis atlantica (Vaillant, 1887) 1 PINGUIPEDIDAE Parapercis fasciatus? (Jenyns, 1840) 1 PINGUIPEDIDAE Parapercis maritzi Anderson, 1992 1 1 PINGUIPEDIDAE Parapercis multifasciata Döderlein, 1884 1 1 1 PINGUIPEDIDAE Parapercis robinsoni Fowler, 1929 1 1 PINGUIPEDIDAE Pinguipes brasilianus Cuvier, 1829 1 1 PINGUIPEDIDAE Pseudopercis numida Miranda-Ribeiro, 1903 1 1 1 PINGUIPEDIDAE Pseudopercis semifasciata (Cuvier, 1829) 1 1 PLATYCEPHALIDAE Platycephalus indicus (Linnaeus, 1758) 1 1 PLATYCEPHALIDAE Solitas gruveli (Pellegrin, 1905) 1 1 POMACANTHIDAE Centropyge acanthops (Norman, 1922) 1 1 POMACANTHIDAE Centropyge argi Woods & Kanazawa, 1951 1 1 1 1 POMACANTHIDAE Centropyge aurantonotus Burgess, 1974 1 1 1 1 1 1 1 POMACANTHIDAE Centropyge resplendens Lubbock & Sankey, 1975 1 POMACANTHIDAE Holacanthus africanus Cadenat, 1951 1 1 1 1 POMACANTHIDAE Holacanthus bermudensis Goode, 1876 1 1 1 1 POMACANTHIDAE Holacanthus ciliaris (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 POMACANTHIDAE Holacanthus tricolor (Bloch, 1795) 1 1 1 1 1 1 1 1 POMACANTHIDAE Pomacanthus arcuatus (Linnaeus, 1758) 1 1 1 1 1 1 POMACANTHIDAE Pomacanthus paru (Bloch, 1787) 1 1 1 1 1 1 1 1 1 1 POMACANTHIDAE Pomacanthus rhomboides (Gilchrist & Thompson, 1908) 1 1 POMACANTHIDAE Pomacanthus semicirculatus (Cuvier, 1831) 1 1 POMACENTRIDAE Abudefduf hoefleri (Steindachner, 1881) 1 1 POMACENTRIDAE Abudefduf luridus (Cuvier, 1830) 1 1 1 1 POMACENTRIDAE Abudefduf saxatilis (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 POMACENTRIDAE Abudefduf sordidus (Forsskål, 1775) 1 1 POMACENTRIDAE Abudefduf taurus (Müller & Troschel, 1848) 1 1 1 1 1 1 1 POMACENTRIDAE Abudefduf vaigiensis (Quoy & Gaimard, 1825) 1 1 POMACENTRIDAE Chromis bermudae Nichols, 1920 1 POMACENTRIDAE Chromis cadenati Whitley, 1951 1 1 POMACENTRIDAE Chromis chromis (Linnaeus, 1758) 1 1 POMACENTRIDAE Chromis cyanea (Poey, 1860) 1 1 1 POMACENTRIDAE Chromis enchrysura Jordan & Gilbert, 1882 1 1 1 1 1 1 1 POMACENTRIDAE Chromis flavicauda (Günther, 1880) 1 1 1 1 POMACENTRIDAE Chromis insolata (Cuvier, 1830) 1 1 1 1 POMACENTRIDAE Chromis jubauna Moura, 1995 1 1 1 1 POMACENTRIDAE Chromis limbata (Valenciennes, 1833) 1 1 1 1 1 POMACENTRIDAE Chromis lubbocki Edwards, 1986 1 POMACENTRIDAE Chromis multilineata (Guichenot, 1853) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 POMACENTRIDAE Chromis sanctaehelenae Edwards, 1987 1 POMACENTRIDAE Chromis scotti Emery, 1968 1 1 1 1 1 POMACENTRIDAE Dascyllus trimaculatus (Rüppell, 1829) 1 1 POMACENTRIDAE Microspathodon chrysurus (Cuvier, 1830) 1 1 1 1 1 1 1 POMACENTRIDAE Microspathodon frontatus Emery, 1970 1 1 POMACENTRIDAE Similiparma hermani (Steindachner, 1887) 1 POMACENTRIDAE Stegastes adustus (Troschel, 1865) 1 1 1 1 POMACENTRIDAE Stegastes diencaeus (Jordan & Rutter, 1897) 1 1 POMACENTRIDAE Stegastes fuscus (Cuvier, 1830) 1 1 1 POMACENTRIDAE Stegastes imbricatus Jenyns, 1840 1 1 1 1 POMACENTRIDAE Stegastes leucostictus (Müller & Troschel, 1848) 1 1 1 1 POMACENTRIDAE Stegastes lubbocki Allen & Smith, 1992 1 POMACENTRIDAE Stegastes otophorus (Poey, 1860) 1 POMACENTRIDAE Stegastes partitus (Poey, 1868) 1 1 1 1 POMACENTRIDAE Stegastes pictus (Castelnau, 1855) 1 1 1 1 1 1 POMACENTRIDAE Stegastes planifrons (Cuvier, 1830) 1 1 1 1 POMACENTRIDAE Stegastes rocasensis (Emery, 1972) 1 1 1 POMACENTRIDAE Stegastes sanctaehelenae (Sauvage, 1879) 1 POMACENTRIDAE Stegastes sanctipauli Lubbock & Edwards, 1981 1 1 POMACENTRIDAE Stegastes sp.(aff_variabilis) 1 1 1 1 1 POMACENTRIDAE Stegastes uenfi Novelli, Nunan & Lima, 2000 1 1 POMACENTRIDAE Stegastes variabilis (Castelnau, 1855) 1 1 PRIACANTHIDAE Cookeolus japonicus (Cuvier, 1829) 1 1 1 1 1 1 1 1 1 1 1 PRIACANTHIDAE Heteropriacanthus cruentatus (Lacepède, 1801) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 PRIACANTHIDAE Priacanthus arenatus Cuvier, 1829 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 PRIACANTHIDAE Priacanthus hamrur (Forsskål, 1775) 1 1 PRIACANTHIDAE Pristigenys alta (Gill, 1862) 1 1 1 1 1 1 PRIACANTHIDAE Pristigenys niphonia (Cuvier, 1829) 1 1 PSEUDOCHROMIDAEHalidesmus scapularis Günther, 1872 1 1 PTERELEOTRIDAE Ptereleotris calliura (Jordan & Gilbert, 1882) 1 1 PTERELEOTRIDAE Ptereleotris helenae (Randall, 1967) 1 1 PTERELEOTRIDAE Ptereleotris randalli Gasparini, Rocha & Floeter, 2001 1 1 1 1 SCARIDAE Cryptotomus roseus Cope, 1871 1 1 1 1 1 1 1 1 1 SCARIDAE Nicholsina usta_collettei Schultz, 1968 1 1 1 SCARIDAE Nicholsina usta_usta (Valenciennes, 1840) 1 1 1 1 1 SCARIDAE Scarus coelestinus Valenciennes, 1840 1 1 1 1 SCARIDAE Scarus coeruleus (Bloch, 1786) 1 1 1 1 SCARIDAE Scarus ghobban Forsskål, 1775 1 1 SCARIDAE Scarus guacamaia Cuvier, 1829 1 1 1 1 1 1 SCARIDAE Scarus hoefleri (Steindachner, 1881) 1 1 1 1 SCARIDAE Scarus iserti (Bloch, 1789) 1 1 1 1 SCARIDAE Scarus taeniopterus Desmarest, 1831 1 1 1 1 SCARIDAE Scarus trispinosus Valenciennes, 1840 1 1 SCARIDAE Scarus vetula Bloch & Schneider, 1801 1 1 1 SCARIDAE Scarus zelindae Moura, Figueiredo & Sazima, 2001 1 1 SCARIDAE Sparisoma amplum (Ranzani, 1842) 1 1 1 1 1 1 SCARIDAE Sparisoma atomarium (Poey, 1861) 1 1 1 1 SCARIDAE Sparisoma aurofrenatum (Valenciennes, 1840) 1 1 1 SCARIDAE Sparisoma axillare (Steindachner, 1878) 1 1 1 1 1 1 SCARIDAE Sparisoma chrysopterum (Bloch & Schneider, 1801) 1 1 1 1 SCARIDAE Sparisoma cretense (Linnaeus, 1758) 1 1 1 1 1 1 1 1 SCARIDAE Sparisoma frondosum (Agassiz, 1831) 1 1 1 1 1 SCARIDAE Sparisoma griseorubra Cervigón, 1982 1 SCARIDAE Sparisoma radians (Valenciennes, 1840) 1 1 1 1 1 1 SCARIDAE Sparisoma rubripinne (Valenciennes, 1840) 1 1 1 1 SCARIDAE Sparisoma sp.(aff_rubripinne) 1 1 1 SCARIDAE Sparisoma strigatum (Günther, 1862) 1 1 SCARIDAE Sparisoma tuiupiranga Gasparini, Joyeux & Floeter, 2003 1 1 1 SCARIDAE Sparisoma viride (Bonnaterre, 1788) 1 1 1 1 SCIAENIDAE Argyrosomus regius (Asso, 1801) 1 1 1 1 1 SCIAENIDAE Equetus lanceolatus (Linnaeus, 1758) 1 1 1 1 1 1 1 SCIAENIDAE Equetus punctatus (Bloch & Schneider, 1801) 1 1 1 SCIAENIDAE Odontoscion dentex (Cuvier, 1830) 1 1 1 1 1 SCIAENIDAE Pareques acuminatus (Bloch & Schneider, 1801) 1 1 1 1 1 SCIAENIDAE Pareques iwamotoi Miller & Woods, 1988 1 1 1 1 SCIAENIDAE Pareques umbrosus (Gmelin, 1789) 1 1 1 1 1 SCIAENIDAE Sciaena umbra (Schultz, 1958) 1 1 1 1 1 1 SCIAENIDAE Umbrina canariensis (Rafinesque, 1810) 1 1 1 1 1 1 1 1 1 SCIAENIDAE Umbrina cirrosa Cuvier, 1829 1 1 1 1 SCIAENIDAE Umbrina ronchus (Longley, 1935) 1 1 1 1 1 1 1 1 1 SCIAENIDAE Umbrina steindachneri (Ginsburg, 1952) 1 SCORPAENIDAE Neomerinthe beanorum (Jordan & Swain, 1885) 1 1 SCORPAENIDAE Neomerinthe folgori Günther, 1868 1 1 1 SCORPAENIDAE Neomerinthe hemingwayi Fowler, 1935 1 1 1 SCORPAENIDAE Phenacoscorpius nebris Eschmeyer, 1965 1 SCORPAENIDAE Pontinus accraensis Norman, 1935 1 1 1 1 SCORPAENIDAE Pontinus castor Poey, 1860 1 1 1 SCORPAENIDAE Pontinus corallinus Miranda-Ribeiro, 1903 1 SCORPAENIDAE Pontinus kuhlii (Bowdich, 1825) 1 1 1 1 1 1 1 SCORPAENIDAE Pontinus leda Eschmeyer, 1969 1 1 SCORPAENIDAE Pontinus longispinis Goode & Bean, 1896 1 1 1 1 1 SCORPAENIDAE Pontinus nematophthalmus (Günther, 1860) 1 1 1 SCORPAENIDAE Pontinus nigropunctatus (Günther, 1868) 1 SCORPAENIDAE Pontinus rathbuni Goode & Bean, 1896 1 1 1 1 1 SCORPAENIDAE Pterois miles (Bennett, 1828) 1 1 SCORPAENIDAE Scorpaena agassizii Goode & Bean, 1896 1 1 1 1 SCORPAENIDAE Scorpaena albifimbria Evermann & Marsh, 1900 1 1 1 SCORPAENIDAE Scorpaena angolensis Norman, 1935 1 1 1 1 SCORPAENIDAE Scorpaena annobonae Eschmeyer, 1969 1 SCORPAENIDAE Scorpaena ascensionis Eschmeyer, 1971 1 SCORPAENIDAE Scorpaena azorica Eschmeyer, 1969 1 1 SCORPAENIDAE Scorpaena bergii Evermann & Marsh, 1900 1 1 1 1 SCORPAENIDAE Scorpaena brachyptera Eschmeyer, 1965 1 1 SCORPAENIDAE Scorpaena brasiliensis Cuvier, 1829 1 1 1 1 1 1 SCORPAENIDAE Scorpaena calcarata Goode & Bean, 1882 1 1 1 1 1 SCORPAENIDAE Scorpaena canariensis (Sauvage, 1878) 1 1 1 SCORPAENIDAE Scorpaena dispar Longley & Hildebrand, 1940 1 1 1 1 1 SCORPAENIDAE Scorpaena elachys Eschmeyer, 1965 1 1 SCORPAENIDAE Scorpaena elongata Cadenat, 1943 1 1 1 1 1 SCORPAENIDAE Scorpaena grandicornis Cuvier, 1829 1 1 1 1 1 1 SCORPAENIDAE Scorpaena inermis Cuvier, 1829 1 1 1 SCORPAENIDAE Scorpaena isthmensis Meek & Hildebrand, 1928 1 1 1 1 1 1 SCORPAENIDAE Scorpaena laevis Troschel, 1866 1 1 1 1 SCORPAENIDAE Scorpaena loppei Cadenat, 1943 1 1 1 SCORPAENIDAE Scorpaena maderensis Valenciennes, 1833 1 1 1 1 1 1 1 SCORPAENIDAE Scorpaena melasma Eschmeyer, 1965 1 SCORPAENIDAE Scorpaena mellissii Günther, 1868 1 SCORPAENIDAE Scorpaena normani Cadenat, 1943 1 1 1 SCORPAENIDAE Scorpaena notata Rafinesque, 1810 1 1 1 1 1 1 1 1 SCORPAENIDAE Scorpaena petricola Eschmeyer, 1965 1 SCORPAENIDAE Scorpaena plumieri Bloch, 1789 1 1 1 1 1 1 1 1 1 SCORPAENIDAE Scorpaena porcus Linnaeus, 1758 1 1 1 1 1 1 SCORPAENIDAE Scorpaena scrofa Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 SCORPAENIDAE Scorpaena stephanica Cadenat, 1943 1 1 1 SCORPAENIDAE Scorpaenodes africanus Pfaff, 1933 1 1 SCORPAENIDAE Scorpaenodes arenai Torchio, 1962 1 1 SCORPAENIDAE Scorpaenodes caribbaeus Meek & Hildebrand, 1928 1 1 1 1 1 SCORPAENIDAE Scorpaenodes elongatus Cadenat, 1950 1 SCORPAENIDAE Scorpaenodes insularis Eschmeyer, 1971 1 1 1 SCORPAENIDAE Scorpaenodes tredecimspinosus (Metzelaar, 1919) 1 1 1 1 1 SCORPAENIDAE Sebastapistes cyanostigma (Bleeker, 1856) 1 1 SCORPAENIDAE Sebastes capensis (Gmelin, 1789) 1 1 SERRANIDAE Acanthistius brasilianus (Cuvier, 1828) 1 1 SERRANIDAE Acanthistius patachonicus (Jenyns, 1840) 1 1 1 SERRANIDAE Acanthistius sebastoides (Castelnau, 1861) 1 1 SERRANIDAE Alphestes afer (Bloch, 1793) 1 1 1 1 1 1 1 1 SERRANIDAE Anthias anthias (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Anthias asperilinguis Günther, 1859 1 1 1 SERRANIDAE Anthias cyprinoides (Katayama & Amaoka, 1986) 1 SERRANIDAE Anthias helenensis Katayama & Amaoka, 1986 1 SERRANIDAE Anthias menezesi Anderson & Heemstra, 1980 1 1 1 SERRANIDAE Anthias nicholsi Firth, 1933 1 1 1 1 SERRANIDAE Anthias salmopunctatus Lubbock & Edwards, 1981 1 SERRANIDAE Anthias tenuis Nichols, 1920 1 1 1 1 SERRANIDAE Anthias woodsi Anderson & Heemstra, 1980 1 1 SERRANIDAE Bathyanthias cubensis (Schultz, 1958) 1 SERRANIDAE Bathyanthias mexicanus (Schultz, 1958) 1 SERRANIDAE Bathyanthias roseus Günther, 1880 1 SERRANIDAE Bullisichthys caribbaeus Rivas, 1971 1 1 SERRANIDAE Callanthias ruber (Rafinesque, 1810) 1 1 1 1 1 1 SERRANIDAE Centropristis fuscula Poey, 1861 1 1 1 SERRANIDAE Centropristis ocyurus (Jordan & Evermann, 1887) 1 1 SERRANIDAE Centropristis philadelphica (Linnaeus, 1758) 1 1 SERRANIDAE Centropristis striata (Linnaeus, 1758) 1 1 SERRANIDAE Cephalopholis cruentata (Lacepède, 1802) 1 1 1 1 SERRANIDAE Cephalopholis fulva (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Cephalopholis furcifer (Valenciennes, 1828) 1 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Cephalopholis nigri (Günther, 1859) 1 1 1 1 SERRANIDAE Cephalopholis taeniops (Valenciennes, 1828) 1 1 1 1 SERRANIDAE Dermatolepis inermis (Valenciennes, 1833) 1 1 1 1 1 1 1 1 SERRANIDAE Diplectrum bivittatum (Valenciennes, 1828) 1 1 1 1 SERRANIDAE Diplectrum formosum (Linnaeus, 1766) 1 1 1 1 1 1 1 1 SERRANIDAE Diplectrum radiale (Quoy & Gaimard, 1824) 1 1 1 1 1 1 SERRANIDAE Dules auriga Cuvier, 1829 1 1 1 SERRANIDAE Epinephelus adscensionis (Osbeck, 1765) 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Epinephelus aeneus (Geoffroy Saint-Hilaire, 1817) 1 1 1 1 1 1 SERRANIDAE Epinephelus andersoni Boulenger, 1903 1 1 SERRANIDAE Epinephelus caninus (Valenciennes, 1843) 1 1 1 1 SERRANIDAE Epinephelus chabaudi (Castelnau, 1861) 1 1 SERRANIDAE Epinephelus costae (Steindachner, 1878) 1 1 1 1 1 1 1 SERRANIDAE Epinephelus drummondhayi Goode & Bean, 1878 1 1 1 SERRANIDAE Epinephelus fasciatus (Forsskål, 1775) 1 1 1 SERRANIDAE Epinephelus flavocaeruleus (Lacepède, 1802) 1 1 SERRANIDAE Epinephelus flavolimbatus Poey, 1865 1 1 1 1 1 SERRANIDAE Epinephelus goreensis (Valenciennes, 1830) 1 1 1 SERRANIDAE Epinephelus guttatus (Linnaeus, 1758) 1 1 1 1 SERRANIDAE Epinephelus haifensis Ben-Tuvia, 1953 1 1 1 1 SERRANIDAE Epinephelus itajara (Lichtenstein, 1822) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Epinephelus lanceolatus (Bloch, 1790) 1 1 SERRANIDAE Epinephelus marginatus (Lowe, 1834) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Epinephelus morio (Valenciennes, 1828) 1 1 1 1 1 1 SERRANIDAE Epinephelus mystacinus (Poey, 1852) 1 1 1 1 1 SERRANIDAE Epinephelus nigritus (Holbrook, 1855) 1 1 1 1 1 1 1 SERRANIDAE Epinephelus niveatus (Valenciennes, 1828) 1 1 1 1 1 1 SERRANIDAE Epinephelus poecilonotus (Temminck & Schlegel, 1842) 1 1 SERRANIDAE Epinephelus rivulatus (Valenciennes, 1830) 1 1 SERRANIDAE Epinephelus striatus (Bloch, 1792) 1 1 1 1 SERRANIDAE Gonioplectrus hispanus (Cuvier, 1828) 1 1 1 1 1 1 SERRANIDAE Hemanthias aureorubens (Longley, 1935) 1 1 SERRANIDAE Hemanthias leptus (Ginsburg, 1952) 1 1 SERRANIDAE Hemanthias vivanus (Jordan & Swain, 1885) 1 1 1 1 SERRANIDAE Holanthias caudalis Trunov, 1976 1 SERRANIDAE Holanthias fronticinctus Günther, 1868 1 SERRANIDAE Hypoplectrus aberrans Poey, 1868 1 1 SERRANIDAE Hypoplectrus chlorurus (Cuvier, 1828) 1 1 SERRANIDAE Hypoplectrus gemma Goode & Bean, 1882 1 1 1 SERRANIDAE Hypoplectrus gummigutta (Poey, 1851) 1 1 SERRANIDAE Hypoplectrus guttavarius (Poey, 1852) 1 1 SERRANIDAE Hypoplectrus indigo (Poey, 1851) 1 1 SERRANIDAE Hypoplectrus nigricans (Poey, 1852) 1 1 1 SERRANIDAE Hypoplectrus providencianus Acero P. & Garzón-Ferreira, 1994 1 SERRANIDAE Hypoplectrus puella (Cuvier, 1828) 1 1 1 1 SERRANIDAE Hypoplectrus sp.(Mayan_hamlet) 1 SERRANIDAE Hypoplectrus sp.(tan_hamlet) 1 1 SERRANIDAE Hypoplectrus unicolor (Walbaum, 1792) 1 1 1 SERRANIDAE Jeboehlkia gladifer Robins, 1967 1 1 SERRANIDAE Liopropoma aberrans (Poey, 1860) 1 1 1 SERRANIDAE Liopropoma carmabi (Randall, 1963) 1 1 1 1 SERRANIDAE Liopropoma eukrines (Starck & Courtenay, 1962) 1 1 SERRANIDAE Liopropoma mowbrayi Woods & Kanazawa, 1951 1 1 1 1 SERRANIDAE Liopropoma rubre Poey, 1861 1 1 1 SERRANIDAE Liopropoma sp.n.(Bahia) 1 SERRANIDAE Liopropoma sp.n.(Stome) 1 SERRANIDAE Meganthias carpenteri Anderson, 2006 1 SERRANIDAE Meganthias natalensis (Fowler, 1925) 1 1 SERRANIDAE Mycteroperca acutirostris (Valenciennes, 1828) 1 1 1 1 1 SERRANIDAE Mycteroperca bonaci (Poey, 1860) 1 1 1 1 1 1 1 1 1 SERRANIDAE Mycteroperca cidi Cervigón, 1966 1 1 SERRANIDAE Mycteroperca fusca (Lowe, 1838) 1 1 1 1 SERRANIDAE Mycteroperca interstitialis (Poey, 1860) 1 1 1 1 1 1 1 SERRANIDAE Mycteroperca microlepis (Goode & Bean, 1879) 1 1 1 1 1 SERRANIDAE Mycteroperca phenax Jordan & Swain, 1884 1 1 1 SERRANIDAE Mycteroperca rubra (Bloch, 1793) 1 1 1 1 SERRANIDAE Mycteroperca tigris (Valenciennes, 1833) 1 1 1 1 1 SERRANIDAE Mycteroperca venenosa (Linnaeus, 1758) 1 1 1 1 1 1 1 SERRANIDAE Nemanthias carberryi Smith, 1954 1 1 SERRANIDAE Paralabrax dewegeri (Metzelaar, 1919) 1 1 SERRANIDAE Parasphyraenops atrimanus Bean, 1912 1 1 SERRANIDAE Parasphyraenops incisus (Colin, 1978) 1 1 SERRANIDAE Plectranthias garrupellus Robins & Starck, 1961 1 1 SERRANIDAE Pronotogrammus martinicensis (Guichenot, 1868) 1 1 1 1 1 SERRANIDAE Pseudogramma gregoryi (Breder, 1927) 1 1 1 1 SERRANIDAE Pseudogramma guineensis (Norman, 1935) 1 1 1 SERRANIDAE Rypticus bistrispinus (Mitchill, 1818) 1 1 1 1 1 1 SERRANIDAE Rypticus bornoi Beebe & Tee-Van, 1928 1 1 SERRANIDAE Rypticus maculatus Holbrook, 1855 1 1 SERRANIDAE Rypticus saponaceus (Bloch & Schneider, 1801) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 SERRANIDAE Rypticus subbifrenatus Gill, 1861 1 1 1 1 1 1 1 1 1 SERRANIDAE Schultzea beta (Hildebrand, 1940) 1 1 1 SERRANIDAE Serraniculus pumilio Ginsburg, 1952 1 1 1 SERRANIDAE Serranus accraensis (Norman, 1931) 1 SERRANIDAE Chelidoperca africana Cadenat, 1960 1 1 SERRANIDAE Serranus annularis (Günther, 1880) 1 1 1 1 1 SERRANIDAE Serranus atricauda Günther, 1874 1 1 1 1 1 1 1 SERRANIDAE Serranus atrobranchus (Cuvier, 1829) 1 1 1 1 1 SERRANIDAE Serranus baldwini (Evermann & Marsh, 1899) 1 1 1 1 1 1 SERRANIDAE Serranus cabrilla (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 SERRANIDAE Serranus chionaraia Robins & Starck, 1961 1 1 1 SERRANIDAE Serranus dewegeri Metzelaar, 1919 1 1 1 SERRANIDAE Serranus flaviventris Gilchrist, 1904 1 1 1 1 1 SERRANIDAE Serranus hepatus (Linnaeus, 1758) 1 1 1 SERRANIDAE Serranus heterurus (Cadenat, 1937) 1 1 SERRANIDAE Serranus knysnaensis Gilchrist, 1904 1 1 1 SERRANIDAE Serranus luciopercanus Poey, 1852 1 SERRANIDAE Serranus maytagi Robins & Starck, 1961 1 1 SERRANIDAE Serranus notospilus Longley, 1935 1 1 1 SERRANIDAE Serranus phoebe Poey, 1851 1 1 1 1 1 1 1 SERRANIDAE Serranus sanctaehelenae Boulenger, 1895 1 1 SERRANIDAE Serranus scriba (Linnaeus, 1758) 1 1 1 1 1 1 SERRANIDAE Serranus sp.n. (Cabo Verde) 1 SERRANIDAE Serranus sp.n. (Stome) 1 SERRANIDAE Serranus sp.n.(cf_notospilus) 1 1 SERRANIDAE Serranus subligarius (Cope, 1870) 1 1 1 SERRANIDAE Serranus tabacarius (Cuvier, 1829) 1 1 1 1 SERRANIDAE Serranus tigrinus (Bloch, 1790) 1 1 1 SERRANIDAE Serranus tortugarum Longley, 1935 1 1 SIGANIDAE Siganus sutor (Valenciennes, 1835) 1 1 SPARIDAE Archosargus probatocephalus (Walbaum, 1792) 1 1 1 1 SPARIDAE Archosargus rhomboidalis (Linnaeus, 1758) 1 1 1 1 1 1 SPARIDAE Argyrops spinifer (Forsskål, 1775) 1 1 SPARIDAE Argyrozona argyrozona (Valenciennes, 1830) 1 1 1 SPARIDAE Boops boops (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 SPARIDAE Boopsoidea inornata Castelnau, 1861 1 1 SPARIDAE Calamus arctifrons Goode & Bean, 1882 1 1 SPARIDAE Calamus bajonado (Bloch & Schneider, 1801) 1 1 1 1 1 1 SPARIDAE Calamus calamus (Valenciennes, 1830) 1 1 1 1 1 1 SPARIDAE Calamus campechanus Randall & Caldwell, 1966 1 SPARIDAE Calamus cervigoni Randall & Caldwell, 1966 1 1 SPARIDAE Calamus leucosteus Jordan & Gilbert, 1885 1 1 SPARIDAE Calamus mu Randall & Caldwell, 1966 1 SPARIDAE Calamus nodosus Randall & Caldwell, 1966 1 1 SPARIDAE Calamus penna (Valenciennes, 1830) 1 1 1 1 1 SPARIDAE Calamus pennatula Guichenot, 1868 1 1 1 1 SPARIDAE Calamus proridens Jordan & Gilbert, 1884 1 1 1 SPARIDAE Cheimerius nufar (Valenciennes, 1830) 1 1 SPARIDAE Chrysoblephus anglicus (Gilchrist & Thompson, 1908) 1 1 SPARIDAE Chrysoblephus cristiceps (Valenciennes, 1830) 1 1 SPARIDAE Chrysoblephus gibbiceps (Valenciennes, 1830) 1 1 SPARIDAE Chrysoblephus laticeps (Valenciennes, 1830) 1 1 SPARIDAE Chrysoblephus puniceus (Gilchrist & Thompson, 1908) 1 1 SPARIDAE Cymatoceps nasutus (Castelnau, 1861) 1 1 SPARIDAE Dentex angolensis Poll & Maul, 1953 1 1 1 1 SPARIDAE Dentex barnardi Cadenat, 1970 1 1 SPARIDAE Dentex canariensis Steindachner, 1881 1 1 1 1 1 SPARIDAE Dentex congoensis Poll, 1954 1 1 SPARIDAE Dentex dentex (Linnaeus, 1758) 1 1 1 1 1 SPARIDAE Dentex gibbosus (Rafinesque, 1810) 1 1 1 1 1 1 1 1 SPARIDAE Dentex macrophthalmus (Bloch, 1791) 1 1 1 1 1 1 1 SPARIDAE Dentex maroccanus Valenciennes, 1830 1 1 1 1 1 SPARIDAE Diplodus annularis (Linnaeus, 1758) 1 1 1 SPARIDAE Diplodus argenteus_argenteus(Valenciennes, 1830) 1 1 1 1 SPARIDAE Diplodus argenteus_caudimacu(Plaoey, 1860) 1 1 1 SPARIDAE Diplodus bellottii (Steindachner, 1882) 1 1 1 1 SPARIDAE Diplodus bermudensis Caldwell, 1965 1 SPARIDAE Diplodus cervinus_cervinus (Lowe, 1838) 1 1 1 1 1 1 SPARIDAE Diplodus cervinus_hottentotus (Smith, 1844) 1 1 SPARIDAE Diplodus fasciatus (Valenciennes, 1830) 1 SPARIDAE Diplodus holbrooki (Bean, 1878) 1 1 1 SPARIDAE Diplodus prayensis (Smith, 1844) 1 SPARIDAE Diplodus puntazzo Steindachner, 1882 1 1 1 1 1 1 1 SPARIDAE Diplodus sargus_ascensionis Steindachner, 1903 1 SPARIDAE Diplodus sargus_cadenati de la Paz, Bauchot & Daget, 1974 1 1 1 1 1 SPARIDAE Diplodus sargus_capensis (Smith, 1844) 1 1 1 1 SPARIDAE Diplodus sargus_helenae (Sauvage, 1879) 1 SPARIDAE Diplodus sargus_lineatus (Valenciennes, 1830) 1 SPARIDAE Diplodus sargus_sargus (Linnaeus, 1758) 1 1 SPARIDAE Diplodus vulgaris (Geoffroy Saint-Hilaire, 1817) 1 1 1 1 1 1 1 1 SPARIDAE Gymnocrotaphus curvidens Günther, 1859 1 1 SPARIDAE Lagodon rhomboides (Linnaeus, 1766) 1 1 1 1 SPARIDAE Lithognathus aureti Smith, 1962 1 1 SPARIDAE Lithognathus lithognathus (Cuvier, 1829) 1 1 1 SPARIDAE Lithognathus mormyrus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 SPARIDAE Oblada melanura (Linnaeus, 1758) 1 1 1 1 1 1 1 1 SPARIDAE Pachymetopon aeneum (Gilchrist & Thompson, 1908) 1 1 SPARIDAE Pachymetopon blochii (Valenciennes, 1830) 1 1 SPARIDAE Pachymetopon grande Günther, 1859 1 1 SPARIDAE Pagellus acarne (Risso, 1827) 1 1 1 1 1 1 1 SPARIDAE Pagellus bellottii_bellottii Steindachner, 1882 1 1 1 1 1 1 SPARIDAE Pagellus bellottii_natalensis Steindachner, 1903 1 1 SPARIDAE Pagellus bogaraveo (Brünnich, 1768) 1 1 1 1 1 SPARIDAE Pagellus erythrinus (Linnaeus, 1758) 1 1 1 1 1 1 1 SPARIDAE Pagrus africanus Akazaki, 1962 1 1 1 1 1 SPARIDAE Pagrus auriga Valenciennes, 1843 1 1 1 1 1 1 1 1 SPARIDAE Pagrus caeruleostictus (Valenciennes, 1830) 1 1 1 1 1 SPARIDAE Pagrus pagrus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 SPARIDAE Petrus rupestris (Valenciennes, 1830) 1 1 SPARIDAE Polysteganus praeorbitalis (Günther, 1859) 1 1 SPARIDAE Polysteganus undulosus (Regan, 1908) 1 1 SPARIDAE Porcostoma dentata (Gilchrist & Thompson, 1908) 1 1 SPARIDAE Pterogymnus laniarius (Valenciennes, 1830) 1 1 SPARIDAE Rhabdosargus globiceps (Valenciennes, 1830) 1 1 1 1 1 SPARIDAE Rhabdosargus holubi (Steindachner, 1881) 1 1 SPARIDAE Rhabdosargus sarba (Forsskål, 1775) 1 1 SPARIDAE Sarpa salpa (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 SPARIDAE Sparodon durbanensis (Castelnau, 1861) 1 1 SPARIDAE Sparus aurata Linnaeus, 1758 1 1 1 1 1 SPARIDAE Spondyliosoma cantharus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 SPARIDAE Spondyliosoma emarginatum (Valenciennes, 1830) 1 1 1 SPARIDAE Stenotomus caprinus Jordan & Gilbert, 1882 1 1 SPARIDAE Stenotomus chrysops (Linnaeus, 1766) 1 SPARIDAE Virididentex acromegalus (Osório, 1911) 1 SYNGNATHIDAE Acentronura dendritica (Barbour, 1905) 1 1 1 1 1 1 SYNGNATHIDAE Anarchopterus criniger (Bean & Dresel, 1884) 1 1 1 1 SYNGNATHIDAE Anarchopterus tectus (Dawson, 1978) 1 1 1 1 1 SYNGNATHIDAE Bryx dunckeri (Metzelaar, 1919) 1 1 1 1 1 1 SYNGNATHIDAE Bryx randalli (Herald, 1965) 1 1 SYNGNATHIDAE Bryx sp.n. 1 SYNGNATHIDAE Cosmocampus albirostris (Kaup, 1856) 1 1 1 1 1 SYNGNATHIDAE Cosmocampus brachycephalus (Poey, 1868) 1 1 1 SYNGNATHIDAE Cosmocampus elucens (Poey, 1868) 1 1 1 1 1 1 SYNGNATHIDAE Cosmocampus hildebrandi (Herald, 1965) 1 1 SYNGNATHIDAE Cosmocampus profundus (Herald, 1965) 1 1 SYNGNATHIDAE Cosmocampus retropinnis Dawson, 1982 1 1 SYNGNATHIDAE Enneacampus ansorgii (Boulenger, 1910) 1 1 1 SYNGNATHIDAE Enneacampus kaupi (Bleeker, 1863) 1 1 SYNGNATHIDAE Entelurus aequoreus (Linnaeus, 1758) 1 SYNGNATHIDAE Halicampus crinitus (Jenyns, 1842) 1 1 1 1 1 1 1 1 1 SYNGNATHIDAE Hippocampus aff._reidi 1 1 1 SYNGNATHIDAE Hippocampus algiricus Kaup, 1856 1 1 1 1 1 SYNGNATHIDAE Hippocampus capensis Boulenger, 1900 1 SYNGNATHIDAE Hippocampus erectus Perry, 1810 1 1 1 1 1 1 1 1 SYNGNATHIDAE Hippocampus guttulatus Cuvier, 1829 1 1 SYNGNATHIDAE Hippocampus hippocampus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 SYNGNATHIDAE Hippocampus reidi Ginsburg, 1933 1 1 1 1 SYNGNATHIDAE Hippocampus zosterae Jordan & Gilbert, 1882 1 1 1 1 SYNGNATHIDAE Micrognathus andersonii (Bleeker, 1858) 1 1 SYNGNATHIDAE Micrognathus erugatus Herald & Dawson, 1974 1 1 SYNGNATHIDAE Minyichthys inusitatus Dawson, 1983 1 1 SYNGNATHIDAE Minyichthys sentus Dawson, 1982 1 1 SYNGNATHIDAE Minyichthys sp.n. 1 SYNGNATHIDAE Nannocampus elegans Smith, 1961 1 1 SYNGNATHIDAE Nerophis lumbriciformis (Jenyns, 1835) 1 SYNGNATHIDAE Nerophis maculatus Rafinesque, 1810 1 1 1 SYNGNATHIDAE Nerophis ophidion (Linnaeus, 1758) 1 1 1 1 1 SYNGNATHIDAE Penetopteryx nanus (Rosén, 1911) 1 1 SYNGNATHIDAE Syngnathoides biaculeatus (Bloch, 1785) 1 1 SYNGNATHIDAE Syngnathus abaster Risso, 1827 1 1 SYNGNATHIDAE Syngnathus acus Linnaeus, 1758 1 1 1 1 1 1 1 1 1 1 SYNGNATHIDAE Syngnathus floridae (Jordan & Gilbert, 1882) 1 1 1 1 SYNGNATHIDAE Syngnathus folletti Herald, 1942 1 1 1 SYNGNATHIDAE Syngnathus fuscus Storer, 1839 1 1 1 SYNGNATHIDAE Syngnathus louisianae Günther, 1870 1 1 1 SYNGNATHIDAE Syngnathus phlegon Risso, 1827 1 1 SYNGNATHIDAE Syngnathus scovelli (Evermann & Kendall, 1896) 1 1 1 1 SYNGNATHIDAE Syngnathus springeri Herald, 1942 1 1 1 SYNGNATHIDAE Syngnathus taenionotus Canestrini, 1871 1 SYNGNATHIDAE Syngnathus tenuirostris Rathke, 1837 1 SYNGNATHIDAE Syngnathus typhle Linnaeus, 1758 1 1 1 1 1 1 SYNGNATHIDAE Syngnathus watermeyeri Smith, 1963 1 SYNODONTIDAE Saurida brasiliensis Norman, 1935 1 1 1 1 1 1 1 1 SYNODONTIDAE Saurida caribbaea Breder, 1927 1 1 1 1 1 1 SYNODONTIDAE Saurida gracilis (Quoy & Gaimard, 1824) 1 1 SYNODONTIDAE Saurida normani Longley, 1935 1 1 1 SYNODONTIDAE Saurida suspicio Breder, 1927 1 1 SYNODONTIDAE Saurida undosquamis (Richardson, 1848) 1 1 SYNODONTIDAE Synodus foetens (Linnaeus, 1766) 1 1 1 1 1 1 SYNODONTIDAE Synodus indicus (Day, 1873) 1 1 SYNODONTIDAE Synodus intermedius (Spix & Agassiz, 1829) 1 1 1 1 1 1 SYNODONTIDAE Synodus poeyi Jordan, 1887 1 1 1 1 SYNODONTIDAE Synodus saurus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 SYNODONTIDAE Synodus synodus (Linnaeus, 1758) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 SYNODONTIDAE Trachinocephalus myops (Forster, 1801) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 TETRAODONTIDAE Amblyrhynchotes honckenii (Bloch, 1785) 1 1 TETRAODONTIDAE Arothron firmamentum (Temminck & Schlegel, 1850) 1 1 1 TETRAODONTIDAE Arothron hispidus (Linnaeus, 1758) 1 1 TETRAODONTIDAE Arothron immaculatus (Bloch & Schneider, 1801) 1 1 TETRAODONTIDAE Arothron inconditus Smith, 1958 1 1 TETRAODONTIDAE Arothron nigropunctatus (Bloch & Schneider, 1801) 1 1 TETRAODONTIDAE Arothron stellatus (Bloch & Schneider, 1801) 1 1 TETRAODONTIDAE Canthigaster bennetti (Bleeker, 1854) 1 1 TETRAODONTIDAE Canthigaster capistrata (Lowe, 1839) 1 1 1 1 1 TETRAODONTIDAE Canthigaster figueiredoi Moura & Castro, 2002 1 1 1 1 1 TETRAODONTIDAE Canthigaster jamestyleri Moura & Castro, 2002 1 1 TETRAODONTIDAE Canthigaster rivulata (Temminck & Schlegel, 1850) 1 1 TETRAODONTIDAE Canthigaster rostrata (Bloch, 1786) 1 1 1 1 TETRAODONTIDAE Canthigaster sanctaehelenae (Günther, 1870) 1 1 TETRAODONTIDAE Canthigaster solandri (Richardson, 1845) 1 1 TETRAODONTIDAE Canthigaster supramacula Moura & Castro, 2002 1 1 TETRAODONTIDAE Chelonodon patoca (Hamilton, 1822) 1 1 TETRAODONTIDAE Ephippion guttifer (Bennett, 1831) 1 1 1 1 1 TETRAODONTIDAE Sphoeroides dorsalis Longley, 1934 1 1 1 1 1 TETRAODONTIDAE Sphoeroides maculatus (Bloch & Schneider, 1801) 1 1 TETRAODONTIDAE Sphoeroides marmoratus (Lowe, 1838) 1 1 1 1 1 1 1 TETRAODONTIDAE Sphoeroides nephelus (Goode & Bean, 1882) 1 1 1 1 TETRAODONTIDAE Sphoeroides pachygaster (Müller & Troschel, 1848) 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 TETRAODONTIDAE Sphoeroides spengleri (Bloch, 1785) 1 1 1 1 1 1 1 1 TETRAODONTIDAE Sphoeroides testudineus (Linnaeus, 1758) 1 1 1 1 1 TETRAODONTIDAE Torquigener balteatus Hardy, 1989 1 1 TRIPTERYGIIDAE Acanthanectes hystrix Holleman & Buxton, 1993 1 TRIPTERYGIIDAE Acanthanectes rufus Holleman & Buxton, 1993 1 TRIPTERYGIIDAE Cremnochorites capensis (Gilchrist & Thompson, 1908) 1 TRIPTERYGIIDAE Enneanectes altivelis Rosenblatt, 1960 1 1 1 1 1 1 1 1 TRIPTERYGIIDAE Enneanectes atrorus Rosenblatt, 1960 1 1 TRIPTERYGIIDAE Enneanectes boehlkei Rosenblatt, 1960 1 1 TRIPTERYGIIDAE Enneanectes jordani (Evermann & Marsh, 1899) 1 1 TRIPTERYGIIDAE Enneanectes n.sp.1 1 TRIPTERYGIIDAE Enneanectes n.sp.2 1 TRIPTERYGIIDAE Enneanectes n.sp.3 1 TRIPTERYGIIDAE Enneanectes pectoralis (Fowler, 1941) 1 1 1 TRIPTERYGIIDAE Enneanectes smithi Lubbock & Edwards, 1981 1 TRIPTERYGIIDAE Helcogramma obtusirostre (Klunzinger, 1871) 1 1 1 TRIPTERYGIIDAE Helcogrammoides cunninghami (Smitt, 1898) 1 TRIPTERYGIIDAE Tripterygion delaisi delaisi Cadenat & Blache, 1970 1 1 1 1 TRIPTERYGIIDAE Tripterygion delaisi xanthosoma Zander & Heymer, 1970 1 1 1 TRIPTERYGIIDAE Tripterygion melanurus Guichenot, 1850 1 1 TRIPTERYGIIDAE Tripterygion tripteronotus (Risso, 1810) 1 1 1 URANOSCOPIDAE Astroscopus guttatus Abbott, 1860 1 URANOSCOPIDAE Astroscopus sexspinosus (Steindachner, 1876) 1 1 URANOSCOPIDAE Astroscopus ygraecum (Cuvier, 1829) 1 1 1 1 1 URANOSCOPIDAE Uranoscopus albesca Günther, 1861 1 1 URANOSCOPIDAE Uranoscopus cadenati Smith, 1966 1 1 URANOSCOPIDAE Uranoscopus polli (Latreille, 1804) 1 1 URANOSCOPIDAE Uranoscopus scaber Linnaeus, 1758 1 1 1 1 1

452 280 810 757 396 382 83 101 117 52 85 82 76 95 133 182 213 172 207 221 282 250 88 296 260 Table S2 APPENDIX – Sister-species table and references (Floeter et al.)

Legends: WA = (North + South) West Atlantic; NWA = Northwestern Atlantic; SWA = Southwestern Atlantic; EA = Eastern Atlantic; TEA = Tropical Eastern Atlantic; MAR = mid-Atlantic Ridge; ASC = Ascension; SHE = St. Helena; CV = Cape Verde Is.; MAC = Macaronesia; IO = Indian Ocean; IWP = Indo-West Pacific.

Figure Family Genus Species Reference Type (ms) ANTENNARIIDAE Antennarius multiocellatus (WA)/pardalis (EA) Pietsch & Grobecker WA-EA (1987) Antennarius ocellatus (NWA)/senegalensis (EA) Pietsch & Grobecker WA-EA (1987) APOGONIDAE Apogon maculatus (NWA)/americanus Look alike NWA-SWA (SWA) AULOSTOMIDAE Aulostomus strigosus (SWA-EA)/chinensis (IP) Bowen et al. (2001) IWP-EA 12 BLENNIIDAE Hypleurochilus pseudoaequipinnis (WA)/ Look alike WA-EA aequipinnis (EA) Ophioblennius atlanticus (EA)/ sp1 (ASC), sp2 Muss et al. (2001) EA-MAR 12 (CV)/macclurei (NWA) CARANGIDAE Hemicaranx amblyrhynchus (WA)/bicolor (EA) Smith-Vaniz, pers WA-EA obs. Trachinotus marginatus (WA)/goreensis (EA) Smith-Vaniz, pers WA-EA obs. CARAPIDAE Carapus bermudensis (WA)/acus (EA) Look alike WA-EA CHAETODONTIDAE Prognathodes aculeatus (NWA)/brasiliensis Hubbs (1963) NWA-SWA (SWA) Prognathodes aya (WA)/marcellae (EA) Hubbs (1963) WA-EA Prognathodes dichrous (MAR)/obliquus (St Paul's Look alike WA-MAR Rocks) CIRRHITIDAE Amblycirrhitus pinos (WA)/earnshawi (ASC) Look alike WA-MAR Cirrhitus atlanticus (EA)/pinnulatus (IWP) Look alike IWP-EA DACTYLOSCOPIDAE Platygillellus rubrocinctus (NWA)/brasiliensis Feitoza (2002) NWA-SWA (SWA) DIODONTIDAE Chilomycterus spinosus (WA)/mauretanicus (EA) Leis (2006) WA-EA DREPANIDAE Drepane africana (EA)/ longimana (IWP) Heemstra & IWP-EA Heemstra (2004) GOBIESOCIDAE Tomicodon fasciatus (NWA)/australis (SWA) Briggs, pers com NWA-SWA GOBIIDAE Elacatinus figaro (SWA)/randalli (NWA) Sazima et al. (1998); NWA-SWA 13 Taylor & Hellberg (2005) GRAMMATIDAE Gramma loreto (NWA)/brasiliensis (SWA) Sazima et al. (1998) NWA-SWA LABRIDAE Bodianus pulchellus/rufus (WA)/insularis Gomon & Lubbock WA-MAR (MAR) (1980) Clepticus parrae (NWA)/brasiliensis Heiser et al. (2000) WA-EA 12 (SWA)/africanus (TEA) Coris julis (MED+NEA+MAC)/atlantica Aurelle et al. (2003) other (TEA) Halichoeres cyanocephalus (NWA)/dimidiatus Rocha (2004) NWA-SWA (SWA) Halichoeres maculipinna (NWA)/penrosei Rocha (2004) NWA-SWA (SWA) Halichoeres radiatus (WA)/brasiliensis (SWA) Rocha et al. (2005) WA-SWA Thalassoma ascensionis (ASC), sanctahelenae Costagliola et al. EA-MAR 12 (SHE)/newtoni (EA) (2005) Thalassoma bifasciatum (NWA)/noronhanum Costagliola et al. NWA-SWA 12 (SWA) (2004) LABRISOMIDAE Malacoctenus triangulatus (WA)/africanus (EA) Look alike WA-EA Paraclinus fasciatus (NWA)/arcanus (SWA) Guimarães & NWA-SWA Bacellar (2002) Paraclinus marmoratus (NWA)/spectator Guimarães & NWA-SWA (SWA) Bacellar (2002) Paraclinus nigripinnis (NWA)/rubicundus Guimarães & NWA-SWA (SWA) Bacellar (2002) Starksia ocellata (NWA)/brasiliensis (SWA) Look alike NWA-SWA LUTJANIDAE Lutjanus apodus (WA)/goreensis (EA) Look alike WA-EA Lutjanus cyanopterus (WA)/agennes (EA) Look alike WA-EA Lutjanus mahogony (WA)/fulgens (EA) Look alike WA-EA MULLIDAE Mullus auratus (NWA)/argentinae (SWA)/ Look alike WA-EA barbatus (NEA)/surmuletus (EA) Pseudupeneus maculatus (WA)/prayensis (EA) Look alike WA-EA OPHICHTHIDAE Myrichthys ocellatus (WA)/pardalis (EA) Look alike WA-EA OPHIDIIDAE Ophidion holbrookii (WA)/barbatum (EA) Look alike WA-EA OPISTOGNATHIDAE Opistognathus aurifrons (NWA)/spn (SWA- Smith-Vaniz, pers NWA-SWA southern NWA) obs. Opistognathus macrognathus (NWA)/brasiliensis Smith-Vaniz, pers NWA-SWA (SWA) obs. OSTRACIIDAE Acanthostracion polygonius-quadricornis (WA)/ Klassen (1995) WA-EA 12 guineensis-notacanthus (EA) POMACANTHIDAE Centropyge argi (NWA)/aurantonotus (NWA- Bowen et al. (2006) WA-MAR 12 SWA-EA)/resplendens (ASC) POMACENTRIDAE Abudefduf saxatilis (WA-EA)/hoefleri (EA) Look alike WA-EA Chromis chromis (MED)/limbata Domingues et al. other 12 (MAC+TEA) (2005) Microspathodon chrysurus (WA)/frontatus (EA) Look alike WA-EA Stegastes aff variabilis (NWA)/variabilis Robertson, NWA-SWA (SWA) unpublished data Stegastes rocasensis (RC-FN)/sanctipauli Robertson, others (SPR) unpublished data Stegastes fuscus (SWA)/imbricatus (EA) Robertson, WA-EA unpublished data Stegastes partitus (CA)/pictus (SWA- Robertson, WA-MAR Venez)/lubbocki (AS)/sanctahelenae unpublished data (SH) PTERELEOTRIDAE Ptereleotris helenae (NWA)/randalli (SWA) Gasparini et al. NWA-SWA (2003) SCARIDAE Nicholsina usta (WA)/collettei (EA) Robertson et al. WA-EA 12 (2006) Scarus coelestinus (NWA)/trispinosus Choat, pers com NWA-SWA (SWA)/guacamaia (WA) Scarus taeniopterus (NWA)/zelindae Choat, pers com NWA-SWA (SWA) Sparisoma cretense (EA)/strigatum (MAR) Robertson et al. EA-MAR 14 (2006) Sparisoma atomarium (NWA)/tuiupiranga Robertson et al. NWA-SWA 14 (SWA) (2006) Sparisoma griseorubra (Venez)/frondosum Robertson et al. NWA-SWA 14 (SWA)/chrysopterum (NWA) (2006) Sparisoma rubripinne (NWA)/axillare (SWA- Robertson et al. NWA-SWA 12, 14 Venez)/spn (EA) (2006) Sparisoma viride (NWA)/amplum (SWA) Robertson et al. NWA-SWA 14 (2006) SERRANIDAE Dermatolepis inermis (WA)/striolata(IO) Craig et al. (2004) others 12, 14 Meganthias carpenteri (EA)/natalensis (SWIO) Anderson (2006) IWP-EA Rypticus saponaceus (WA)/ sp.n. (EA) Carlin et al. (2003) WA-EA Cephalopholis nigri (EA)/ argus (IWP) Craig & Hastings IWP-EA (2007) Mycteroperca acutirostris (WA)/ rubra (EA, coast)/ Heemstra (1991); WA-EA fusca (MAC+CV) Heemstra & Randall (1993) SYNGNATHIDAE Hippocampus reidi (WA)/algiricus (EA) Casey et al. (2004) WA-EA 12 TETRAODONTIDAE Canthigaster rostrata (NWA)/figueiredoi Moura & Castro WA-EA (SWA)/capistrata (MAC+CV)/ (2002) supramacula (EA) Sphoeroides spengleri (WA)/marmoratus (EA) Moura & Castro WA-EA (2002)

References

Anderson, W.D., Jr. (2006) Meganthias carpenteri, new species of fish from the eastern Atlantic

Ocean, with a key to eastern Atlantic Anthiinae (Perciformes: Serranidae). Proceedings of the

Biological Society of Washington, 119, 404–417.

Aurelle, D., Guillemaud, T., Afonso, P., Morato, T., Wirtz, P., Santos, R.S. & Cancela, M.L. (2003)

Genetic study of Coris julis (Osteichtyes, Perciformes, Labridae) evolutionary history and

dispersal abilities. Comptes Rendus Biologies, 326, 771–785.

Bath, H. (1994) Untersuchung der Arten Hypleurochilus geminatus (Wood 1825), H. fissicornis (Quoy

& Gaimard 1824) und H. aequipinnis (Günther 1861), mit Revalidation von Hypleurochilus

multifilis (Girard 1858) und Beschreibung von zwei neuen Arten. Senckenbergiana biologica, 74,

59–85.

Bernardi, G. & Lape, J. (2005) Tempo and mode of speciation in the Baja California disjunct fish

species Anisotremus davidsonii. Molecular Ecology, 14, 4085–4096.

Bowen, B.W., Bass, A.L., Rocha, L.A., Grant, W.S. & Robertson, D.R. (2001) Phylogeography of the

trumpetfish (Aulostomus): Ring species complex on a global scale? Evolution, 55, 1029–1039.

Bowen, B.W., Muss, A.J., Rocha, L.A. & Grant, W.S. (2006) Shallow mtDNA coalescence in Atlantic

pygmy angelfish (genus Centropyge) indicates a recent invasion from the Indian Ocean. Journal of

Heredity, 97, 1–12.

Carlin, J.L., Robertson, D.R. & Bowen, B.W. (2003) Ancient divergences and recent connections in

two tropical Atlantic reef fishes: Epinephelus adscensionis and Rypticus saponaceous (Percoidei:

Serranidae). Marine Biology, 143, 1057–1069. Casey, S.P., Hall, H.J., Stanley, H.F. & Vincent, A.C.J. (2004) The origin and evolution of seahorses

(genus Hippocampus): a phylogenetic study using the cytochrome b gene of mitochondrial DNA.

Molecular Phylogenetics and Evolution, 30, 261–272.

Costagliola, D., Robertson, D.R., Guidetti, P., Stefanni, S., Wirtz, P., Heiser, J.B. & Bernardi, G.

(2004) Evolution of coral reef fish Thalassoma spp. (Labridae). 2. Evolution of the eastern

Atlantic species. Marine Biology, 144, 377–383.

Craig, M.T., Hastings, P.A. & Pondella, D.J. (2004) Speciation in the Central American Seaway: the

importance of taxon sampling in the identification of trans-isthmian geminate pairs. Journal of

Biogeography, 31, 1085–1091.

Craig, M.T. & Hastings, P.A. (2007) A molecular phylogeny of the groupers of the subfamily

Epinephelinae (Serranidae) with a revised classification of the Epinephelini. Ichthyological

Research, 54, 1–17.

Domingues, V.S., Bucciarelli, G., Almada, V.C. & Bernardi, G. (2005) Historical colonization and

demography of the Mediterranean damselfish, Chromis chromis. Molecular Ecology, 14, 4051–

4063.

Domingues, V.S., Santos, R.S., Brito, A., & Almada, V. (2006) Historical population dynamics and

demography of the eastern Atlantic pomacentrid Chromis limbata (Valenciennes, 1833).

Molecular Phylogenetics and Evolution, 40, 139–147.

Domingues, V.S., Almada, V.C. Santos, R.S., Brito, A. & Bernardi, G. (2007) Phylogeography and

evolution of the triplefin Tripterygion delaisi (Pisces, Blennioidei). Marine Biology, 150, 509–

519.

Feitoza, B.M. (2002) Platygillellus brasiliensis n. sp. (Perciformes: Dactyloscopidae), the third species

of the genus from the Atlantic. Aqua Journal of Ichthyology and Aquatic Biology 6(1), 21-28.

Gasparini, J.L., Rocha, L.A. & Floeter, S.R. (2001) Ptereleotris randalli n. sp., a new dartfish

(Gobioidei: Microdesmidae) from the Brazilian Coast. Aqua Journal of Ichthyology and Aquatic

Biology. 4(3): 109–114.

Gomon, M.F. & Lubbock, R. (1980) A new hogfish of the genus Bodianus (Teleostei: Labridae) from

islands of the mid-Atlantic ridge. Northeast Gulf Science, 3 [for 1979], 104–111. Guimarães, R.Z.P. (1999) Revision, phylogeny and comments on biogeography of soapfishes of the

genus Rypticus (Teleostei: Serranidae). Bulletin of Marine Science 65, 337–379.

Guimarães, R.Z.P & Bacelar, A.C.L.H. (2002) Review of the Brazilian species of Paraclinus

(Teleostei: Labrisomidae), with descriptions of two new species and revalidation of Paraclinus

rubicundus (Starks). Copeia 2002, 419–427.

Hadley-Hansen, P.E. (1986) Revision of the tripterygiid fish genus Helcogramma, including

descriptions of four new species. Bulletin of Marine Science, 38, 313–354.

Heemstra, P.C. (1991) A taxonomic revision of the Eastern Atlantic groupers (Pisces: Serranidae).

Boletim Museu Municipal do Funchal, 43, 5–71.

Heemstra, P.C. & Randall, J.E. (1993) FAO Species Catalogue, Vol. 16. Groupers of the world

(Family Serranidae, SubfamilyEpinephelinae). An annotated and illustrated catalogue of the

grouper, rockcod, hind, coral grouper and lyretail species known to date. FAO Fish. Syn., 125(16):

1–382. FAO: Rome.

Heemstra, P.C. & Heemstra, E. (2004) Coastal Fishes of Southern Africa. Grahamstown: NISC-

SAIAB.

Heiser, J.B., Moura, R.L. & Robertson, D.R. (2000) Two new species of creole wrasse (Labridae:

Clepticus) from opposite sides of the Atlantic. Aqua Journal of Ichthyology and Aquatic Biology,

4, 67–76.

Hubbs, C.L. (1963) Chaetodon aya and related deep-living butterflyfishes: their variation, dis-

tribution and synonyms. Bull. Mar. Sci. Gulf Caribbean 13, 133–192.

Klassen, G.J. (1995) Phylogeny and biogeography of the Ostraciinae (Tetraodontiformes: Ostraciidae)

Bulletin of Marine Science, 57, 393–441.

Leis, J.M. (2006) Nomenclature and distribution of the species of the porcupinefish family

Diodontidae (Pisces, Teleostei). Memoirs of Museum Victoria 63, 77–90

Moura, RL & Castro, R.M.C. (2002) Revision of Atlantic sharpnose pufferfishes (Tetraodontiformes:

Tetraodontidae: Canthigaster) with description of three new species. Proc. Biol. Soc. Wash., 115,

32–50. Muss, A., Robertson, D.R., Stepien, C.A., Wirtz, P. & Bowen, B.W. (2001) Phylogeography of

Ophioblennius: the role of ocean currents and geography in reef fish evolution. Evolution, 55,

561–572.

Pietsch, T.W. & Grobecker, D.B. (1987) Frogfishes of the world. Systematics, zoogeography, and

behavioral ecology. Stanford University Press, Stanford, California. 420 p.

Robertson, D.R., Karg, F., Moura, R.L., Victor, B.C. & Bernardi, G. (2006) Mechanisms of speciation

and faunal enrichment in Atlantic parrotfishes. Molecular Phylogenetics and Evolution, 40, 795–

807.

Rocha, L.A. (2004) Mitochondrial DNA and Color Pattern Variation in Three Western Atlantic

Halichoeres (Labridae), with the Revalidation of Two Species. Copeia, 2004, 770–782.

Rocha, L.A., Robertson, D.R., Roman, J. & Bowen, B.W. (2005) Ecological speciation in tropical reef

fishes. Proceedings of the Royal Society London B, 272, 573–579.

Sazima, I., Moura, R.L. & Rosa, R.S. (1997) Elacatinus figaro sp. n. (Perciformes: Gobiidae), a new

cleaner goby from the coast of Brazil. Aqua, Journal of Ichthyology and Aquatic Biology, 2 , 33–

38.

Sazima I., Gasparini, J.L. & Moura, R.L. (1998) Gramma brasiliensis, a new basslet from the western

South Atlantic (Perciformes: Grammatidae). Aqua, Journal of Ichthyology and Aquatic Biology, 3,

39–43.

Summerer, M., Janel, R. & Sturmbauer, C. (2001) Mitochondrial phylogeny and phylogeography

affinities of seabreans of the genus Diplodus (Sparidae). Journal of Fish Biology, 10, 1–23.

Taylor, M.S. & Hellberg, M.E. (2005) Marine radiations at small geographic scales: speciation in

neotropical reef gobies (Elacatinus). Evolution, 59, 374–385.