Adult habitat preferences, larval dispersal, and the comparative phylogeography of three Atlantic surgeonfishes (Teleostei: Acanthuridae)

Although many reef fishes of the tropical Atlantic are widely distributed, there are large discontinuities that may strongly influence phylogeographical patterns. The freshwater outflow of the Amazon basin is recognized as a major barrier that produces a break between Brazilian and Caribbean faunas. The vast oceanic distances between Brazil and the mid-Atlantic ridge islands represent another formidable barrier. To assess the relative importance of these barriers, we compared a fragment of the mitochondrial DNA (mtDNA) cytochrome b gene among populations of three species of Atlantic surgeonfishes: Acanthurus bahianus, A. chirurgus and A. coeruleus. These species have similar life histories but different adult habitat preferences. The mtDNA data show no population structure between Brazil and the mid-Atlantic islands, indicating that this oceanic barrier is readily traversed by the pelagic larval stage of all three surgeonfishes, which spend approximately 45-70 days in the pelagic environment. The Amazon is a strong barrier to dispersal of A. bahianus (d = 0.024, phiST = 0.724), a modest barrier for A. coeruleus (phiST = 0.356), and has no discernible effect as a barrier for A. chirurgus. The later species has been collected on soft bottoms with sponge habitats under the Amazon outflow, indicating that relaxed adult habitat requirements enable it to readily cross that barrier. A limited ability to use soft bottom habitats may also explain the low (but significant) population structure in A. coeruleus. In contrast, A. bahianus has not been collected over deep sponge bottoms, and rarely settles outside shallow reefs. Overall, adult habitat preferences seem to be the factor that differentiates phylogeographical patterns in these reef-associated species.     

Long distance dispersal and connectivity in amphi-Atlantic corals at regional and basin scales

Among Atlantic scleractinian corals, species diversity is highest in the Caribbean, but low diversity and high endemism are observed in various peripheral populations in central and eastern Atlantic islands and along the coasts of Brazil and West Africa. The degree of connectivity between these distantly separated populations is of interest because it provides insight into processes at both evolutionary and ecological time scales, such as speciation, recruitment dynamics and the persistence of coral populations. To assess connectivity in broadly distributed coral species of the Atlantic, DNA sequence data from two nuclear markers were obtained for six coral species spanning their distributional ranges. At basin-wide scales, significant differentiation was generally observed among populations in the Caribbean, Brazil and West Africa. Concordance of patterns in connectivity among co-distributed taxa indicates that extrinsic barriers, such as the Amazon freshwater plume or long stretches of open ocean, restrict dispersal of coral larvae from region to region. Within regions, dispersal ability appears to be influenced by aspects of reproduction and life history. Two broadcasting species, Siderastrea siderea and Montastraea cavernosa, were able to maintain gene flow among populations separated by as much as 1,200 km along the coast of Brazil. In contrast, brooding species, such as Favia gravida and Siderastrea radians, had more restricted gene flow along the Brazilian coast.     

Atlantic reef fish biogeography and evolution

Aim To understand why and when areas of endemism (provinces) of the tropical Atlantic Ocean were formed, how they relate to each other, and what processes have contributed to faunal enrichment. Location Atlantic Ocean. Methods The distributions of 2605 species of reef fishes were compiled for 25 areas of the Atlantic and southern Africa. Maximum‐parsimony and distance analyses were employed to investigate biogeographical relationships among those areas. A collection of 26 phylogenies of various Atlantic reef fish taxa was used to assess patterns of origin and diversification relative to evolutionary scenarios based on spatio‐temporal sequences of species splitting produced by geological and palaeoceanographic events. We present data on faunal (species and genera) richness, endemism patterns, diversity buildup (i.e. speciation processes), and evaluate the operation of the main biogeographical barriers and/or filters. Results Phylogenetic (proportion of sister species) and distributional (number of shared species) patterns are generally concordant with recognized biogeographical provinces in the Atlantic. The highly uneven distribution of species in certain genera appears to be related to their origin, with highest species richness in areas with the greatest phylogenetic depth. Diversity buildup in Atlantic reef fishes involved (1) diversification within each province, (2) isolation as a result of biogeographical barriers, and (3) stochastic accretion by means of dispersal between provinces. The timing of divergence events is not concordant among taxonomic groups. The three soft (non‐terrestrial) inter‐regional barriers (mid‐Atlantic, Amazon, and Benguela) clearly act as ‘filters’ by restricting dispersal but at the same time allowing occasional crossings that apparently lead 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.     

Mechanisms of speciation and faunal enrichment in Atlantic parrotfishes

Relationships based on mtDNA and nDNA sequences were used to assess effects of two major geographic barriers (the >30 myo Atlantic ocean and the approximately 11 myo Amazon-Orinoco outflow) on speciation among Atlantic parrotfishes (Sparisoma and Nicholsina). Allopatric distributions of sister taxa implicate isolating actions of both barriers in all recent speciation in these fishes, with no clear indications that any speciation resulted from other mechanisms. Molecular clock estimates of the timing of lineage splits indicate that both barriers acted by limiting dispersal well after they formed, although the Amazon barrier also may have been a vicariance agent. Fluctuations in sealevel, climate, and ocean-current dynamics over the past approximately 10 my likely produced marked variation in the effectiveness of both barriers, but particularly the Amazon barrier, allowing intermittent dispersal leading to establishment and allopatric speciation. A dynamic Amazon barrier represents a major engine of West Atlantic faunal enrichment that has repeatedly facilitated bidirectional dispersal, allopatric speciation, and remixing of the Caribbean and Brazilian faunas.     

The southwestern Atlantic reef fish fauna: composition and zoogeographic patterns

The Brazilian coast, the Caribbean and the tropical South Atlantic oceanic islands reef ichthyofauna separated into two major clusters: (1) the western Atlantic continental margin and Bermuda, further divided into (1a) the western North Atlantic, and (1b) the Brazilian coast; and (2) the South Atlantic oceanic islands, also divided in (2a) the Brazilian offshore islands, and (2b) Ascension and St Helena. Species geographic ranges suggest the recognition of only two western Atlantic zoogeographic provinces for tropical marine shore fishes. A Brazilian Province, including the offshore localities of Atol das Rocas, Fernando de Noronha, Trindade and St Paul's Rocks, and an expanded West Indian Province that includes Bermuda. Ascension and St Helena should be regarded as a separated Central Atlantic Province.     

Distribution patterns of the genus Pacifigorgia (Octocorallia: Gorgoniidae): track compatibility analysis and parsimony analysis of endemicity

Aim We analysed the distribution patterns of the eastern Pacific octocoral genus Pacifigorgia and deduced its ancestral distribution to determine why Pacifigorgia is absent from the Gulf of Mexico, the Caribbean of central America, and the Antilles. We also examined the current patterns of endemism for Pacifigorgia to look for congruence between hot spots of endemism in the genus and generally recognized areas of endemism for the eastern Pacific. Location The tropical eastern Pacific and western Atlantic, America. Methods We used track compatibility analysis (TCA) and parsimony analysis of endemicity (PAE) to derive ancestral distribution patterns and hot spots of endemism, respectively. Distributional data for Pacifigorgia were gathered from several museum collections and from fieldwork, particularly in the Pacific of Costa Rica and Panama. Results A single generalized track joined the three main continental eastern Pacific biogeographical provinces and the western Atlantic. This track can be included within a larger eastern Atlantic–eastern Pacific transoceanic track that may be the oldest transoceanic track occurring in the region. PAE results designate previously recognized eastern Pacific biogeographical provinces as Pacifigorgia hot spots of endemism. The number of endemic species, which for other taxonomic groups is similar among the eastern Pacific provinces, is higher in the Panamic province for Pacifigorgia. Main conclusions We propose that the absence of Pacifigorgia from the Gulf of Mexico, the Caribbean of central America, and the Antilles is the result of an ancient absence of the genus from these areas rather than the consequence of a major, recent, extinction episode. The Cortez province and the Mexican province appear together as a result of either non‐response to vicariance or dispersal across the Sinaloan Gap. We posit that the Central American Gap acts as a barrier that separates the Panamic province from the northern Cortez–Mexican province.     

Historical biogeography of South American freshwater fishes

Aim To investigate biogeographical patterns of the obligate freshwater fish order Characiformes. Location South America. Methods Parsimony analysis of endemicity, likelihood analysis of congruent geographical distribution, and partition Bremer support were used. Results Areas of endemism are deduced from parsimony analysis of endemicity, and putative dispersal routes from a separate analysis of discordant patterns of distribution. Main conclusions Our results demonstrate the occurrence of 11 major areas of endemism and support a preferential eastern–western differentiation of the characiforms in the Amazonian region, contrasting with the southern–northern differentiation of terrestrial organisms. The areas of endemism identified seem to be deeply influenced by the distribution of the emerged land during the 100‐m marine highstand that occurred during the late Miocene and allow us to hypothesize the existence of eight aquatic freshwater refuges at that time. The raw distribution of non‐endemic species supports nine patterns of species distribution across the 11 areas of endemism, two of which support a southern–northern differentiation in the eastern part of the Amazon. This result shows that the main channel of the Amazon limited dispersal between tributaries from each bank of the river. The levels of endemism further demonstrate that the aquatic freshwater refuges promoted allopatric speciation and later allowed the colonization of the lowlands. By contrast, the biogeographical pattern found in the western part of the Amazon is identified as a result of the Miocene Andean foreland dynamic and the uplift of the palaeoarches that promoted allopatric divergence across several sedimentary basins by the establishment of disconnected floodplains. The assessment of conflicting species distributions also shows the presence of seven putative dispersal routes between the Amazon, Orinoco and Paraná rivers. Our findings suggest that, rather than there being a single predominant process, the establishment of the modern South American freshwater fish biotas is the result of an interaction between marine incursions, uplift of the palaeoarches, and historical connections allowing cross‐drainage dispersal.     

The implications of Tertiary and Quaternary sea level rise events for avian distribution patterns in the lowlands of northern South America

Aim To assess the correspondence between current avian distributions in the lowlands of northern South America with respect to the hypothesized importance of sea level rise and other events over the past 15 million years on speciation. Location Tropical lowlands of north‐western South America. Methods To establish which bird taxa may have originated in each area of endemism, I examined the ranges of all bird species occurring in the Pacific and the Caribbean lowlands. To determine land and sea distribution during a sea level rise of 100 m in north‐western South America and eastern Central America, I traced the 100 m contour line from the Geoatlas of Georama and the Global 30‐Arc‐Second Elevation Data (GTOPO30) produced by the US Geological Survey. Results During a sea level rise of ～100 m, marine incursions would have occurred from the Caribbean Sea and the Pacific Ocean. Several areas of tropical forest and dry/arid vegetation would have been isolated, currently known as the Darién, Chocó, Nechí, Catatumbo, Tumbesian and Guajiran areas of animal and plant endemism. Main conclusions A large part of the high diversity of forest and nonforest birds and other animals and plants in the Pacific rain forest and the Caribbean woodland likely arose as the result of sea level rises, dispersal and other geological and climatic events.     

Assessing Genetic Diversity of Brazilian Reef Fishes by Chromosomal and DNA Markers

Little is known on genetics of Brazilian coral reef fish and most of this information is limited to chromosome characterization of major representative species. The diploid chromosome number in marine fish varies from 2n= 22–26 to 2n = 240–260. Despite of this apparent diversity, most studied marine species have a diploid complement with 48 acrocentric chromosomes. This latter trend is mostly observed among Perciformes, an important major taxon of coral reef fishes. Studies in the families Pomacentridae, Pomacanthidae and Chaetodontidae, for example, have shown a common karyotype pattern entirely formed by 48 uniarmed chromosomes. However, rare numerical and structural chromosome polymorphisms and cryptic chromosome rearrangements involving heterochromatin segments and/or nucleolar organizing sites have been reported among such fishes. Although new chromosome forms can contribute to the establishment of genetically isolated populations, their role in reef fish speciation at marine realm still is an open question. More recently, genomic DNA analyses using RAPD and microsatellites, and sequencing and RFLP of mitochondrial DNA have increasingly been used in Atlantic reef fish species. Genetic homogeneity over wide geographical ranges has been reported for different fish groups, in contrast to several cases of population substructuring related to environmental constraints or evolutionary history. Amazonas outflow and upwelling on the Southeastern coast of Brazil are believed to be strong barriers to dispersal of some reef species. Moreover, it is suggested that the pattern of speciation and population structure at South Atlantic is quite distinctive from Pacific Ocean, even when comparing closely related taxa. Further genetic studies are strongly encouraged in Brazilian reef fishes in order to provide a reliable scenario of the genetic structure in this important and diverse fish group.     

Repeated invasions into the twilight zone: evolutionary origins of a novel assemblage of fishes from deep Caribbean reefs

Mesophotic and deeper reefs of the tropics are poorly known and underexplored ecosystems worldwide. Collectively referred to as the ‘twilight zone’, depths below ~30–50 m are home to many species of reef fishes that are absent from shallower depths, including many undescribed and endemic species. We currently lack even a basic understanding of the diversity and evolutionary origins of fishes on tropical mesophotic reefs. Recent submersible collections in the Caribbean have provided new specimens that are enabling phylogenetic reconstructions that incorporate deep‐reef representatives of tropical fish genera. Here, we investigate evolutionary depth transitions in the family Gobiidae (gobies), the most diverse group of tropical marine fishes. Using divergence‐time estimation coupled with stochastic character mapping to infer the timing of shallow‐to‐deep habitat transitions in gobies, we demonstrate at least four transitions from shallow to mesophotic depths. Habitat transitions occurred in two broad time periods (Miocene, Pliocene–Pleistocene), and may have been linked to the availability of underutilized niches, as well as the evolution of morphological/behavioural adaptations for life on deep reefs. Further, our analysis shows that at least three evolutionary lineages that invaded deep habitats subsequently underwent speciation, reflecting another unique mode of radiation within the Gobiidae. Lastly, we synthesize depth distributions for 95 species of Caribbean gobies, which reveal major bathymetric faunal breaks at the boundary between euphotic and mesophotic reefs. Ultimately, our study is the first rigorous investigation into the origin of Caribbean deep‐reef fishes and provides a framework for future studies that utilize rare, deep‐reef specimens.     

New records of the snow bass Serranus chionaraia (Perciformes: Serranidae) confirm an established population in the Brazilian Province

The snow bass Serranus chionaraia is a small-bodied reef fish presumed to be restricted to the Caribbean Province, with a single specimen captured south of the Amazon River mouth. Recent surveys with baited remote underwater stereo-video systems detected the species c. 1900 km southward. Meristic and morphometric characters of two specimens examined in this study confirmed the species identity, which greatly extends the speciesʼ range southward on the eastern coast of Brazil and indicates the presence of an established population of S. chionaraia in the Brazilian Province.     

Scorched mussels (Brachidontes spp., Bivalvia: Mytilidae) from the tropical and warm‐temperate southwestern Atlantic: the role of the Amazon River in their speciation

Antitropicality is a distribution pattern where closely related taxa are separated by an intertropical latitudinal gap. Two potential examples include Brachidontes darwinianus (south eastern Brazil to Uruguay), considered by some authors as a synonym of B. exustus (Gulf of Mexico and the Caribbean), and B. solisianus, distributed along the Brazilian coast with dubious records north of the intertropical zone. Using two nuclear (18S and 28S rDNA) and one mitochondrial gene (mtDNA COI), we aimed to elucidate the phylogeographic and phylogenetic relationships among the scorched mussels present in the warm‐temperate region of the southwest Atlantic. We evaluated a divergence process mediated by the tropical zone over alternative phylogeographic hypotheses. Brachidontes solisianus was closely related to B. exustus I, a species with which it exhibits an antitropical distribution. Their divergence time was approximately 2.6 Ma, consistent with the intensification of Amazon River flow. Brachidontes darwinianus, an estuarine species is shown here not to be related to this B. exustus complex. We suspect ancestral forms may have dispersed from the Caribbean to the Atlantic coast via the Trans‐Amazonian seaway (Miocene). The third species, B rodriguezii is presumed to have a long history in the region with related fossil forms going back to the Miocene. Although scorched mussels are very similar in appearance, their evolutionary histories are very different, involving major historical contingencies as the formation of the Amazon River, the Panama Isthmus, and the last marine transgression.     

An evaluation of the use of branching fire-corals (Millepora spp.) as refuge by reef fish in the Abrolhos Bank, eastern Brazil

Branching scleractinians provide fish with critical refuge against predation. While in the Caribbean and Indo-Pacific regions they are conspicuous elements of the reef community, in the tropical Southwestern Atlantic (Brazil) branching corals are lacking and massive forms dominate. The only branching forms that occur in Brazil are stinging fire-corals of genus Millepora (Hydrozoa). Preliminary observations showed that several fish species seek refuge within fire-corals in Brazil, suggesting that they may replace the functional role played by scleractinians in other regions. Here, the association of fishes to individual fire-coral colonies (Millepora alcicornis and M. brasiliensis) and the relationship between fire-coral cover and fish abundance was studied in eastern Brazil. A total of 38 fish species from 18 families (37?% of the regional species pool) were recorded within fire-corals. Most records were of relatively small-bodied fishes and juveniles of large-bodied fishes. Comparative observations between day and night indicated that fire-corals are used as sleeping sites by a few species. Volume of colonies was positively correlated with fish abundance and species richness. Over a large spatial scale (tens of kilometers) fire-coral cover had a positive and significant influence on the abundance of three fish species only (Abudefduf saxatilis, Acanthurus coeruleus and Microspatodon chrysurus). Although physical attributes of colonies and fire-coral cover may help to account for some of the spatial variability in fish abundance, fire-corals may be considered as an alternative instead of a critical microhabitat for fish. The advantages and drawbacks of living in a branching and stinging environment are discussed.     

Titi monkey biogeography: Parallel Pleistocene spread by Plecturocebus and Cheracebus into a post‐Pebas Western Amazon

Titi monkeys, subfamily Callicebinae, are a diverse, species‐rich group of Neotropical primates with an extensive range across South America. Their distribution in space and time makes them an interesting primate model for addressing questions of Neotropical historical biogeography. Our aim was to reconstruct the biogeographic history of Callicebinae to better understand their diversification patterns and the history of their colonisation of South America since the late Miocene. We reconstructed a time‐calibrated phylogeny of 19 titi species under Bayesian inference using two mitochondrial and 11 nuclear loci. Species were assigned across eight Neotropical areas of endemism, and statistical biogeographic methods implemented in BioGeoBEARS were employed to estimate ancestral areas using 12 biogeographic models. Our results indicate that the most recent common ancestor to extant titi monkeys was widespread from the present‐day Andean foothills in the Colombian Amazon, through the wet and dry savannas of Bolivia and Brazil, to the southern Atlantic forest of eastern Brazil. Genus‐level divergences were characterised by vicariance of ancestral range in the late Miocene. Species‐level diversification in Cheracebus and the Plecturocebus moloch group occurred as they spread across the Amazon in the Pleistocene and were largely characterised by a sequential, long‐distance “island‐hopping” dispersal model of speciation from a narrow area of origin through jump dispersal across rivers. This study comprises the first large‐scale investigation of the evolutionary history of titi monkeys in the context of Amazonian and South American historical biogeography and sheds light on the processes that generated the great diversity found among Callicebinae.     

Latitudinal gradients in Atlantic reef fish communities: trophic structure and spatial use patterns

Trophic strategies and spatial use habits were investigated in reef fish communities. The results supported the hypothesis of differential use of food resources among tropical and higher latitude reef fishes, i.e . the number of species and relative abundance of fishes relying on relatively low‐quality food significantly decreased from tropical to temperate latitudes. The species : genus ratio of low‐quality food consumers increased toward the tropics, and was higher than the overall ratio considering all fishes in the assemblages. This supports the view that higher speciation rates occurred among this guild of fishes in warm waters. It was also demonstrated that density of herbivorous fishes (the dominant group relying on low‐quality food resources) in the western Atlantic decreased from tropical to temperate latitudes. Spatial use and mobility varied with latitude and consequently reef type and complexity. Fishes with small‐size home ranges predominated on tropical coral reefs.     

Biogeography and the structure of coral reef fish communities on isolated islands

Aim To determine the applicability of biogeographical and ecological theory to marine species at two remote island locations. This study examines how biogeography, isolation and species geographic range size influence patterns of species richness, endemism, species composition and the abundance of coral reef fishes. Location Christmas Island and the Cocos (Keeling) Islands in the tropical eastern Indian Ocean. Methods Published species lists and underwater visual surveys were used to determine species richness, endemism, species composition and abundance of reef fishes at the islands. These data were statistically compared with patterns of species composition and abundance from the neighbouring ‘mainland’ Indonesian region. Results The two isolated reef fish communities were species‐poor and contained a distinct taxonomic composition with an overrepresentation of species with high dispersal potential. Despite low species richness, we found no evidence of density compensation, with population densities on the islands similar to those of species‐rich mainland assemblages. The mix of Indian and Pacific Ocean species and the proportional representations of the various regional faunas in the assemblages were not influenced by the relative proximity of the islands to different biogeographical provinces. Moreover, species at the edge of their range did not have a lower abundance than species at the centre of their range, and endemic species had substantially higher abundances than widespread species. At both locations, endemism was low (less than 1.2% of the community); this may be because the locations are not sufficiently isolated or old enough to promote the evolution of endemic species. Main conclusions The patterns observed generally conform to terrestrial biogeographical theory, suggesting that similar processes may be influencing species richness and community composition in reef fish communities at these remote islands. However, species abundances differed from typical terrestrial patterns, and this may be because of the life history of reef fishes and the processes maintaining isolated populations.     

Composition and structure of reef fish communities in Paraíba State,north‐eastern Brazil

The composition and trophic structure of reef fish communities in two natural and two artificial reefs along the coast of Paraíba State in north‐eastern Brazil were investigated. A total of 114 species of fish belonging to 47 families were recorded during 120 stationary visual surveys, slightly less than half (46·55%) of which were recorded at all four surveyed localities. Most species are widely distributed on the western Atlantic coast, but several are endemic to Brazil. The greatest diversity and equitability indexes were recorded at the reefs of Sapatas and Cabeço dos Cangulos, whereas the greatest richness and abundance were found at the Queimado shipwreck. The Alvarenga shipwreck reef had the least richness, diversity and equitability. The four localities studied had very similar ichthyofaunas, especially in relation to species composition. The reefs along the Paraíba coast are considered priority conservation areas by the Brazilian Ministry of the Environment, and the information generated by this study will be useful for comparison with other reefs in the region and can be directly applied to programmes seeking to protect and manage these environments.     

Regional biogeography of shallow reef fish and macro-invertebrate communities in the Galapagos archipelago

Aim To delineate biogeographical patterns in Galapagos shallow‐water reef fauna at regional scales. Location Galapagos Islands. Methods Fishes and macro‐invertebrates were quantitatively censused using underwater visual techniques along more than 500 transects at defined depth strata across the Galapagos archipelago. Data were analysed using multivariate techniques to define regional patterns and identify species typical of different regions. Results Subtidal communities of fishes and macro‐invertebrates on shallow reefs differed consistently in species composition across the Galapagos archipelago, with three major biogeographical groupings: (1) the ‘far‐northern area’ containing the islands of Darwin and Wolf, (2) the ‘central/south‐eastern area’, including the east coast of Isabela, and (3) the ‘western area’, encompassing Fernandina and western Isabela. In addition, the northern islands of Pinta, Marchena and Genovesa form a separate region in the central/south‐eastern area, and Bahia Elizabeth and Canal Bolivar separate from other parts of the western area. The far‐northern bioregion is characterized by high fish species richness overall, including a high proportion of species of Indo‐Pacific origin. However, very few endemic fishes or species with distributions extending south from Ecuador (‘Peruvian’ species) are present, and the bioregion also possesses relatively low species richness of mobile macro‐invertebrate taxa. By contrast, the ‘western’ bioregion possesses disproportionately high numbers of endemic fish taxa, high numbers of cool‐temperate Peruvian fish species, and high invertebrate species richness, but very few species of Indo‐Pacific origin. The Bahia Elizabeth/Canal Bolivar bioregion possesses more endemic species and fewer species with Peruvian affinities than coasts within the western bioregion. The northern bioregion of Pinta, Marchena and Genovesa represents an overlap zone with affinities to both the far‐northern and south‐eastern islands. The south‐eastern bioregion includes species from a variety of different sources, particularly ‘Panamic’ species with distributions extending north to Central America. Main conclusions On the basis of congruent divisions for reef fish and macro‐invertebrate communities, the Galapagos archipelago can be separated into three major biogeographical areas, two of which can be further subdivided into two regions. Each of these five bioregions possesses communities characterized by a distinctive mix of species derived from Indo‐Pacific, Panamic, Peruvian and endemic source areas. The conservation significance of different regions is not reflected in counts of total species richness. The regions with the lowest overall fish species richness possess a temperate rather than tropical climate and highest levels of endemism.     

Island biogeography of Caribbean coral reef fish

Aim The goal of our study was to test fundamental predictions of biogeographical theories in tropical reef fish assemblages, in particular relationships between fish species richness and island area, isolation and oceanographic variables (temperature and productivity) in the insular Caribbean. These analyses complement an analogous and more voluminous body of work from the tropical Indo‐Pacific. The Caribbean is more limited in area with smaller inter‐island distances than the Indo‐Pacific, providing a unique context to consider fundamental processes likely to affect richness patterns of reef fish. Location Caribbean Sea. Methods We compiled a set of data describing reef‐associated fish assemblages from 24 island nations across the Caribbean Sea, representing a wide range of isolation and varying in land area from 53 to 110,860 km². Regression‐based analyses compared the univariate and combined effects of island‐specific physical predictors on fish species richness. Results We found that diversity of reef‐associated fishes increases strongly with increasing island area and with decreasing isolation. Richness also increases with increasing nearshore productivity. Analyses of various subsets of the entire data set reveal the robustness of the richness data and biogeographical patterns. Main conclusions Within the relatively small and densely packed Caribbean basin, fish species richness fits the classical species–area relationship. Richness also was related negatively to isolation, suggesting direct effects of dispersal limitation in community assembly. Because oceanic productivity was correlated with isolation, however, the related effects of system‐wide productivity on richness cannot be disentangled. These results highlight fundamental mechanisms that underlie spatial patterns of biodiversity among Caribbean coral reefs, and which are probably also are functioning in the more widespread and heterogeneous reefs of the Indo‐Pacific.