Coral reefs as drivers of cladogenesis: expanding coral reefs, cryptic extinction events, and the development of biodiversity hotspots

Diversification rates within four conspicuous coral reef fish families (Labridae, Chaetodontidae, Pomacentridae and Apogonidae) were estimated using Bayesian inference. Lineage through time plots revealed a possible late Eocene/early Oligocene cryptic extinction event coinciding with the collapse of the ancestral Tethyan/Arabian hotspot. Rates of diversification analysis revealed elevated cladogenesis in all families in the Oligocene/Miocene. Throughout the Miocene, lineages with a high percentage of coral reef-associated taxa display significantly higher net diversification rates than expected. The development of a complex mosaic of reef habitats in the Indo-Australian Archipelago (IAA) during the Oligocene/Miocene appears to have been a significant driver of cladogenesis. Patterns of diversification suggest that coral reefs acted as a refuge from high extinction, as reef taxa are able to sustain diversification at high extinction rates. The IAA appears to support both cladogenesis and survival in associated lineages, laying the foundation for the recent IAA marine biodiversity hotspot.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Influence of the geography of speciation on current patterns of coral reef fish biodiversity across the Indo‐Pacific

The role of speciation processes in shaping current biodiversity patterns represents a major scientific question for ecologists and biogeographers. Hence, numerous methods have been developed to determine the geography of speciation based on co‐occurrence between sister‐species. Most of these methods rely on the correlation between divergence time and several metrics based on the geographic ranges of sister‐taxa (i.e. overlap, asymmetry). The relationship between divergence time and these metrics has scarcely been examined in a spatial context beyond regression curves. Mapping this relationship across spatial grids, however, may unravel how speciation processes have shaped current biodiversity patterns through space and time. This can be particularly relevant for coral reef fishes of the Indo‐Pacific since the origin of the exceptional concentration of biodiversity in the Indo‐Australian Archipelago (IAA) has been actively debated, with several alternative hypotheses involving species diversification and dispersal. We reconstructed the phylogenetic relationships between three species‐rich families of coral reef fish (Chaetodontidae, Labridae, Pomacentridae) and calculated co‐occurrence metrics between closely related lineages of those families. We demonstrated that repeated biogeographic processes can be identified in present‐day species distribution by projecting co‐occurrence metrics between related lineages in a geographical context. Our study also evidence that sister‐species do not co‐occur randomly across the Indo‐Pacific, but tend to overlap their range within the IAA. We identified the imprint of two important biogeographic processes that caused this pattern in 48% of the sister‐taxa considered: speciation events within the IAA and repeated divergence between the Indian and Pacific Ocean, with subsequent secondary contact in the IAA.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Responses of coral reef fishes to past climate changes are related to life‐history traits

Coral reefs and their associated fauna are largely impacted by ongoing climate change. Unravelling species responses to past climatic variations might provide clues on the consequence of ongoing changes. Here, we tested the relationship between changes in sea surface temperature and sea levels during the Quaternary and present‐day distributions of coral reef fish species. We investigated whether species‐specific responses are associated with life‐history traits. We collected a database of coral reef fish distribution together with life‐history traits for the Indo‐Pacific Ocean. We ran species distribution models (SDMs) on 3,725 tropical reef fish species using contemporary environmental factors together with a variable describing isolation from stable coral reef areas during the Quaternary. We quantified the variance explained independently by isolation from stable areas in the SDMs and related it to a set of species traits including body size and mobility. The variance purely explained by isolation from stable coral reef areas on the distribution of extant coral reef fish species largely varied across species. We observed a triangular relationship between the contribution of isolation from stable areas in the SDMs and body size. Species, whose distribution is more associated with historical changes, occurred predominantly in the Indo‐Australian archipelago, where the mean size of fish assemblages is the lowest. Our results suggest that the legacy of habitat changes of the Quaternary is still detectable in the extant distribution of many fish species, especially those with small body size and the most sedentary. Because they were the least able to colonize distant habitats in the past, fish species with smaller body size might have the most pronounced lags in tracking ongoing climate change.     

Species-specific habitat distribution of coral reef fish assemblages in relation to habitat characteristics in an Okinawan coral reef

We determined the species-specific habitat associations of coral reef fishes and environmental characteristics in an Okinawan coral reef in Japan. We focused on three families (Pomacentridae, Gobiidae and Labridae) and attempted to determine differences in habitat utilization. We selected six sites along the coast of Amitori Bay, from the entrance to the innermost part, in order to cover a wide range of habitat characteristics (exposed habitat, semi-exposed habitat and sheltered habitat). The species diversity of coral assemblages was greater at the exposed and semi-exposed habitats, whereas branching coral mostly covered the sheltered habitat. The environmental factors that determine the species-specific spatial association in fishes differed among families. Both biological characteristics (coral morphology and coral species diversity) and physical characteristics (water depth and wave exposure) affected the spatial association of pomacentrids and gobiids. In contrast, physical characteristics such as substrate complexity and water depth affected the species-specific spatial association of labrid species. Further study is needed to determine the ecological factors that regulate the species-specific habitat preference in Okinawan coral reefs.     

Diel differences in the seagrass fish assemblages of a Caribbean island in relation to adjacent habitat types

The fish fauna of Thalassia testudinum (König) seagrass beds was studied at two sites in the Grand Cul-de-Sac Marin Bay (Guadeloupe, French West Indies). The first seagrass bed was located near a coral reef and the second was near coastal mangroves. Both habitats were sampled during day and night, using a purse-seine and a trap net. A total of 98 species belonging to 36 families were observed. Distance-based redundancy analyses revealed two site-specific assemblages of fishes. Diel assemblage shifts were more pronounced in the seagrass beds near coral reefs than in those near mangroves, due to the existence of nocturnal trophic incursions of coral reef fishes into seagrass beds. First-order carnivores dominated the trophic structure of the fish assemblages during both day and night. At night, Haemulidae, Holocentridae and Apogonidae took the place of Labridae, Chaetodontidae and Mullidae present by day near the reef. This switch did not occur near the coast where the exchanges between seagrass beds and mangrove appear to be less important than with the reef ecosystem. Thus, it appears that the adjacent seascape habitat setting affects the intensity in diel variability of the seagrass bed fish community.     

Molecular systematics of the damselfishes (Teleostei: Pomacentridae): Bayesian phylogenetic analyses of mitochondrial and nuclear DNA sequences

Damselfishes in the family Pomacentridae represent one of the few families of reef fishes found on coral reefs irrespective of location. At a local scale, damselfishes are often the most abundant coral reef fish, and their study has provided much of our current understanding of the ecology of tropical reef animals. The study of phylogenetic relationships among the Pomacentridae has lagged ecological investigation of the group, thus limiting historical perspective on the remarkable species richness of the family. In this study, we used 1989bp of DNA sequence representing three mitochondrial genes and 1500bp of the single copy nuclear RAG1 region to infer hypotheses of relationship for the group. Our analysis includes 103 Pomacentridae species in 18 genera, and three of the four named subfamilies: Amphriprioninae, Chrominae, and Pomacentrinae. The Bayesian method of phylogenetic reconstruction was applied to the data, because even with a large number of sequences it is an efficient means of analysis that provides intuitive measures of support for tree topologies and for the parameters of the nucleotide substitution model. Four Pomacentridae clades were identified with high statistical support whether the data were analyzed from a mtDNA, RAG1 or combined perspective, and in all analyses the current subfamilial classification of the Pomacentridae was rejected. At the genus level, Amphiprion, Chromis, and Chrysiptera were also rejected as natural groups. Abudefduf, Amblyglyphidodon, Dascyllus, Neoglyphidodon, Neopomacentrus, and Pomacentrus were each strongly supported as monophyletic genera but the support for monophyly is nonetheless compromised by sample size, except in the case of Dascyllus and Abudefduf for which we have sampled almost all of the described species.     

Length–weight relationships of cryptic reef fishes from the southwestern Gulf of California, Mèxico

Length–weight relationships of 33 cryptic reef fishes from the southwestern Gulf of California were estimated. The most common families were Gobiidae (seven species), Labrisomidae (four), Pomacentridae (three), and Tripterygiidae (three). These are the first length–weight parameters reported for 32 of these species. Specimens were sampled during a comparative ecological study of community structure of fish assemblages associated with coral heads, rocky walls, and artificial reefs in Bahía de La Paz, Baja California Sur. The b values varied between 2.63 and 3.61. New maximum length records for five of the species were established in the study.     

Diversification and functional evolution of reef fish feeding guilds

A core eco‐evolutionary aim is to better understand the factors driving the diversification of functions in ecosystems. Using phylogenetic, trophic, and functional information, we tested whether trophic habits (i.e. feeding guilds) affect lineage and functional diversification in two major radiations of reef fishes. Our results from wrasses (Labridae) and damselfishes (Pomacentridae) do not fully support the ‘dead‐end’ hypothesis that specialisation leads to reduce speciation rates because the tempo of lineage diversification did not substantially vary among guilds in both fish families. Our findings also demonstrate a tight relationship between trophic habits and functional roles held by fish in reef ecosystems, which is not associated with a variation in the tempo of functional diversification among guilds. By illustrating the pivotal importance of the generalist feeding strategy during the evolutionary history of reef fishes, our study emphasises the role of this feeding guild as a reservoir for future diversity.     

Alimentary tract morphology and diet in three coral reef fish families

Alimentary tract morphology and its relationship with diet was examined in three coral reef fish families: Labridae, Chaetodontidae and Pomacentridae. Convergent patterns of dietary‐related morphological variation were evident across the three families, with corallivores possessing long and narrow intestines, carnivores relatively short and wide intestines and herbivores relatively long and wide intestines. Between‐family differences were also evident: chaetodontids typically possessed long and narrow intestines, pomacentrids long and wide intestines and labrids generally possessed short and wide intestines. Overall, intestinal length variation was most strongly attributable to diet, whilst intestinal diameter varied most widely among families. Relative intestinal length appears as a useful predictor of dietary variation, particularly within families. Relative intestinal diameter appears as a less reliable measure, but rather, a useful adjunct to intestinal length comparisons.     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Assessing the diversity and abundances of larvae and juveniles of coral reef fish: a synthesis of six sampling techniques

Due to an increasing emphasis for fish population survey and regulation, efficient tools for evaluating the abundance and diversity of fish from various life stages are needed, especially for coral reef species that present a high taxonomic diversity. The characteristics of six different techniques used for sampling pelagic larvae (a plankton-net and two light-traps), newly settled juveniles (one type of artificial reef), and older juveniles (an underwater seine net in seagrass and macroalgal beds, and rotenone poisoning in coral patches) are described in this study. Larvae belonging to 70 families and juveniles belonging to 34 families were collected. An analysis of similarity (ANOSIM) showed that the taxonomic composition of assemblages collected with the plankton-net and the two light-traps were overlapping but clearly different, due to the higher occurrence of Gobiidae in the plankton-net and of Pomacentridae in both light-traps. Larvae being 2–4 mm standard length (SL) dominated in the plankton-net, whereas larvae being 9–11 mm SL dominated in both light-traps. Pomacentridae juveniles were more abundant in rotenone samples, whereas Labridae dominated in the underwater seine. Juvenile fish collected with the artificial reefs, the underwater seine, and rotenone poisoning largely overlapped in size, with mean sizes of 22, 38, and 33 mm SL, respectively. Seven families were caught by the six sampling techniques, but with unequal success. This study provides ecologists and managers with a unique review of six techniques for sampling a wide range of developmental stages of young fish in different habitats of a coral reef lagoon.

Extended breeding and recruitment periods of fishes on a low latitude coral reef

The temporal dynamics of fish recruitment to equatorial Indo-Pacific coral reefs are not well known. This paper documents fish recruitment over a 2.5-year period in Kimbe Bay (PNG) and shows that it is much less seasonal than is typically described for higher latitude coral reefs. Two families, wrasses (Labridae) and damselfishes (Pomacentridae), which accounted for 90% of all non-cryptic reef fish settlers, exhibited contrasting patterns. Most wrasse species had year-round recruitment with irregular peaks in abundance between November and May. Damselfish species showed a wider range of recruitment patterns, but most had negligible recruitment during the wet season (December–February), followed by one or two recruitment peaks between May and November. Species with longer seasonal recruitment periods exhibited higher cumulative levels of recruitment. For three focal damselfish species, reproductive output was reduced during the wet season, but this alone did not account for the low recruitment at this time. The lack of damselfish recruitment during the wet season is hypothesised to be due to a combination of reduced reproductive output and increased larval mortality associated with monsoonal conditions. The results indicate that there are consistent family-wide recruitment strategies that may play a significant role in the dynamics of populations in equatorial waters.     

Do coral-reef fish faunas have a distinctive taxonomic structure?

Do the highly diverse fish faunas that associate with coral reefs have distinguishing taxonomic and ecological characteristics, as proposed by Choat and Bellwood (1991) and Bellwood (1996)? Does a 50?my old (Eocene) fossil fish fauna from Italy represent a coral-reef fish assemblage that provides unique information about the evolution of such assemblages, as claimed by Bellwood (1996)? I compared the structure of the reef fish faunas of adjacent tropical regions rich and poor in coral reefs, in both America and Polynesia, and found that they exhibit no substantive differences in relative species richness among families of typical “coral-reef” fishes. While coral-rich regions have larger reef fish faunas, a variety of factors probably contribute to such differences. Thus coral-reef fish faunas may lack a distinctive taxonomic structure. A similar comparative approach would be useful for assessing whether assemblages of fishes on coral reefs have distinctive ecological characteristics. Based on patterns of habitat use by modern tropical shorefishes, the Italian Eocene fauna includes few definite reef fishes, and may well consist primarily of non-reef fishes preserved in a non-reef habitat. Until we know more about the environment in which those fossils were preserved, that fauna can contribute little to understanding how coral reef fish assemblages have evolved.     

Species composition,habitat configuration and seasonal changes of coral reef fish assemblages in western Mexico

In spite of their ecological and economic importance, reef fishes from the coast of Oaxaca, Mexico are rarely studied, therefore precluding their management and conservation. In order to identify the set of habitat characteristics/environmental conditions that predict major shifts in fish assemblages in space and time, a stationary census (5′, φ = 5 m) was conducted on a semi‐monthly basis from 2006 to 2009 at patch reefs along the coast. Habitat configuration was gathered using 25 m long point‐intersect transects (data every 25 cm), recording all underlying coral species and substrate characteristics (rocks, sand, algal mats, rubble or dead corals). Recorded were 65 452 fishes grouped in 11 orders, 36 families, 65 genera and 89 species. Labridae (nine species), Pomacentridae (eight species) and Serranidae (seven species) were the most frequent families. Abundance is severely skewed among species; four species Thalassoma lucasanum, Chromis atrilobata, Apogon pacificus and Stegastes acapulcoensis comprise nearly 59% of the fish abundance, 11 species contribute 30%, whereas most of the species (75) can be considered as rare since they contribute <1% each to the total. Species richness and family‐level assemblage composition are similar to those recorded elsewhere in the eastern Pacific. Non‐parametric multivariate analysis of variance demonstrated that changes of diversity metrics might be associated with environmental differences on the scale of hundreds of meters to kilometers, as well as coupled with major changes on oceanographic variables throughout time, exerting meaningful changes on reef‐related fish assemblages.     

Diversity of pectoral fin structure and function in fishes with labriform propulsion

Aquatic propulsion generated by the pectoral fins occurs in many groups of perciform fishes, including numerous coral reef families. This study presents a detailed survey of pectoral fin musculoskeletal structure in fishes that use labriform propulsion as the primary mode of swimming over a wide range of speeds. Pectoral fin morphological diversity was surveyed in 12 species that are primarily pectoral swimmers, including members of all labroid families (Labridae, Scaridae, Cichlidae, Pomacentridae, and Embiotocidae) and five additional coral reef fish families. The anatomy of the pectoral fin musculature is described, including muscle origins, insertions, tendons, and muscle masses. Skeletal structures are also described, including fin shape, fin ray morphology, and the structure of the radials and pectoral girdle. Three novel muscle subdivisions, including subdivisions of the abductor superficialis, abductor profundus, and adductor medialis were discovered and are described here. Specific functional roles in fin control are proposed for each of the novel muscle subdivisions. Pectoral muscle masses show broad variation among species, particularly in the adductor profundus, abductor profundus, arrector dorsalis, and abductor superficialis. A previously undescribed system of intraradial ligaments was also discovered in all taxa studied. The morphology of these ligaments and functional ramifications of variation in this connective tissue system are described. Musculoskeletal patterns are interpreted in light of recent analyses of fin behavior and motor control during labriform swimming. Labriform propulsion has apparently evolved independently multiple times in coral reef fishes, providing an excellent system in which to study the evolution of pectoral fin propulsion.     

Can differences in the structure of larval,juvenile and adult coral‐reef fish assemblages be detected at the family level?

Processes occurring at the end of the larval stage are of major importance in shaping spatial structure of fish assemblages in coral reefs. However, because of the difficulty in identifying larvae to species, many studies dealing with these stages are limited to the family level. It remains unknown if variation in the spatial structure of coral‐reef fish assemblages across life stages can be detected at such a coarse taxonomic level. Two different surveys conducted in a similar area of New Caledonia, Southwest Pacific, provided the opportunity to compare the structure of coral‐reef fish assemblages collected as pre‐settlement larvae, juveniles and adults along a coast to barrier reef gradient. Adult and juvenile fish were sampled using underwater visual counts (UVC) during the warm seasons of 2004 and 2005. Pre‐settlement larvae were sampled with light‐traps during the same seasons. In order to standardize data between sampling methods, analyses were conducted on the relative abundance (for larvae) and density (for juveniles and adults) of 21 families commonly collected with both methods. Relative abundances/densities of families were analysed as a function of life stage (larvae, juveniles or adults), large‐scale spatial location (coast, lagoon or barrier) and years (2004, 2005) using non‐parametric multidimensional scaling (nMDS) and permutational multivariate analysis of variance (permanova ). Kruskal–Wallis tests were then used to examine differences among life stages and locations for individual families. Different levels of spatial and temporal variability characterized fish assemblages from different life stages, and differences among life stages were detected at all locations and years. Differences among life stages were also significant at the level of individual families. Overall results indicate that studies conducted at the family level may efficiently reveal changes in coral‐reef fish spatial structure among successive life stages when large spatial scales are considered.     

Effects of depth and bottom communities on the distribution of highly territorial reef fish in the northwestern region of Cuba

The main goal of the study was to investigate changes in reef fish species abundance and its correlation with selected environmental variables. Three main questions guided the analyses: (i) Do the analytical methods have a significant influence on the results; (ii) What are the main assemblage variation axes; and (iii) What are the factors correlating best with assemblage composition. Highly territorial fish assemblages of several coral reefs along the northwestern region of Cuba were assessed using a stationary point visual census technique. A total of 39 701 individuals of 26 species from the families Chaetodontidae, Labridae and Pomacentridae were counted in 841 censuses; almost 80% of the fish belonged to just four species: bluehead (Thalassoma bifasciatum), bicolor damselfish (Stegastes partitus), slippery dick (Halichoeres bivittatus) and blue chromis (Chromis cyanea). Several multivariate techniques (cluster analysis, non‐metric multidimensional scaling and canonical correspondence analysis) were used to explore main patterns in assemblage composition variation. It was found that the results did not differ significantly when analyzing the same data set. Furthermore, it was shown that the combined use of different multivariate techniques enhanced the interpretation of fish assemblage composition changes. Depth was the main variable explaining variation in the composition of fish assemblages in the studied reefs. The densities of corals and gorgonians were also strongly associated with depth. Sponge density made an additional significant contribution to the explanatory model. Results of this research could be used as a baseline reference for future analyses of the impact of human actions in the study area. (e.g. oil spills, change in fishing intensity, other pollution events, tourism development).     

Local phylogenetic divergence and global evolutionary convergence of skull function in reef fishes of the family Labridae

The Labridae is one of the most structurally and functionally diversified fish families on coral and rocky reefs around the world, providing a compelling system for examination of evolutionary patterns of functional change. Labrid fishes have evolved a diverse array of skull forms for feeding on prey ranging from molluscs, crustaceans, plankton, detritus, algae, coral and other fishes. The species richness and diversity of feeding ecology in the Labridae make this group a marine analogue to the cichlid fishes. Despite the importance of labrids to coastal reef ecology, we lack evolutionary analysis of feeding biomechanics among labrids. Here, we combine a molecular phylogeny of the Labridae with the biomechanics of skull function to reveal a broad pattern of repeated convergence in labrid feeding systems. Mechanically fast jaw systems have evolved independently at least 14 times from ancestors with forceful jaws. A repeated phylogenetic pattern of functional divergence in local regions of the labrid tree produces an emergent family-wide pattern of global convergence in jaw function. Divergence of close relatives, convergence among higher clades and several unusual 'breakthroughs' in skull function characterize the evolution of functional complexity in one of the most diverse groups of reef fishes.