Diversification of the sleepers (Gobiiformes: Gobioidei: Eleotridae) and evolution of the root gobioid families.

Eleotridae (sleepers) and five smaller families are the earliest diverging lineages within Gobioidei. Most inhabit freshwaters in and around the Indo-Pacific, but Eleotridae also includes species that have invaded the Neotropics as well as several inland radiations in the freshwaters of Australia, New Zealand, and New Guinea. Previous efforts to infer phylogeny of these families have been based on sets of mitochondrial or nuclear loci and have yielded uncertain resolution of clades within Eleotridae. We expand the taxon sampling of previous studies and use genomic data from nuclear ultraconserved elements (UCEs) to infer phylogeny, then calibrate the hypothesis with recently discovered fossils. Our hypothesis clarifies ambiguously resolved relationships, provides a timescale for divergences, and indicates the core crown Eleotridae diverged over a short period 24.3-26.3 Ma in the late Oligocene. Within Eleotridae, we evaluate diversification dynamics with BAMM and find evidence for an overall slowdown in diversification over the past 35 Ma, but with a sharp increase 3.5 Ma in the genus Mogurnda, a clade of brightly colored species found in the freshwaters of Australia and New Guinea.

Species delineation and systematics of a hemiclonal hybrid complex in Australian freshwaters (Gobiiformes: Gobioidei: Eleotridae: Hypseleotris).

The rivers of southeastern Australia host a species complex within the carp gudgeon genus Hypseleotris that includes parental species and hemiclonal hybrid lineages. These hemiclones can be difficult to distinguish from their parent taxa, making delineation of species unusually difficult. We approach this historical taxonomic problem by using single nucleotide polymorphism (SNP) genotyping to distinguish individuals of each species and hemiclones, enabling us to quantify the variation among evolutionary lineages and assign names to the species. Hypseleotris klunzingeri remains valid and does not have any hemiclones. We describe Hypseleotris bucephala and Hypseleotris gymnocephala from the Murray-Darling Basin and Hypseleotris acropinna from the Murray-Darling as well as eastern coastal streams north of the Mary River, part of the range attributed to H. galii. We further split H. galii to distinguish a species from the Mary River, Hypseleotris moolooboolaensis. We designate a neotype and redescribe H. galii due to uncertainty about the source and species identity of specimens used in the original description. We reconcile previous taxonomies, provide new common names for parental species, and advocate using the scientific names of both parents when referring to the hemiclone hybrids to avoid confusion with previous common names that did not distinguish parental taxa and hemiclones.

Prolonged morphological expansion of spiny-rayed fishes following the end-Cretaceous.

Spiny-rayed fishes (Acanthomorpha) dominate modern marine habitats and account for more than a quarter of all living vertebrate species. Previous time-calibrated phylogenies and patterns from the fossil record explain this dominance by correlating the origin of major acanthomorph lineages with the Cretaceous-Palaeogene mass extinction. Here we infer a time-calibrated phylogeny using ultraconserved elements that samples 91.4% of all acanthomorph families and investigate patterns of body shape disparity. Our results show that acanthomorph lineages steadily accumulated throughout the Cenozoic and underwent a significant expansion of among-clade morphological disparity several million years after the end-Cretaceous. These acanthomorph lineages radiated into and diversified within distinct regions of morphospace that characterize iconic lineages, including fast-swimming open-ocean predators, laterally compressed reef fishes, bottom-dwelling flatfishes, seahorses and pufferfishes. The evolutionary success of spiny-rayed fishes is the culmination of multiple species-rich and phenotypically disparate lineages independently diversifying across the globe under a wide range of ecological conditions.

Patterns of Phenotypic Evolution Associated with Marine/Freshwater Transitions in Fishes.

Evolutionary transitions between marine and freshwater ecosystems have occurred repeatedly throughout the phylogenetic history of fishes. The theory of ecological opportunity predicts that lineages that colonize species-poor regions will have greater potential for phenotypic diversification than lineages invading species-rich regions. Thus, transitions between marine and freshwaters may promote phenotypic diversification in trans-marine/freshwater fish clades. We used phylogenetic comparative methods to analyze body size data in nine major fish clades that have crossed the marine/freshwater boundary. We explored how habitat transitions, ecological opportunity, and community interactions influenced patterns of phenotypic diversity. Our analyses indicated that transitions between marine and freshwater habitats did not drive body size evolution, and there are few differences in body size between marine and freshwater lineages. We found that body size disparity in freshwater lineages is not correlated with the number of independent transitions to freshwaters. We found a positive correlation between body size disparity and overall species richness of a given area, and a negative correlation between body size disparity and diversity of closely related species. Our results indicate that the diversity of incumbent freshwater species does not restrict phenotypic diversification, but the diversity of closely related taxa can limit body size diversification. Ecological opportunity arising from colonization of novel habitats does not seem to have a major effect in the trajectory of body size evolution in trans-marine/freshwater clades. Moreover, competition with closely related taxa in freshwaters has a greater effect than competition with distantly related incumbent species.

Phylogeny, diversification, and biogeography of a hemiclonal hybrid system of native Australian freshwater fishes (Gobiiformes: Gobioidei: Eleotridae: Hypseleotris).

BACKGROUND: Carp gudgeons (genus Hypseleotris) are a prominent part of the Australian freshwater fish fauna, with species distributed around the western, northern, and eastern reaches of the continent. We infer a calibrated phylogeny of the genus based on nuclear ultraconserved element (UCE) sequences and using Bayesian estimation of divergence times, and use this phylogeny to investigate geographic patterns of diversification with GeoSSE. The southeastern species have hybridized to form hemiclonal lineages, and we also resolve relationships of hemiclones and compare their phylogenetic placement in the UCE phylogeny with a hypothesis based on complete mitochondrial genomes. We then use phased SNPs extracted from the UCE sequences for population structure analysis among the southeastern species and hemiclones. RESULTS: Hypseleotris cyprinoides, a widespread euryhaline species known from throughout the Indo-Pacific, is resolved outside the remainder of the species. Two Australian radiations comprise the bulk of Hypseleotris, one primarily in the northwestern coastal rivers and a second inhabiting the southeastern region including the Murray-Darling, Bulloo-Bancannia and Lake Eyre basins, plus coastal rivers east of the Great Dividing Range. Our phylogenetic results reveal cytonuclear discordance between the UCE and mitochondrial hypotheses, place hemiclone hybrids among their parental taxa, and indicate that the genus Kimberleyeleotris is nested within the northwestern Hypseleotris radiation along with three undescribed species. We infer a crown age for Hypseleotris of 17.3 Ma, date the radiation of Australian species at roughly 10.1 Ma, and recover the crown ages of the northwestern (excluding H. compressa) and southeastern radiations at 5.9 and 7.2 Ma, respectively. Range-dependent diversification analyses using GeoSSE indicate that speciation and extinction rates have been steady between the northwestern and southeastern Australian radiations and between smaller radiations of species in the Kimberley region and the Arnhem Plateau. Analysis of phased SNPs confirms inheritance patterns and reveals high levels of heterozygosity among the hemiclones. CONCLUSIONS: The northwestern species have restricted ranges and likely speciated in allopatry, while the southeastern species are known from much larger areas, consistent with peripatric speciation or allopatric speciation followed by secondary contact. Species in the northwestern Kimberley region differ in shape from those in the southeast, with the Kimberley species notably more elongate and slender than the stocky southeastern species, likely due to the different topographies and flow regimes of the rivers they inhabit.

Delayed Adaptive Radiation among New Zealand Stream Fishes: Joint Estimation of Divergence Time and Trait Evolution in a Newly Delineated Island Species Flock.

Adaptive radiations are generally thought to occur soon after a lineage invades a region offering high levels of ecological opportunity. However, few adaptive radiations beyond a handful of exceptional examples are known, so a comprehensive understanding of their dynamics is still lacking. Here, we present a novel case of an island species flock of freshwater fishes with a radically different tempo of adaptive history than that found in many popular evolutionary model systems. Using a phylogenomic data set combined with simultaneous Bayesian estimation of divergence times and trait-based speciation and extinction models, we show that the New Zealand Gobiomorphus gudgeons comprise a monophyletic assemblage, but surprisingly, the radiation did not fully occupy freshwater habitats and explosively speciate until more than 10 myr after the lineage invaded the islands. This shift in speciation rate was not accompanied by an acceleration in the rate of morphological evolution in the freshwater crown clade relative to the other species, but is correlated with a reduction in head pores and scales as well as an increase in egg size. Our results challenge the notion that clades always rapidly exploit ecological opportunities in the absence of competing lineages. Instead, we demonstrate that adaptive radiation can experience a slow start before undergoing accelerated diversification and that lineage and phenotypic diversification may be uncoupled in young radiations. [Adaptive radiation; Eleotridae; freshwater; Gobiomorphus; New Zealand.].

Supermatrix phylogeny resolves goby lineages and reveals unstable root of Gobiaria.

Gobies, sleepers, and cardinalfishes represent major clades of a species rich radiation of small bodied, ecologically diverse percomorphs (Gobiaria). Molecular phylogenetics has been crucial to resolving broad relationships of sleepers and gobies (Gobioidei), but the phylogenetic placements of cardinalfishes and nurseryfishes, as reciprocal or sequential sister clades to Gobioidei, are uncertain. In order to evaluate relationships among and within families we used a phylogenetic data mining approach to generate densely sampled trees inclusive of all higher taxa. We utilized conspecific amino acid homology to improve alignment accuracy, included ambiguously identified taxa to increase taxon sampling density, and resampled individual gene alignments to filter rogue sequences before concatenation. This approach yielded the most comprehensive tree yet of Gobiaria, inferred from a sparse (17 percent-complete) supermatrix of one ribosomal and 22 protein coding loci (18,065 characters), comprised of 50 outgroup and 777 ingroup taxa, representing 32 percent of species and 68 percent of genera. Our analyses confirmed the lineage-based classification of gobies with strong support, identified sleeper clades with unforeseen levels of systematic uncertainty, and quantified competing phylogenetic signals that confound resolution of the root topology. We also discovered that multilocus data completeness was related to maximum likelihood branch support, and verified that the phylogenetic uncertainty of shallow relationships observed within goby lineages could largely be explained by supermatrix sparseness. These results demonstrate the potential and limits of publicly available sequence data for producing densely-sampled phylogenetic trees of exceptionally biodiverse groups.

Co-evolution of cleaning and feeding morphology in western Atlantic and eastern Pacific gobies.

Cleaning symbioses are mutualistic relationships where cleaners remove and consume ectoparasites from their clients. Cleaning behavior is rare in fishes and is a highly specialized feeding strategy only observed in around 200 species. Cleaner fishes vary in their degree of specialization, ranging from species that clean as juveniles or facultatively as adults, to nearly obligate or dedicated cleaners. Here, we investigate whether these different levels of trophic specialization correspond with similar changes in feeding morphology. Specifically, we model the evolution of cleaning behavior across the family Gobiidae, which contains the most speciose radiation of dedicated and facultative cleaner fishes. We compared the cranial morphology and dentition of cleaners and non-cleaners across the phylogeny of cleaning gobies and found that facultative cleaners independently evolved four times and have converged on an intermediate morphology relative to that of dedicated cleaners and non-cleaning generalists. This is consistent with their more flexible feeding habits. Cleaner gobies also possess a distinct tooth morphology, which suggests they are adapted for scraping parasites off their clients and show little similarity to other cleaner clades. We propose that evolutionary history and pre-adaptation underlie the morphological and ecological diversification of cleaner fishes.

Morphometric convergence among European sand gobies in freshwater (Gobiiformes: Gobionellidae).

The five genera of sand gobies inhabit the seas and freshwaters of Europe and western Asia and occupy habitats ranging from fully marine to exclusively freshwater. In this study, we use geometric morphometrics to quantify body shape among sand gobies, in order to investigate how shape has evolved and how it is related to habitat. We also compare body shape between preserved museum specimens and fresh specimens, to determine whether or not fixation and storage in ethanol introduce detectable bias. We confirm that the fixed specimens exhibit significant shape changes as compared to fresh specimens, and so, we perform the bulk of our analyses exclusively on fixed specimens. We find that Economidichthys, Orsinigobius, and Pomatoschistus occupy distinct regions of morphospace. Knipowitschia and Ninnigobius have intermediate forms that overlap with Pomatoschistus and Orsinigobius, but not Economidichthys. This pattern is also in rough accordance with their habitats: Pomatoschistus is fully marine, Economidichthys fully freshwater, and the others fresh with some brackish tolerance. We augment a recent phylogeny of sand gobies with data for P. quagga and interpret morphometric shape change on that tree. We then evaluate convergence in form among disparate lineages of freshwater species by constructing a phylomorphospace and applying pattern-based (convevol) measures of convergence. We find that freshwater taxa occupy a mostly separate region of morphospace from marine taxa and exhibit significant convergence in form. Freshwater taxa are characterized by relatively larger heads and stockier bodies than their marine relatives, potentially due to a common pattern of heterochronic size reduction.

Comparison of genetic structure in co-occurring freshwater eleotrids (Actinopterygii: Philypnodon) reveals cryptic species, likely translocation and regional conservation hotspots.

Freshwater systems are naturally fragmented and heterogeneous habitats that promote genetic sub-division and speciation for aquatic biota. Here we provide a novel nuclear genetic perspective (49 allozyme loci) complimented with updated mitochondrial data for the eleotrid genus Philypnodon to investigate broad genetic sub-structure across south-eastern Australia as a foundation for management and conservation. The genus is nominally comprised of two small benthic fishes with contrasting physical and ecological traits, namely the Flathead Gudgeon P. grandiceps and the Dwarf Flathead Gudgeon P. macrostomus. Extensive sample coverage included 99 sites across 5 major drainage divisions and 48 river basins. Nuclear markers revealed strong, geographically-based divergence and sub-structure, contrasting with shallower but largely congruent patterns for mtDNA. The results flag that each nominal species represents a hyper-cryptic species complex, including both broadly distributed and narrow-range taxa, with complicated biogeographic patterns. Predictions on dispersal and genetic structure based on ecological traits were only partially supported and varied by region, with the potential signature of human-assisted translocation evident in several catchments. Further intensive sampling in an important area of high genetic diversity, coastal south-east Queensland, is recommended to better resolve species boundaries and conservation units. The findings provide new insights on regional ecology and biogeography, demonstrating that even supposedly common species can, in reality, have complex conservation and management needs.

Explosive diversification of marine fishes at the Cretaceous-Palaeogene boundary.

The Cretaceous-Palaeogene (K-Pg) mass extinction is linked to the rapid emergence of ecologically divergent higher taxa (for example, families and orders) across terrestrial vertebrates, but its impact on the diversification of marine vertebrates is less clear. Spiny-rayed fishes (Acanthomorpha) provide an ideal system for exploring the effects of the K-Pg on fish diversification, yet despite decades of morphological and molecular phylogenetic efforts, resolution of both early diverging lineages and enormously diverse subclades remains problematic. Recent multilocus studies have provided the first resolved phylogenetic backbone for acanthomorphs and suggested novel relationships among major lineages. However, these new relationships and associated timescales have not been interrogated using phylogenomic approaches. Here, we use targeted enrichment of >1,000 ultraconserved elements in conjunction with a divergence time analysis to resolve relationships among 120 major acanthomorph lineages and provide a new timescale for acanthomorph radiation. Our results include a well-supported topology that strongly resolves relationships along the acanthomorph backbone and the recovery of several new relationships within six major percomorph subclades. Divergence time analyses also reveal that crown ages for five of these subclades, and for the bulk of the species diversity in the sixth, coincide with the K-Pg boundary, with divergences between anatomically and ecologically distinctive suprafamilial clades concentrated in the first 10 million years of the Cenozoic.

Patterns of divergence in fish species separated by the Isthmus of Panama.

BACKGROUND: The Pleistocene closure of Isthmus of Panama, separating the basins of the Eastern Pacific and the Caribbean Sea, created a unique natural experiment that reveals how marine faunas respond to environmental change. To explore how fishes have been affected by this tectonic event, I compare transisthmian patterns in phylogeny and morphology for geminate lineages in two families, Eleotridae (sleepers) and Apogonidae (cardinalfishes). RESULTS: Time-calibrated phylogenies for these families show different diversification patterns. In Eleotridae, several independent shallow instances of transisthmian divergences occur, with one or a few species on either side of the Isthmus. Among Apogonidae, a single clade of Eastern Pacific species is nested within a broad Caribbean radiation that also includes a species known from the Mediterranean. Divergence time estimates for taxa isolated by closure of the Isthmus are broadly congruent. Hypotheses dated with deeper, fossil-based legacy calibrations put the divergences in the Miocene at 7.4-15.1 Ma, while those estimated with a shallow biogeographic calibration of final Isthmus closure range from 5.1 to 9.9 Ma, in the late Miocene/early Pliocene. Eleotridae are more euryhaline than Apogonidae, but do not exhibit shallower transisthmian divergences. In both families, descendent lineages on either side of the Isthmus of Panama exhibit significant shape differences, although that distinction disappears for Apogonidae when I apply a correction for phylogenetic relationships. To evaluate the tempo and mode of continuous character evolution, I fit several single and multiple rate evolutionary models to morphometric data reconstructed on the Apogonidae phylogeny. I find that the most highly favored model, as estimated on both legacy and isthmus calibrated hypotheses, is a multiple rate Ornstein-Uhlbeck model, with a mosaic of rate shifts postulated for shape changes among fishes in the Caribbean and Eastern Pacific. CONCLUSIONS: Although many transisthmian taxa have been compared and their phylogenies calibrated to estimate the dates associated with population sundering, few studies correlate these timing estimates with morphological change. I show that in transisthmian fish lineages, morphometric distinctions are detectable across the Isthmus, and that rates and patterns of shape change have also shifted, with variable manifestations across the body and between the Caribbean and Eastern Pacific.

Molecular phylogeny of Percomorpha resolves Trichonotus as the sister lineage to Gobioidei (Teleostei: Gobiiformes) and confirms the polyphyly of Trachinoidei.

The percomorph fish clade Gobiiformes is a worldwide, tropical and temperate radiation with species occupying nearly all aquatic, and some semi-terrestrial, habitats. Early molecular phylogenetic studies led to the discovery of Gobiiformes, which contains Gobioidei, the gobies and sleepers, and a clade (Apogonoidei) consisting of Apogonidae and Kurtus, the cardinalfishes and nurseryfishes. Gobioidei is consistently resolved as monophyletic in molecular studies, and includes eight families whose members range from waterfall climbing stream gobies to several prominent lineages inhabiting coral reefs. The sister taxon to Gobioidei is also reliably resolved as Apogonoidei. Despite the consistent support for gobiiform monophyly in molecular studies, it is not known if percomorph lineages unsampled in molecular phylogenetic studies are closely related to Gobioidei or Apogonoidei. Here we assemble a large dataset of DNA sequence from ten protein-coding genes, sampling widely across Acanthomorpha and Percomorpha, including Gobioidei, Apogonidae, and Kurtus, along with representatives of all twelve families comprising the former Trachinoidei. The phylogenies inferred from the nuclear gene sequences show that Trachinoidei is polyphyletic, with constituent lineages spread widely among several major percomorph clades. Most notably, the sanddivers (Trichonotus) are resolved as the sister lineage of Gobioidei. This study clarifies the phylogenetic relationships of lineages previously classified in Trachinoidei, identifies Trichonotus as the sister lineage of gobies, provides a molecular phylogeny of the major lineages of Gobioidei, and offers suggested changes to percomorph classification.

Identification of the notothenioid sister lineage illuminates the biogeographic history of an Antarctic adaptive radiation.

BACKGROUND: Antarctic notothenioids are an impressive adaptive radiation. While they share recent common ancestry with several species-depauperate lineages that exhibit a relictual distribution in areas peripheral to the Southern Ocean, an understanding of their evolutionary origins and biogeographic history is limited as the sister lineage of notothenioids remains unidentified. The phylogenetic placement of notothenioids among major lineages of perciform fishes, which include sculpins, rockfishes, sticklebacks, eelpouts, scorpionfishes, perches, groupers and soapfishes, remains unresolved. We investigate the phylogenetic position of notothenioids using DNA sequences of 10 protein coding nuclear genes sampled from more than 650 percomorph species. The biogeographic history of notothenioids is reconstructed using a maximum likelihood method that integrates phylogenetic relationships, estimated divergence times, geographic distributions and paleogeographic history. RESULTS: Percophis brasiliensis is resolved, with strong node support, as the notothenioid sister lineage. The species is endemic to the subtropical and temperate Atlantic coast of southern South America. Biogeographic reconstructions imply the initial diversification of notothenioids involved the western portion of the East Gondwanan Weddellian Province. The geographic disjunctions among the major lineages of notothenioids show biogeographic and temporal correspondence with the fragmentation of East Gondwana. CONCLUSIONS: The phylogenetic resolution of Percophis requires a change in the classification of percomorph fishes and provides evidence for a western Weddellian origin of notothenioids. The biogeographic reconstruction highlights the importance of the geographic and climatic isolation of Antarctica in driving the radiation of cold-adapted notothenioids.

Ecology and evolution affect network structure in an intimate marine mutualism.

Elucidating patterns and causes of interaction among mutualistic species is a major focus of ecology, and recent meta-analyses of terrestrial networks show that network-level reciprocal specialization tends to be higher in intimate mutualisms than in nonintimate mutualisms. It is largely unknown, however, whether this pattern holds for and what factors affect specialization in marine mutualisms. Here we present the first analysis of network specialization ([Formula: see text]) for marine mutualistic networks. Specialization among eight Indo-Pacific networks of obligate mutualistic gobies and shrimps was indistinguishable from that among comparably intimate terrestrial mutualisms (ants-myrmecophytes) and higher than that among nonintimate ones (seed dispersers). Specialization was affected by variability in habitat use for both gobies and shrimps and by phylogenetic history for shrimps. Habitat use was phylogenetically conserved among shrimp, and thus effects of shrimp phylogeny on partner choice were mediated in part by habitat. By contrast, habitat use and pairing patterns in gobies were not related to phylogenetic history. This asymmetry appears to result from evolutionary constraints on partner use in shrimps and convergence among distantly related gobies to utilize burrows provided by multiple shrimp species. Results indicate that the evolution of mutualism is affected by life-history characteristics that transcend environments and that different factors constrain interactions in disparate ecosystems.

Phylogenetic placement of the European sand gobies in Gobionellidae and characterization of gobionellid lineages (Gobiiformes: Gobioidei).

The Mediterranean, northeastern Atlantic, and inland freshwaters of Europe and the Ponto-Caspian region host a distinct fauna of gobiiform fishes, including the sand gobies (Pomatoschistus Gill and related genera), all of which have been classified in the most diverse goby group, the family Gobiidae. Recent molecular phylogenetic analyses have suggested that the sand gobies are not gobiids, and are instead part of their sister clade Gobionellidae (Thacker and Roje 2011). Phylogenetic analysis of Pomatoschistus in the context of both gobiid and gobionellid taxa indicates that Pomatoschistus is part of Gobionellidae, specifically the Mugilogobius lineage. Gobionellidae includes 93 genera, which are arrayed into four lineages (Stenogobius, Mugilogobius, Periophthalmus and Northern Pacific). These lineages exhibit variation in characters of the jaw and suspensorium, including the shapes and relative positions of the palatine, quadrate, and ectopterygoid. The observations of the palatopterygoid complex in Gobionellidae of Harrison (1989) and Larson (2001) are supported and augmented. Gobionellidae generally exhibit suspensoria that are overall more elongated and gracile than those of gobiids: the palatine/ectopterygoid pair features a very short (Periophthalmus lineage) or elongate, pointed palatine (Mugilogobius, Northern Pacific, and Stenogobius lineages), with a relatively slender ectopterygoid and a short quadrate articulation. In Gobiidae, the palatine extends about halfway along the length of the ectopterygoid, and the ectopterygoid generally features a large, flat articulation with the quadrate. Suspensoria of Pomatoschistus and relatives are similar to those of other taxa in the Mugilogobius lineage. Placement of Pomatoschistus and relatives in Gobionellidae rather than Gobiidae is significant in that it indicates that sand gobies are not closely related to other European gobies, and has implications for any comparative evolutionary or biogeographic studies.

Phylogeny and evolution of Indo-Pacific shrimp-associated gobies (Gobiiformes: Gobiidae).

Despite the ubiquity of obligate mutualisms on coral reef ecosystems, little is known about the evolution of many participating species. The shrimp gobies, known primarily from the coral reef habitats of the Indo-Pacific, are small benthic fishes that participate in a remarkable mutualism with alpheid shrimp. In this mutualism, the shrimp build and maintain a burrow that is guarded by the goby, and the shrimp and goby engage in an intricate tactile communication system. The mutualism is obligate for most shrimp gobies as participating species are highly vulnerable to predation when separated from a shrimp partner. We use phylogenetic analysis of nuclear and mitochondrial DNA sequence data to infer evolutionary relationships among shrimp gobies, and between shrimp gobies and their non-mutualistic gobiid relatives. We show that the mutualist shrimp association has arisen twice among gobies, once in a clade composed of Amblyeleotris, Ctenogobiops, and Vanderhorstia, and a second time in a clade including Cryptocentrus, Mahidolia, Tomiamichthys and Stonogobiops. We then compare the evolution of traits within each shrimp goby clade and consider their intrarelationships. We document cryptic diversity among shrimp gobies, with three distinct clades delineated among Mahidolia mysticina specimens captured at the same locality, paired with the same shrimp species. Mahidolia is placed as sister to the Cryptocentrus species Cryptocentrus cinctus; both exhibit pronounced dichromatism, occurring in both brown and yellow (xanthic) forms. We additionally clarify species identities within Amblyeleotris, confirming that widespread similar species Amblyeleotris fasciata, Amblyeleotris steinitzi and Amblyeleotris wheeleri are all distinct. We hypothesize that the flexibility of gobiid gobies and alpheid shrimp to interact with mutualist partners, as well as the apparently highly beneficial nature of mutualism between them, has contributed to the dual evolution of shrimp-association among Indo-Pacific gobies.

Phylogeny of cardinalfishes (Teleostei: Gobiiformes: Apogonidae) and the evolution of visceral bioluminescence.

The cardinalfishes (Apogonidae) are a diverse clade of small, mostly reef-dwelling fishes, for which a variety of morphological data have not yielded a consistent phylogeny. We use DNA sequence to hypothesize phylogenetic relationships within Apogonidae and among apogonids and other acanthomorph families, to examine patterns of evolution including the distribution of a visceral bioluminescence system. In conformance with previous studies, Apogonidae is placed in a clade with Pempheridae, Kurtidae, Leiognathidae, and Gobioidei. The apogonid genus Pseudamia is recovered outside the remainder of the family, not as sister to the superficially similar genus Gymnapogon. Species sampled from the Caribbean and Western Atlantic (Phaeoptyx, Astrapogon, and some Apogon species) form a clade, as do the larger-bodied Glossamia and Cheilodipterus. Incidence of visceral bioluminescence is found scattered throughout the phylogeny, independently for each group in which it is present. Examination of the fine structure of the visceral bioluminescence system through histology shows that light organs exhibit a range of morphologies, with some composed of complex masses of tubules (Siphamia, Pempheris, Parapriacanthus) and others lacking tubules but containing chambers formed by folds of the visceral epithelium (Acropoma, Archamia, Jaydia, and Rhabdamia). Light organs in Siphamia, Acropoma, Pempheris and Parapriacanthus are distinct from but connected to the gut; those in Archamia, Jaydia, and Rhabdamia are simply portions of the intestinal tract, and are little differentiated from the surrounding tissues. The presence or absence of symbiotic luminescent bacteria does not correlate with light organ structure; the tubular light organs of Siphamia and chambered tubes of Acropoma house bacteria, those in Pempheridae and the other Apogonidae do not.

Phylogeography of marine mutualists: parallel patterns of genetic structure between obligate goby and shrimp partners.

The survival of many organisms depends on interspecific, mutualistic interactions. Hence, it is important to assess the phylogeography of multiple mutualistic species simultaneously to gain insight into how their metapopulations persist, spread and recover from disturbance. Consequently, we sequenced mitochondrial DNA (cytochrome b) from a gobiid fish (Ctenogobiops feroculus) and its mutualistic partner, an alpheid snapping shrimp (Alpheus djeddensis). These obligate mutualists are common in shallow coral reef lagoons, and we collected individuals from 11 Indo-Pacific islands that were likely exposed to different disturbance histories due to sea level fluctuations associated with glaciation events [geological studies indicate that eastern oceanic islands (Cook Islands, French Polynesia) were more disturbed than western continental plate islands that have deeper, more extensive lagoons (Okinawa, Fiji)]. Both phylogenetic and population genetic analyses indicated that gobies from Okinawa and Fiji were genetically distinct both from each other and those in the Cook Islands and French Polynesia. Shrimp, by contrast, grouped into two distinct genetic groups that showed no geographic structure. Restricting the analysis to one clade of shrimp, which contained two-thirds of the individuals, showed that these shrimp had very similar phylogeographical structure to the gobies. Mismatch distributions demonstrated that both gobies and shrimp experienced a recent, rapid population expansion into French Polynesia. Finally, nucleotide and haplotype diversities per sample location were significantly correlated between mutualists and tended to be higher on western continental than eastern oceanic islands. Our results suggest that these mutualists recovered from a major disturbance by colonizing South Pacific islands in synchrony.

Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei).

Morphological character analyses indicate that Rhyacichthyidae, Odontobutidae, Eleotridae, and Xenisthmidae are the basal families within the perciform suborder Gobioidei. This study uses DNA sequence data to infer the relationships of genera within these families, as well as determine the placement of more derived gobioids (family Gobiidae) and the identity of the outgroup to Gobioidei. Complete sequences of the mitochondrial ND1, ND2, COI, and cyt b genes (4397 base pairs) are analyzed for representatives of 27 gobioid genera and a variety of perciform and scorpaeniform outgroup candidates; the phylogeny is rooted with a beryciform as a distal outgroup. The single most parsimonious tree that results indicates that, of the outgroups sampled, the perciform family Apogonidae is most closely related to Gobioidei. Gobioidei is monophyletic, and Rhyacichthys aspro is the most basal taxon. The remainder of Gobioidei is resolved into clades corresponding to the families Odontobutidae (plus Milyeringa) and Eleotridae+Xenisthmidae+Gobiidae. Within Eleotridae, the subfamily Butinae (minus Milyeringa) is paraphyletic with respect to Gobiidae, and Eleotrinae is paraphyletic with respect to Xenisthmidae. Other than these groupings, the primary disagreement with the current morphology-based classification is that the molecular data indicate that the troglodytic Milyeringa should be placed in Odontobutidae, not Butinae, although support for this placement is weak. The most basal lineage of Gobioidei is known from the freshwaters of the Indo-Pacific, with marine-dwelling lineages arising several times independently in the group. The phylogeny also indicates that different gobioid lineages are distributed in Asia, Africa, Madagascar and the Neotropics. Five sister pairs of basal gobioid species inhabit Atlantic and Pacific drainages of Panama, with widely varying divergences.