James L. Van Tassell

Recent invasion of the tropical Atlantic by an Indo-Pacific coral reef fish

The last tropical connection between Atlantic and Indian–Pacific habitats closed c. 2 million years ago (Ma), with the onset of cold‐water upwelling off southwestern Africa. Yet comparative morphology indicates more recent connections in several taxa, including reef‐associated gobies (genus Gnatholepis). Coalescence and phylogenetic analyses of mtDNA cytochrome b sequences demonstrate that Gnatholepis invaded the Atlantic during an interglacial period ∼145 000 years ago (d = 0.0054), colonizing from the Indian Ocean to the western Atlantic, and subsequently to the central (∼100 000 years ago) and eastern Atlantic (∼30 000 years ago). Census data show a contemporary range expansion in the northeastern Atlantic linked to global warming.

RAPID SPECIATION AND ECOLOGICAL DIVERGENCE IN THE AMERICAN SEVEN-SPINED GOBIES (GOBIIDAE, GOBIOSOMATINI) INFERRED FROM A MOLECULAR PHYLOGENY

Abstract.— The American seven‐spined gobies (Gobiidae, Gobiosomatini) are highly diverse both in morphology and ecology with many endemics in the Caribbean region. We have reconstructed a molecular phylogeny of 54 Gobio‐somatini taxa (65 individuals) based on a 1646‐bp region that includes the mitochondrial 12S rRNA, tRNA‐Val, and 16S rRNA genes. Our results support the monophyly of the seven‐spined gobies and are in agreement with the existence of two major groups within the tribe, the Gobiosoma group and the Microgobius group. However, they reject the monophyly of some of the Gobiosomatini genera. We use the molecular phylogeny to study the dynamics of speciation in the Gobiosomatini by testing for departures from the constant speciation rate model. We observe a burst of speciation in the early evolutionary history of the group and a subsequent slowdown. Our results show a split among clades into coastal‐estuarian, deep ocean, and tropical reef habitats. Major habitat shifts account for the early significant acceleration in lineage splitting and speciation rate and the initial divergence of the main Gobiosomatini clades. We found that subsequent diversification is triggered by behavior and niche specializations at least in the reef‐associated clades. Overall, our results confirm that the diversity of Gobiosomatini has arisen during episodes of adaptive radiation, and emphasize the importance of ecology in marine speciation.

Biodiversity Assessment of the Fishes of Saba Bank Atoll, Netherlands Antilles

Biodiversity surveys were conducted on Saba Bank, Netherlands Antilles, to assess ichthyofaunal richness and to compare with published surveys of other Caribbean localities. The primary objective was to estimate the total species richness of the Saba Bank ichthyofauna. A variety of sampling techniques was utilized to survey the fish species of both the visually accessible megafauna and the camouflaged and small-sized species comprising the cryptic ichthyofauna. Based on results presented herein, the number of species known on Saba Bank is increased from 42 previously known species to 270 species. Expected species-accumulation curves demonstrate that the current estimate of species richness of fishes for Saba Bank under represents the actual richness, and our knowledge of the ichthyofauna has not plateaued. The total expected fish-species richness may be somewhere between 320 and 411 species. The Saba Bank ichthyofaunal assemblage is compared to fish assemblages found elsewhere in the Caribbean. Despite the absence of shallow or emergent shore habitats like mangroves, Saba Bank ranks as having the eighth highest ichthyofaunal richness of surveyed localities in the Greater Caribbean. Some degree of habitat heterogeneity was evident. Fore-reef, patch-reef, and lagoonal habitats were sampled. Fish assemblages were significantly different between habitats. Species richness was highest on the fore reef, but 11 species were found only at lagoonal sites. A comprehensive, annotated list of the fishes currently known to occur on Saba Bank, Netherland Antilles, is provided and color photographs of freshly collected specimens are presented for 165 of the listed species of Saba Bank fishes to facilitate identification and taxonomic comparison with similar taxa at other localities. Coloration of some species is shown for the first time. Preliminary analysis indicates that at least six undescribed new species were collected during the survey and these are indicated in the annotated list.

A comparative study of two goby shrimp associations in the Caribbean Sea

In Curacao (Netherland Antilles, Caribbean Sea), two species of gobies, Nes longus (N. longus) (Nichols 1914) and Ctenogobius saepepallens (C. saepepallens) (Gilbert & Randall 1968) with their associated shrimp Alpheus floridanus (A. floridanus) (Kingsley 1878) were observed. Data were collected on feeding behaviour, distances from their burrows, and interactions among conspecifics. Results confirm that the partnership of C. saepepallens and its shrimp is facultative, but show differences to former observations. Possible reasons for this are discussed.

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.

Extreme gender flexibility: Using a phylogenetic framework to infer the evolution of variation in sex allocation, phylogeography, and speciation in a genus of bidirectional sex changing fishes (Lythrypnus, Gobiidae)

The genus Lythrypnus is a group of marine gobies that exhibit extreme gender flexibility as bidirectional sex changers. The genus consists of 20 described species and several undescribed species that are distributed in the Americas. Five species have been characterized with respect to sex allocation and gonad morphology. The hormonal, morphological, and behavioral aspects of sex change have been studied extensively for one species, L. dalli. These data, however, have not been interpreted in an evolutionary context because a phylogenetic hypothesis has not previously been proposed for the genus Lythrypnus. We propose the first phylogenetic hypothesis for the genus based on molecular data from three mitochondrial genes (12s, ND2, and Cytb), one nuclear gene (Rag1) and one nuclear intron (S7). We also include three previously undescribed Lythrypnus species. Our results support the monophyly of the genus with L. heterochroma, an Atlantic species, as the basal taxon. After the divergence of L. heterochroma, there are two main clades, one comprised of species distributed in the Atlantic, the other comprised of species distributed in the Pacific. These data indicate an Atlantic origin for the genus, followed by divergence after the closure of the Isthmus of Panama. Our data also support the monophyly of three previously described species complexes, the L. rhizophora complex and L. dalli complex in the Pacific, and the L. mowbrayi complex in the Atlantic. We mapped patterns of sex allocation within this genus onto the fully resolved and supported topology, and found that sexual plasticity and gender flexibility is likely a synapomorphy for the genus. Overall our results create a well-supported framework to understand the phylogeography of the genus, and to interpret the evolution of sex allocation in Lythrypnus gobies.

Invasion of the Indo-Pacific blenny Omobranchus punctatus (Perciformes: Blenniidae) on the Atlantic Coast of Central and South America

We examined 308 specimens of the Indo-Pacific blenniid Omobranchus punctatus deposited in four museum collections, and analyzed data on their collection locations to assess its invasion on the Atlantic coast of Central and South America. This species occurs in shoreline estuarine and marine habitats in the Indo-West Pacific. Previous sampling and recent records in the Tropical West Atlantic from 1930 to 2004 produced 20 records for: Panama, Colombia, Venezuela, Trinidad and Brazil. In this work, we provide data on 17 new records for the Gulfs of Venezuela and Paria in Venezuela, as well as four records for Maranhao and Para states in NE Brazil. The temporal pattern of collections (1930 - 2009) and the proximity of most localities to ports and zones of ship traffic indicate that O. punctatus was initially introduced to the Atlantic by ships travelling from India to Trinidad. Within Brazil the introduction is linked to shipping connected to petroleum platforms. In Maranhao and Para the

Status of Gobiosoma (Teleostei: Gobiidae) from Brazil: description of a new species, redescription of G. hemigymnum , molecular phylogeny of the genus, and key to Atlantic species

It is unclear how many species of Gobiosoma occur in Brazil and what their geographic distributions are. Here we combine data from a comprehensive morphological survey and a molecular analysis to clarify this uncertain taxonomy and place Brazilian Gobiosoma within a phylogenetic framework. Recent collections in Brazil, from the states of Ceará to Santa Catarina, and in Uruguay yielded two allopatric species of Gobiosoma that are distinct in genetics, meristics, morphometrics, scale pattern and coloration. Comparisons were made with types and specimens of Gobiosoma hemigymnum, Garmannia mediocricula, Gobiosoma spilotum and Gobiosoma parri and all other known species of Gobiosoma. We place G. parri in synonomy with G. hemigymnum with a distribution of Rio de Janeiro to Uruguay and Argentina. The northern species, that extends from the states of Espírito Santo to Ceará, is described as a new species, Gobiosoma alfiei. A key to the Atlantic species of Gobiosoma is provided.

Redescription of the goby genus Gobiosoma (Teleostei: Gobiidae: Gobiosomatini), with the synonymy of the genus Enypnias

Previous classifications of the American gobiid genus Gobiosoma (Teleostei: Gobiidae: Gobiosomatini) have rendered the genus polyphyletic or paraphyletic based on our current understanding of the Gobiosomatini phylogeny. We offer a new definition for Gobiosoma that renders the genus monophyletic. Our new classification synonymizes the genus Enypnias with Gobiosoma and provides characters to distinguish Gobiosoma from allied genera, including Elacatinus and Tigrigobius. Redescriptions and colour photographs of the poorly known species Gobiosoma aceras (formerly Enypnias aceras) and Gobiosoma seminudum (formerly Enypnias seminudus) are provided.

Cryptic lineage divergence in marine environments: genetic differentiation at multiple spatial and temporal scales in the widespread intertidal goby Gobiosoma bosc

Abstract The adaptive radiation of the seven‐spined gobies (Gobiidae: Gobiosomatini) represents a classic example of how ecological specialization and larval retention can drive speciation through local adaptation. However, geographically widespread and phenotypically uniform species also do occur within Gobiosomatini. This lack of phenotypic variation across large geographic areas could be due to recent colonization, widespread gene flow, or stabilizing selection acting across environmental gradients. We use a phylogeographic approach to test these alternative hypotheses in the naked goby Gobiosoma bosc, a widespread and phenotypically invariable intertidal fish found along the Atlantic Coast of North America. Using DNA sequence from 218 individuals sampled at 15 localities, we document marked intraspecific genetic structure in mitochondrial and nuclear genes at three main geographic scales: (i) between Gulf of Mexico and Atlantic Coast, (ii) between the west coast of the Florida peninsula and adjacent Gulf of Mexico across the Apalachicola Bay, and (iii) at local scales of a few hundred kilometers. Clades on either side of Florida diverged about 8 million years ago, whereas some populations along the East Cost show divergent phylogroups that have differentiated within the last 200,000 years. The absence of noticeable phenotypic or ecological differentiation among lineages suggests the role of stabilizing selection on ancestral phenotypes, together with isolation in allopatry due to reduced dispersal and restricted gene flow, as the most likely explanation for their divergence. Haplotype phylogenies and spatial patterns of genetic diversity reveal frequent population bottlenecks followed by rapid population growth, particularly along the Gulf of Mexico. The magnitude of the genetic divergence among intraspecific lineages suggests the existence of cryptic species within Gobiosoma and indicates that modes of speciation can vary among lineages within Gobiidae.