Diarmaid Ó Foighil

Hidden Floridian biodiversity: mitochondrial and nuclear gene trees reveal four cryptic species within the scorched mussel, Brachidontes exustus, species complex

The well‐documented Floridian ‘Gulf/Atlantic’ marine genetic disjunction provides an influential example of vicariant cladogenesis along a continental coastline for major elements of a diverse nearshore fauna. We are engaged in a two‐part study that aims to place this disjunction into a regional Caribbean Basin phylogenetic perspective using the scorched mussel Brachidontes exustus as an exemplar. Our first step, documented here, is to thoroughly characterize the genetic structure of Floridian scorched mussel populations using mitochondrial (mt) and nuclear markers. Both sets of markers recovered the expected disjunction involving sister clades distributed on alternate flanks of peninsular Florida and lineage‐specific mt molecular clocks placed its origin in the Pliocene. The two sister clades had distinct population genetic profiles and the Atlantic clade appears to have experienced an evolutionarily recent bottleneck, although plots of the relative estimates of N through time are consistent with its local persistence through the last Ice Age Maximum. Our primary novel result, however, was the discovery that the Gulf/Atlantic disjunction represents but one of three cryptic, nested genetic discontinuities represented in Floridian scorched mussel populations. The most pronounced phylogenetic split distinguished the Gulf and Atlantic sister clades from two additional nested cryptic sister clades present in samples taken from the southern Florida tropical marine zone. Floridian populations of B. exustus are composed of four cryptic taxa, a result consistent with the hypothesis that the Gulf/Atlantic disjunction in this morphospecies is but one of multiple latent regional genetic breakpoints.

Molecular phylogeny of the livebearing Goodeidae (Cyprinodontiformes).

The Goodeinae is a speciose group of viviparous freshwater fishes endemic to the Mesa Central of Mexico. The current taxonomy of the group is based on morphology associated with viviparity and several of the groupings are questionable. We sequenced part of the mitochondrial cytochrome c oxidase subunit I (COI) gene (627bp) and control region (approximately 430bp aligned) of representatives of 36 species (all genera) of goodeid fishes in order to establish phylogenetic relationships among the taxa. Findings support the monophyly of the Goodeidae, the sister-group relationship of the Empetrichthyinae and Goodeinae, and the relationship of Profundulus to the Goodeidae. All goodeine genera but Xenotoca were recovered as monophyletic. Many of the higher-level relationships within the group contradict the findings of previous studies based upon morphology. The rate of molecular change in COI (0.9% per Myr), calibrated with the fossil record and geological data, suggests an approximate age for the Goodeidae of 16.5Myr. The majority of divergence within the Goodeinae appears to have occurred during the Miocene, with subsequent cladogenesis in the Pliocene and Pleistocene. Most recent speciation appears allopatric. River piracy, particularly involving the Rio Ameca basin, has played a significant role in the diversification of the Goodeinae.

Placing the Floridian marine genetic disjunction into a regional evolutionary context using the scorched mussel, Brachidontes exustus, species complex

Abstract The well‐documented Floridian Gulf/Atlantic marine genetic disjunction provides an influential example of presumed vicariant cladogenesis along a continental coastline for major elements of a diverse nearshore fauna. However, it is unclear if this disjunction represents a local anomaly for regionally distributed morphospecies, or if it is merely one of many such cryptic phylogenetic splits that underlay their assumed genetic cohesiveness. We aimed to place the previously characterized scorched mussel Gulf/Atlantic genetic disjunction into a regional phylogenetic perspective by incorporating genotypes of nominal conspecifics sampled throughout the Caribbean Basin as well as those of eastern Pacific potential geminate species. Our results show it to be one of multiple latent regional genetic disjunctions, involving five cryptic Caribbean species, that appear to be the product of a long history of regional cladogenesis. Disjunctions involving three stem lineages clearly predate formation of the Isthmus of Panama and of the Caribbean Sea, although four of the five cryptic species have within‐basin sister relationships. Surprisingly, the Atlantic clade was also found to be widespread in the southern Caribbean, and ancestral demography calculations through time for Atlantic coast‐specific genotypes are consistent with a northward range extension after the last glacial maximum. Our new data seriously undermine the hypothesis of a Floridian vicariant genesis and imply that the scorched mussel Gulf/Atlantic disjunction represents a case of geographic and temporal pseudocongruence. All five Caribbean Basin cryptic species exhibited an intriguing pattern of predominantly allopatric distribution characterized by distinct geographic areas of ecological dominance, often adjoining those of sister taxa. This pattern of distribution is consistent with allopatric speciation origins, coupled with restricted postspeciation range extensions. Several lines of indirect evidence favor the hypothesis that the predominantly allopatric distributions are maintained over evolutionary time scales, primarily by postrecruitment ecological filters rather than by oceanographic barriers to larvalmediated gene flow.

Evolution of asexuality in the cosmopolitan marine clam Lasaea

The marine clam genus Lasaea is unique among marine bivalves in that it contains both sexual and asexual lineages. We employed molecular tools to infer intrageneric relationships of geographically restricted sexual versus cosmopolitan asexual forms. Polymerase chain reaction primers were used to amplify and sequence homologous 624 nucleotide fragments of COIII from polyploid, asexual, direct‐developing individuals representing northeastern Pacific, northeastern Atlantic, Mediterranean, southern Indian Ocean, and Australian populations. DNA sequences also were obtained from the two known diploid congeners, the Australian sexual, indirect developer, Lasaea australis, and an undescribed meiotic Australian direct developer.

Prehistoric inter-archipelago trading of Polynesian tree snails leaves a conservation legacy

Inter-archipelago exchange networks were an important aspect of prehistoric Polynesian societies. We report here a novel genetic characterization of a prehistoric exchange network involving an endemic Pacific island tree snail, Partula hyalina. It occurs in the Society (Tahiti only), Austral and Southern Cook Islands. Our genetic data, based on museum, captive and wild-caught samples, establish Tahiti as the source island. The source lineage is polymorphic in shell coloration and contains a second nominal species, the dark-shelled Partula clara, in addition to the white-shelled P. hyalina. Prehistoric inter-island introductions were non-random: they involved white-shelled snails only and were exclusively inter-archipelago in scope. Partulid shells were commonly used in regional Polynesian jewellery, and we propose that the white-shelled P. hyalina, originally restricted to Tahiti, had aesthetic value throughout these archipelagoes. Demand within the Society Islands could be best met by trading dead shells, but a low rate of inter-archipelago exchange may have prompted the establishment of multiple founder populations in the Australs and Southern Cooks. The alien carnivorous land snail Euglandina rosea has recently devastated populations of all 61 endemic species of Society Island partulid snails. Southern Cooks and Australs P. hyalina now represent the only unscathed wild populations remaining of this once spectacular land snail radiation.

Sympatric Australian Lasaea species (Mollusca: Bivalvia) differ in their ploidy levels, reproductive modes and developmental modes

The cosmopolitan marine bivalve genus kisaea is predominantly composed of highly polyploid asexual lineages with one exception: the diploid, sexual Australian species L. australis. Two undescribed, direct-deireloping congeners co-occur with the indirect-developing L. austruliJ on the rocky intertidal of southeastern Australia. One of these, L. colmani sp. nov., is also diploid and sexual. The other direct-developing congener is an asexual polyploid composed of a variety of clonal lineages. All three sympatric Australian h a e a congeners are morphologically distinguishable, although prodissoconch distinctions are required to separate large polyploid clams from equivalently-sized L. australis. Similarities in mitochondria1 gene sequence and in shell morphology suggest that L. australis and the Australian sytnpatric polyploid clones share an exclusive common ancestor despite differing in developmental mode, ploidy and reproductive mode. However, detailed karyological analyses failed to identify a chromosome set morphologically similar to that of L. australis among the sympatric Australian polypoid complement. We propose that generation of the polyploid Australian clones (presumably by hybridization) was followed by radical karyological rearrangement.

Parallel changes in genital morphology delineate cryptic diversification of planktonic nudibranchs.

The relative roles of geographical and non-geographical barriers in the genesis of genetic isolation are highly debated in evolutionary biology, yet knowing how speciation occurs is essential to our understanding of biodiversity. In the open ocean, differentiating between the two is particularly difficult, because of the high levels of gene flow found in pelagic communities. Here, we use molecular phylogenetics to test the hypothesis that geography is the primary isolating mechanism in a clade of pelagic nudibranchs, Glaucinae. Our results contradict allopatric expectations: the cosmopolitan Glaucus atlanticus is panmictic, whereas the Indo-Pacific Glaucus marginatus contains two pairs of cryptic species with overlapping distributions. Within the G. marginatus species complex, a parallel reproductive change has occurred in each cryptic species pair: the loss of a bursa copulatrix. Available G. marginatus data are most consistent with non-geographical speciation events, but we cannot rule out the possibility of allopatric speciation, followed by iterative range extension and secondary overlap. Irrespective of ancestral range distributions, our results implicate a central role for reproductive character differentiation in glaucinin speciation—a novel result in a planktonic system.

Geographically Widespread, Non-hybridizing, Sympatric Strains of the Hermaphroditic, Brooding Clam Lasaea in the Northeastern Pacific Ocean

We have studied phenotypic variation in six enzymes of Lasaea, a taxonomically complex genus of small brooding clams, from nine northeastern Pacific sites. Each of the individuals examined produced one of five combinations of electromorph patterns. Lasaea phenotypes could be differentiated into two main types, one containing two and the other three phenotype combinations. Samples from each population contained from one to three phenotype combinations and there was no evidence for crossbreeding among phenotypes. These results are strongly at variance with random mating expectations and indicate that the phenotype combinations represent reproductively isolated strains. This is substantiated by a more detailed study of the McNeill Bay, British Columbia, population where both main strains coexist. Electrophoretic characterization of Lasaea from individual 100 cm2 samples of barnacle cover revealed that strains are not spatially segregated. Progeny of (1) pair mating experiments, (2) brooding field individuals...

Triton's trident: cryptic Neogene divergences in a marine clam (Lasaea australis) correspond to Australia's three temperate biogeographic provinces

The southern coast of Australia is composed of three distinct biogeographic provinces distinguished primarily by intertidal community composition. Several ecological mechanisms have been proposed to explain their formation and persistence, but no consensus has been reached. The marine clam Lasaea australis is arguably the most common bivalve on southern Australian rocky shores and occurs in all three provinces. Here, we tested if this species exhibits cryptic genetic structuring corresponding to the provinces and if so, what mechanisms potentially drove its divergence. Variation in two mitochondrial genes (16S and COIII) and one nuclear gene (ITS2) was assayed to test for genetic structuring and to reconstruct the clams phylogenetic history. Our results showed that L. australis is comprised of three cryptic mitochondrial clades, each corresponding almost perfectly to one of the three biogeographic provinces. Divergence time estimates place their cladogenesis in the Neogene. The trident‐like topology and Neogene time frame of L. australis cladogenesis are incongruent with Quaternary vicariance predictions: a two‐clade topology produced by Pleistocene Bass Strait land bridge formation. We hypothesize that the interaction of the Middle Miocene Climate Transition with the specific geography of the southern coastline of Australia was the primary cladogenic driver in this clam lineage. Additional in‐depth studies of the endemic southern Australian marine biota across all three provinces are needed to establish the generality of this proposed older framework for regional cladogenesis.

Molecular and morphological systematics of neustonic nudibranchs (Mollusca : Gastropoda : Glaucidae : Glaucus), with descriptions of three new cryptic species

Abstract. A recent molecular phylogenetic study on Glaucus, a genus of neustonic aeolid nudibranchs, revealed undescribed cryptic diversity. Glaucus atlanticus is sister to the traditional species Glaucus marginatus, which is a complex of four genetically distinct cryptic species (Informal clade ‘Marginatus’). The present paper revises the systematics of Glaucus and provides formal descriptions for three new species in the informal clade ‘Marginatus’ substantiated by species delimitation analyses. Molecular and morphological evidence confirms that the type species of Glaucus, Glaucus atlanticus, has a cosmopolitan subtropical distribution and is characterised by having a uniseriate ceratal arrangement, a penial spine and a longitudinal, medial silver stripe on the sole of the foot. Examination of type material indicates that the name G. marginatus should be retained for the most widespread of these species, found in the Indian and Pacific Oceans. This species is characterised by molecular diagnostic characters as well as the presence of a bursa copulatrix. Glaucus marginatus is sister to the undescribed species Glaucus bennettae, sp. nov., which is found in the South Pacific Ocean and lacks a bursa copulatrix. The other two undescribed species, Glaucus thompsoni, sp. nov. and Glaucus mcfarlanei, sp. nov. are only known from the North Pacific Ocean, and are characterised by molecular diagnostic characters as well as possessing and lacking a bursa copulatrix, respectively. Because sister species of Glaucus differ in their reproductive anatomy, we hypothesise that mating behaviour has played a role in cladogenesis in this group. ZooBank Publication code: http://zoobank.org/References/E352E264-A440-4AF1-8565-B57B7EEE25BC