The tropical history and future of the Mediterranean biota and the West African enigma

Aim Since the opening of the Suez Canal in 1869, many tropical taxa from the Indo‐West Pacific (IWP) realm have entered the Mediterranean Sea, which is experiencing rising temperatures. My aims are: (1) to compare biogeographically this tropical transformation of the Mediterranean biota with the tropical faunas of the Mediterranean and adjacent southern European and West African seas during the Late Oligocene to Pliocene interval; (2) to infer the relative contributions of the tropical eastern Atlantic and IWP to the tropical component of the marine biota in southern Europe; and (3) to understand why West Africa is not now a major source of warm‐water species. Location Southern Europe, including the Mediterranean Sea, and the coast of tropical West Africa. Methods I surveyed the literature on fossil and living shell‐bearing molluscs to infer the sources and fates of tropical subgenus‐level taxa living in southern Europe and West Africa during the Late Oligocene to Pliocene interval. Results Ninety‐four taxa disappeared from the tropical eastern Atlantic (including the Mediterranean) but persisted elsewhere in the tropics, mainly in the IWP (81 taxa, 86%) and to a lesser extent in tropical America (36 taxa, 38%). Nine taxa inferred to have arrived in the tropical eastern Atlantic from the west after the Pliocene did not enter the Mediterranean. The modern West African fauna is today isolated from that of other parts of the marine tropics. Main conclusions Taxa now entering the Mediterranean through the Suez Canal are re‐establishing a link with the IWP that last existed 16 million years ago. This IWP element, which evolved under oligotrophic conditions and under a regime of intense anti‐predatory selection, will continue to expand in the increasingly warm and increasingly oligotrophic Mediterranean. The IWP source fauna contrasts with the tropical West African biota, which evolved under productive conditions and in a regime of less anti‐predatory specialization. Until now, the post‐Pliocene West African source area has been isolated from the Mediterranean by cold upwelling. If further warming should reduce this barrier, as occurred during the productive and warm Early Pliocene, the Mediterranean could become a meeting place for two tropical faunas of contrasting source conditions.     

Origins and diversification of Indo-West Pacific marine fauna: evolutionary history and biogeography of turban shells (Gastropoda, Turbinidae)

The present study aimed to assess the consequences of tectonic events and temperature regime on the diversification of Indo‐West Pacific (IWP) turban shell species. Bayesian and parsimony methods were used to construct a phylogenetic hypothesis using sequence data from three genes for the turban shell genus Turbo and for a larger data set including representatives of all known genera in the subfamily Turbininae. Phylogenetic analyses indicate that all IWP Turbo species form a single clade approximately 68 Myr in age, predating the closure of the Tethys Sea and therefore predating the physical separation of the IWP from other biogeographical regions. All but one of the IWP subgenera tested in Turbo also predate the closure of the Tethys Sea. Fossil evidence for Marmarostoma, the largest subgenus, confirms that at least some Turbo lineages currently restricted to the IWP were previously more widespread. The combination of the phylogeny with the fossil evidence suggests that present day diversity in IWP Turbo is the result of the evolutionary persistence within the IWP of several, morphologically distinct lineages, some of which were more widespread in the Oligocene. Some IWP lineages show significant increases in diversification in the early Miocene, probably as a result of the increased availability of both shallow‐water habitats due to tectonic plate movements and increased carbonate platforms in the central IWP resulting from coincident diversification of zooxanthellate corals. The IWP is therefore behaving as both a cradle of diversity (with new species originating in situ) and a museum of diversity (with lineages that predate its isolation also being maintained). Bayesian and parsimony analyses of the subfamily recovered five clades and mapping the temperature regime (tropical or temperate) of each species onto the molecular tree using parsimony suggested that temperate habitat is an ancestral character in at least four of the clades. This result was also supported by Bayesian reconstruction of ancestral states. Astralium (the fifth clade) may also prove to have a temperate origin, but this could not be determined with certainty given the available data. The origin of diversity in tropical regions is of particular interest because it has been suggested that the tropics are the source of many evolutionary novelties and have provided a species pool, from which temperate regions were populated. The present study suggests that Turbininae may be an exception to this rule. The tree shape also suggests that temperature has had an effect on speciation rates; temperate Turbininae are apparently evolving more slowly or suffering more extinction than their tropical sister clades, which show greater diversity. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 573–592.     

Australian barnacles (cirripedia: thoracica), distributions and biogeographical affinities

Currently, 279 barnacle species are recognized in Australia waters. The barnacle fauna of tropical Australia exhibits high species diversity (221), with a high incidence of tropical species (87 Indo-west Pacific [IWP], 16 West Pacific and 65 Indo-Malayan), a low species endemicity (8), and 44 cosmopolitan and 1 Australasian species. Conversely, that of temperate Australia shows lower species diversity (129), with a lower incidence of tropical species (26 IWP, 10 West Pacific and 25 Indo-Malayan), higher species endemicity (23), 37 cosmopolitan, 6 Australasian species, and 3 Australasian/Antarctic species. Distributions corroborate the general patterns demonstrated by the shallow-water biota of northern tropical and southern temperate Australian biogeographic provinces. Tropical and temperate provinces grade into each other in a broad overlap zone along both the western and eastern Australian coasts. This overlap zone is essentially a transitional region, with the gradual replacement of a tropical barnacle fauna in the north by a predominantly temperate barnacle fauna in the south. Both western and eastern Australian coasts are bounded by major poleward-flowing warm currents that have considerable influence on the marine flora and fauna, distributing tropical species of many taxa much farther south than could be predicted by latitude. Currently, 16 barnacle species introduced into Australian waters are identified, although this number may increase in the future due to new port developments and increased shipping arrivals.     

Taxonomic diversity gradients through geological time

Abstract. There is evidence from the fossil record to suggest that latitudinal gradients in taxonomic diversity may be time‐invariant features, although almost certainly not on the same scale as that seen at the present day. It is now apparent that both latitudinal and longitudinal gradients increased dramatically in strength through the Cenozoic era (i.e. the last 65 my) to become more pronounced today than at any time in the geological past. Present‐day taxonomic diversity gradients, in both the marine and terrestrial realms, are underpinned by the tropical radiations of a comparatively small number of species‐rich clades. Quite why these particular taxa proliferated through the Cenozoic is uncertain, but it could be that at least part of the explanation involves the phenomenon of evolutionary escalation. This is, in essence, a theory of biological diversification through evolutionary feedback mechanisms between predators and prey; first one develops an adaptive advantage, and then the other. However, there may also have been some form of extrinsic control on the process of tropical diversification, and this was most likely centred on the phenomenon of global climate change. This is especially so over the last 15 my Various Late Cenozoic (Neogene) vicariant events effectively partitioned the tropics into a series of high diversity centres, or foci. It has been suggested that, in the largest of these in the marine realm (the Indo‐West Pacific or IWP centre), a critical patterns of islands acted as a template for rapid speciation during glacioeustatic sea level cycles. The same process occurred in the Atlantic, Caribbean and East Pacific (ACEP) centre, though on a lesser scale. Tropical terrestrial diversity may also have been promoted by rapid range expansions and contractions in concert with glacial cycles (a modified refugium hypothesis). We are beginning to appreciate that an integrated sequence of Neogene tectonic and climatic events greatly influenced the formation of contemporary taxonomic diversity patterns.     

High dispersal capacity and biogeographic breaks shape the genetic diversity of a globally distributed reef‐dwelling calcifier

Understanding the role of dispersal and adaptation in the evolutionary history of marine species is essential for predicting their response to changing conditions. We analyzed patterns of genetic differentiation in the key tropical calcifying species of large benthic foraminifera Amphistegina lobifera to reveal the evolutionary processes responsible for its biogeographic distribution. We collected specimens from 16 sites encompassing the entire range of the species and analyzed hypervariable fragments of the 18S SSU rDNA marker. We identified six hierarchically organized genotypes with mutually exclusive distribution organized along a longitudinal gradient. The distribution is consistent with diversification occurring in the Indo‐West Pacific (IWP) followed by dispersal toward the periphery. This pattern can be explained by: (a) high dispersal capacity of the species, (b) habitat heterogeneity driving more recent differentiation in the IWP, and (c) ecological‐scale processes such as niche incumbency reinforcing patterns of genotype mutual exclusion. The dispersal potential of this species drives the ongoing range expansion into the Mediterranean Sea, indicating that A. lobifera is able to expand its distribution by tracking increases in temperature. The genetic structure reveals recent diversification and high rate of extinction in the evolutionary history of the clade suggesting a high turnover rate of the diversity at the cryptic level. This diversification dynamic combined with high dispersal potential, allowed the species to maintain a widespread distribution over periods of geological and climatic upheaval. These characteristics are likely to allow the species to modify its geographic range in response to ongoing global warming without requiring genetic differentiation.     

Dispersal barriers in tropical oceans and speciation in Atlantic and eastern Pacific sea urchins of the genus Echinometra

Echinometra is a pantropical sea urchin made famous through studies of phylogeny, speciation, and genetic structure of the Indo-West Pacific (IWP) species. We sequenced 630 bp of the cytochrome oxidase I (COI) mitochondrial gene to provide comparable information on the eastern Pacific and Atlantic species, using divergence between those separated by closure of the Isthmus of Panama 3.1 million years ago (Ma) to estimate dates for cladogenic events. Most recently (1.27-1. 62 Ma), the Atlantic species E. lucunter and E. viridis diverged from each other, at a time in the Pleistocene that sea levels fell and Caribbean coral speciation and extinction rates were high. An earlier split, assumed to have been coincident with the completion of the Isthmus of Panama, separated the eastern Pacific E. vanbrunti from the Atlantic common ancestor. Transisthmian COI divergence similar to that in the sea urchin genus Eucidaris supports this assumption. The most ancient split in Echinometra occurred between the IWP and the neotropical clades, due to cessation of larval exchange around South Africa or across the Eastern Pacific Barrier. Gene flow within species is generally high; however, there are restrictions to genetic exchange between E. lucunter populations from the Caribbean and those from the rest of the Atlantic. Correlation between cladogenic and vicariant events supports E. Mayr's contention that marine species, despite their high dispersal potential, form by means of geographical separation. That sympatric, nonhybridizing E. lucunter and E. viridis were split so recently suggests, however, that perfection of reproductive barriers between marine species with large populations can occur in less than 1.6 million years (Myr).     

Marine shelf habitat: biogeography and evolution

We synthesize the evolutionary implications of recent advances in the fields of phylogeography, biogeography and palaeogeography for shallow‐water marine species, focusing on marine speciation and the relationships among the biogeographic regions and provinces of the world. A recent revision of biogeographic provinces has resulted in the recognition of several new provinces and a re‐evaluation of provincial relationships. These changes, and the information that led to them, make possible a clarification of distributional dynamics and evolutionary consequences. Most of the new conclusions pertain to biodiversity hotspots in the tropical Atlantic, tropical Indo‐West Pacific, cold‐temperate North Pacific, and the cold Southern Ocean. The emphasis is on the fish fauna, although comparative information on invertebrates is utilized when possible. Although marine biogeographic provinces are characterized by endemism and thus demonstrate evolutionary innovation, dominant species appear to arise within smaller centres of high species diversity and maximum interspecies competition. Species continually disperse from such centres of origin and are readily accommodated in less diverse areas. Thus, the diversity centres increase or maintain species diversity within their areas of influence, and are part of a global system responsible for the maintenance of biodiversity over much of the marine world.     

Phenotypic plasticity across 50 MY of evolution: Drosophila wing size and temperature

We studied the response in wing size to rearing at different temperatures of nine strains of Drosophila representing six species. The species varied in their natural habitats from tropical to temperate and one cosmopolitan. The evolutionary divergence of the species spans 50 million years. While some quantitative differences were found, all species responded to temperature very similarly: females increased an average of ∼11% and males ∼14% when reared at 19 °C compared to 25 °C. The phenotypic plasticity in wing size in response to temperature appears to be a fixed trait in Drosophila across long evolutionary time and diverse ecological settings. This likely reflects the close relationship between wing area (and thus wing loading) and insect body mass that is a crucial factor for flight regardless of ecology and is, thus, maintained across long evolutionary time.     

Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae)

Most of the 29 living species of Potamididae show a close association with mangroves. The trees provide the snails with shelter, protection from predators, a solid substrate and sometimes food. Using sequences from three genes (nuclear 18S rRNA and 28S rRNA, mitochondrial COI) we derive a molecular phylogeny and recognize six living genera (Terebralia, Telescopium, Tympanotonos, Cerithidea, Cerithideopsis, Cerithideopsilla). The oldest modern genera (Terebralia, Cerithideopsis) appeared in the Tethyan realm in the Middle Eocene, shortly after the origin of mangrove trees. Whereas most potamidid genera are now restricted to either the Indo-West Pacific (IWP) or to the eastern Pacific plus Atlantic (EPA), sister clades of Cerithideopsis survive in both realms. Based on a reinterpretation of the fossil record (particularly of the monotypic Tympanotonos and extinct Potamides), and parsimonious reconstruction of ancestral habitats, we suggest that the living potamidids are an adaptive radiation that has always been closely associated with mangroves. The specialized tree-climbing groups Cerithidea and Cerithideopsis were independently derived from mud-dwelling ancestors. Cerithideopsilla cingulata (a species complex in the IWP) and 'Potamides' conicus (in the Mediterranean and Indian Ocean) form a single clade within the genus Cerithideopsilla. This refutes the hypothesis that 'P.'conicus is the sole relict of the Tethyan Potamides that has occurred in the Mediterranean region since the Palaeocene. Instead, the phylogeny and fossil record suggest that an ancestor of Cerithideopsilla conica with planktotrophic larvae dispersed from the IWP to the Mediterranean in the Middle Miocene, that its direct development evolved in the Mediterranean during the Pliocene, and that it reinvaded the Indian Ocean during the Plio-Pleistocene.     

Key innovations and the ecology of macroevolution

The origin or evolutionary `success' of taxa is often attributed to key innovations-aspects of organismal phenotype that promote diversification. Different ways of delimiting taxa and measuring `success' (i.e. number or longevity of species, morphological variety or differential control of energy) give rise to different ideas of how key innovations might operate. Key innovations may enhance competitive ability, relax adaptive trade-offs or permit exploitation of a new productive resource base. Recent key innovation studies comparing species richness in extant sister clades may miss important observations possible only with consideration of the fossil record, traditional higher taxa and phenotypic diversity.     

Key innovations and island colonization as engines of evolutionary diversification: a comparative test with the Australasian diplodactyloid geckos

The acquisition of key innovations and the invasion of new areas constitute two major processes that facilitate ecological opportunity and subsequent evolutionary diversification. Using a major lizard radiation as a model, the Australasian diplodactyloid geckos, we explored the effects of two key innovations (adhesive toepads and a snake‐like phenotype) and the invasion of new environments (island colonization) in promoting the evolution of phenotypic and species diversity. We found no evidence that toepads had significantly increased evolutionary diversification, which challenges the common assumption that the evolution of toepads has been responsible for the extensive radiation of geckos. In contrast, a snakelike phenotype was associated with increased rates of body size evolution and, to a lesser extent, species diversification. However, the clearest impact on evolutionary diversification has been the colonization of New Zealand and New Caledonia, which were associated with increased rates of both body size evolution and species diversification. This highlights that colonizing new environments can drive adaptive diversification in conjunction or independently of the evolution of a key innovation. Studies wishing to confirm the putative link between a key innovation and subsequent evolutionary diversification must therefore show that it has been the acquisition of an innovation specifically, not the colonization of new areas more generally, that has prompted diversification.     

Do key innovations unlock diversification? A case-study on the morphological and ecological impact of pharyngognathy in acanthomorph fishes

Key innovations may allow lineages access to new resources and facilitate the invasion of new adaptive zones, potentially influencing diversification patterns. Many studies have focused on the impact of key innovations on speciation rates, but far less is known about how they influence phenotypic rates and patterns of ecomorphological diversification. We use the repeated evolution of pharyngognathy within acanthomorph fishes, a commonly cited key innovation, as a case study to explore the predictions of key innovation theory. Specifically, we investigate whether transitions to pharyngognathy led to shifts in the rate of phenotypic evolution, as well as shifts and/or expansion in the occupation of morphological and dietary space, using a dataset of 8 morphological traits measured across 3,853 species of Acanthomorpha. Analyzing the 6 evolutionarily independent pharyngognathous clades together, we found no evidence to support pharyngognathy as a key innovation; however, comparisons between individual pharyngognathous lineages and their sister clades did reveal some consistent patterns. In morphospace, most pharyngognathous clades cluster in areas that correspond to deeper-bodied morphologies relative to their sister clades, while occupying greater areas in dietary space that reflects a more diversified diet. Additionally, both Cichlidae and Labridae exhibited higher univariate rates of phenotypic evolution compared with their closest relatives. However, few of these results were exceptional relative to our null models. Our results suggest that transitions to pharyngognathy may only be advantageous when combined with additional ecological or intrinsic factors, illustrating the importance of accounting for lineage-specific effects when testing key innovation hypotheses. Moreover, the challenges we experienced formulating informative comparisons, despite the ideal evolutionary scenario of multiple independent evolutionary origins of pharyngognathous clades, illustrates the complexities involved in quantifying the impact of key innovations. Given the issues of lineage specific effects and rate heterogeneity at macroevolutionary scales we observed, we suggest a reassessment of the expected impacts of key innovations may be warranted.     

Species Diversity,Phylogeny and Large Scale Biogeographic Patterns of the Genus Padina (Phaeophyceae,Dictyotales)

The brown algal genus Padina (Dictyotales, Phaeophyceae) is distributed worldwide in tropical and temperate seas. Global species diversity and distribution ranges, however, remain largely unknown. Species‐level diversity was reassessed using DNA‐based, algorithmic species delineation techniques based on cox3 and rbcL sequence data from 221 specimens collected worldwide. This resulted in estimates ranging from 39 to 61 putative species (ESUs), depending on the technique as well as the locus. We discuss the merits, potential pitfalls, and evolutionary and biogeographic significance of algorithmic species delineation. We unveil patterns whereby ESUs are in all but one case restricted to either the Atlantic or Indo‐Pacific Ocean. Within ocean basins we find evidence for the vast majority of ESUs to be confined to a single marine realm. Exceptions, whereby ESUs span up to three realms, are located in the Indo‐Pacific Ocean. Patterns of range‐restricted species likely arise by repeated founder events and subsequent peripatric speciation, hypothesized to dominate speciation mechanisms for coastal marine organisms in the Indo‐Pacific. Using a three‐gene (cox3, psaA and rbcL), relaxed molecular clock phylogenetic analysis we estimated divergence times, providing a historical framework to interpret biogeographic patterns.     

Molecular systematics of cowries (Gastropoda: Cypraeidae) and diversification patterns in the tropics

This study produces a nearly comprehensive phylogeny for the marine gastropod group Cypraeidae (cowries) and uses this topology to examine diversification patterns in the tropics. The dataset is based on molecular sequence data from two mitochondrial genes and includes 210 evolutionary significant units (ESUs) from 170 recognized species (>80%). Systematics for the group is revised based on well‐supported clades, and tree topology is generally consistent with previously proposed classification schemes. Three new genera are introduced ( Cryptocypraea gen. nov , Palmulacypraea gen. nov , and Contradusta gen. nov ) and two previous genera are resurrected (Perisserosa and Eclogavena). One new tribe is proposed (Bistolidini). Topologies produced by a range of transition:transversion (Ti:Tv) weighting schemes in parsimony are pooled and evaluated using maximum likelihood criteria. Extensive geographical coverage shows persistent, large‐scale geographical structure in sister‐groups. Genetic divergence between subspecies is often equivalent or even greater than that between recognized species. Using ESUs as a metric, diversity throughout the Indo‐West Pacific (IWP) increases by 38%. Intra‐ and inter‐regional diversification patterns show that the IWP is the centre for speciation in cowries. The other major tropical regions of the world are inhabited by a predominantly relictual fauna; from a cowrie's eye‐view. Good dispersal ability begets larger ranges, increased extinction resistance and morphological stasis; whereas shorter larval duration results in smaller ranges, higher speciation rates, but also higher turnover. Larval duration and dispersal ability appear correlated with ocean productivity as taxa with longer‐lived larvae are associated with oligotrophic conditions; whereas taxa with shorter larval durations are associated with eutrophic, continental conditions. This tendency is carried to the extreme in temperate or upwelling regions where a planktonic phase is completely lost and crawl‐away larvae evolve multiple times. A strong phylogenetic trend supports these observations as lineages leading up to and including the derived Indo‐West Pacific Erroneinae clade contain taxa predominantly restricted to continental habitats and have undergone the greatest evolutionary radiations in their respective regions. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79, 401– 459.     

Combined community ecology and floristics,a synthetic study on the upper montane evergreen broad-leaved forests in Yunnan,southwestern China

The upper montane evergreen broad-leaved forest in Yunnan occurs mainly in the zone of persistent cloud and has a discontinuous, island-like, distribution. It is diverse, rich in endemic species, and likely to be sensitive to climate change. Six 1-ha sampling plots were established across the main distribution area of the upper montane evergreen broad-leaved forest in Yunnan. All trees with d.b.h. > 1 cm in each plot were identified. Patterns of seed plant distributions were quantified at the specific, generic and family levels. The forests are dominated by the families Fagaceae, Lauraceae, Theaceae and Magnoliaceae, but are very diverse with only a few species shared between sites. Floristic similarities at the family and generic level were high, but they were low at the specific level, with species complementarity between plots. Diversity varied greatly among sites, with greater species richness and more rare species in western Yunnan than central Yunnan. The flora is dominated by tropical biogeographical elements, mainly the pantropic and the tropical Asian distributions at the family and genus levels. In contrast, at the species level, the flora is dominated by the southwest or the southeast China distributions, including Yunnan endemics. This suggests that the flora of the upper montane forest in Yunnan could have a tropical floristic origin, and has adapted to cooler temperatures with the uplift of the Himalayas. Due to great sensitivity to climate, high endemism and species complementarity, as well as the discontinuous, island-like, distribution patterns of the upper montane forest in Yunnan, the regional conservation of the forest is especially needed.     

The marine indo-west pacific break: contrasting the resolving power of mitochondrial and nuclear genes

Simultaneous studies of both nuclear and mitochondrial markerswere undertaken in two widespread Indo-West Pacific (IWP) marineinvertebrates to compare and contrast the ability of these markersto resolve genetic structure. In particular, we were interestedin the resolution of a genetic break between the Indian andPacific Oceans due to historical isolation. Sequence variationfrom the nuclear gene encoding myosin heavy chain (MyHC) andthe mitochondrial gene cytochrome oxidase I (COI) were examinedfor the snapping shrimp Alpheus lottini from wide-ranging populationsthroughout the Indian and Pacific Oceans. A previously identifiedgenetic break between oceans based on COI sequences appearsto have been an artifact caused by the inadvertent inclusionof pseudogene sequences; our new COI data provide evidence onlyof a break between IWP and East Pacific populations. Distributionof a single nucleotide polymorphism in MyHC, on the other hand,shows evidence of a cline between Indian and Pacific Oceans.New allozyme and mtDNA sequence data were also obtained forthe starfish Linckia laevigata. Allozyme data show a clear geneticbreak between Indian Ocean populations and Pacific (includingwestern Australian) populations, whereas the distribution ofmtDNA haplotypes shows a region of overlap in the central IWP.Comparisons of our data for both Alpheus and Linckia with datafrom other population genetic studies in the IWP suggest thatnuclear markers (allozymes, sequence data and morphologicalcharacters) may in some instances reveal historical patternsof genetic population structure whereas mtDNA variation betterreflects present day patterns of gene flow.     

GLOBAL SYSTEMATIC AND PHYLOGENETIC ANALYSIS OF SARGASSUM IN THE GULF OF MEXICO,CARIBBEAN AND PACIFIC BASIN

Sargassum is one of the most species‐rich genera in the brown algae with over 400 described species worldwide. The bulk of these species occurs in Pacific‐Indian ocean waters with only a small portion found on the Atlantic side of the Isthmus of Panama. Sargassum also has one of the most subdivided and complex taxonomic systems used within the algae. Systematic distinctions within the genus are further complicated by high rates of phenotypic variability in several key morphological characters. Molecular analyses in such systems should allow testing of systematic concepts while providing insights into speciation and evolutionary patterns. Global molecular phylogenetic analyses using both conserved and variable regions of the Rubisco operon (rbcL and rbcL‐IGS‐rbcS) were performed with species from the Gulf of Mexico, Caribbean, and Pacific basin. Results confirm earlier analyses based on rbcL‐IGS‐rbcS from Pacific species at the subgeneric and sectional level while providing additional insights into the systematics and phylogenetics on a global scale. For example, species east of the Isthmus of Panama form a distinct well‐resolved clade within the tropical subgenus. This result in sharp contrast to traditional systematic treatments but provides a window into the evolutionary history of this genus in the Pacific and Atlantic Ocean basins and a possible means to time speciation events.     

IUCN classification zones concord with, but underestimate, the population genetic structure of the zebra shark Stegostoma fasciatum in the Indo-West Pacific

The Indo-West Pacific (IWP), from South Africa in the western Indian Ocean to the western Pacific Ocean, contains some of the most biologically diverse marine habitats on earth, including the greatest biodiversity of chondrichthyan fishes. The region encompasses various densities of human habitation leading to contrasts in the levels of exploitation experienced by chondrichthyans, which are targeted for local consumption and export. The demersal chondrichthyan, the zebra shark, Stegostoma fasciatum , is endemic to the IWP and has two current regional International Union for the Conservation of Nature (IUCN) Red List classifications that reflect differing levels of exploitation: 'Least Concern' and 'Vulnerable'. In this study, we employed mitochondrial ND4 sequence data and 13 microsatellite loci to investigate the population genetic structure of 180 zebra sharks from 13 locations throughout the IWP to test the concordance of IUCN zones with demographic units that have conservation value. Mitochondrial and microsatellite data sets from samples collected throughout northern Australia and Southeast Asia concord with the regional IUCN classifications. However, we found evidence of genetic subdivision within these regions, including subdivision between locations connected by habitat suitable for migration. Furthermore, parametric F _ST analyses and Bayesian clustering analyses indicated that the primary genetic break within the IWP is not represented by the IUCN classifications but rather is congruent with the Indonesian throughflow current. Our findings indicate that recruitment to areas of high exploitation from nearby healthy populations in zebra sharks is likely to be minimal, and that severe localized depletions are predicted to occur in zebra shark populations throughout the IWP region.     

MODULATION OF VERY-LONG CHAIN (C28) HIGHLY UNSATURATED FATTY ACIDS IN PROROCENTRUM MININUM (DINOPHYCEAE) BY SELENIUM

Sargassum is one of the most species-rich genera in the brown algae with over 400 described species worldwide. The bulk of these species occurs in Pacific-Indian ocean waters with only a small portion found on the Atlantic side of the Isthmus of Panama. Sargassum also has one of the most subdivided and complex taxonomic systems used within the algae. Systematic distinctions within the genus are further complicated by high rates of phenotypic variability in several key morphological characters. Molecular analyses in such systems should allow testing of systematic concepts while providing insights into speciation and evolutionary patterns. Global molecular phylogenetic analyses using both conserved and variable regions of the Rubisco operon ( rbc L and rbc L-IGS-rbcS) were performed with species from the Gulf of Mexico, Caribbean, and Pacific basin. Results confirm earlier analyses based on rbc L-IGS- rbc S from Pacific species at the subgeneric and sectional level while providing additional insights into the systematics and phylogenetics on a global scale. For example, species east of the Isthmus of Panama form a distinct well-resolved clade within the tropical subgenus. This result in sharp contrast to traditional systematic treatments but provides a window into the evolutionary history of this genus in the Pacific and Atlantic Ocean basins and a possible means to time speciation events.     

Molecular phylogenies and historical biogeography of a circumtropical group of gastropods (Genus: Nerita): implications for regional diversity patterns in the marine tropics

To determine how historical processes, namely speciation, extinction, and dispersal, have contributed to regional species diversity patterns across the marine tropics, we examined the biogeographical history of a circumtropical genus of intertidal gastropods. A species-level phylogeny of Nerita, representing approximately 87% of extant species, was developed from 1608bp of mitochondrial (COI and 16S) and nuclear (ATPSalpha) markers. Phylogenetic relationships generally corresponded to prior classifications; however, comprehensive sampling revealed a number of previously undetected ESUs. Using the resulting tree as a framework, we combined geographical distributions and fossil evidence to reconstruct ancestral ranges, produce a time-calibrated chronogram, and estimate diversification rates. Analyses revealed two monophyletic eastern Pacific+Atlantic (EPA) clades, each of which likely split from an Indo-West Pacific (IWP) sister clade prior to an early Miocene Tethys Seaway closure. More recent diversification throughout the IWP appears to have been driven by both vicariance and dispersal events; EPA diversity has been further shaped by speciation across the Central American Seaway prior to its closure and dispersal across the Atlantic. Despite the latter, inter-regional dispersal has been rare, and likely contributes little to regional diversity patterns. Similarly, infrequent transitions into temperate regions combined with reduced diversification rates may explain low diversity in West and South Pacific clades. Since origination, Nerita diversification appears remarkably constant, with the exception of a lag in the late Eocene-early Oligocene and elevated rates in the late Oligocene-early Miocene. However, a comparison among regions suggested that IWP clades have experienced, on average, higher rates of speciation. Fossil evidence indicates that the EPA likely witnessed greater extinction relative to the IWP. We propose that regional differences in species diversity in Nerita have been largely shaped by differential rates of speciation and extinction.