Molecular genetic assessment of bipolarity in the anchovy genus Engraulis

To test three hypotheses accounting for bipolar distributions in Engraulis , seven of eight taxa (except E. eurystole ) were surveyed with allozymes (34 loci) and 521 bp of the mitochondrial DNA cytochrome b gene. Both allozymes and mtDNA sequences revealed deep separations between New World and Old World anchovies with a mean allozyme genetic distance D  = 1·26 and net mtDNA sequence divergence d  = 15%. These values reflected separations of 5 to 10 million years. Contrary to previous phylogenetic hypotheses, which place north‐east Pacific E. mordax and south‐east Pacific E. ringens as sister taxa, the south‐west Atlantic E. anchoita and E. ringens are most closely related to each other. The north‐east Pacific E. mordax is the closest lineage to the Old World Engraulis , a group of taxa showing low divergences typical of population‐level separations (mean D  = 0·06; mean d  = 0·87%). Bipolarities of sister taxa in the east Atlantic and west Pacific reflect recent dispersals. Bipolarities in the east Pacific and west Atlantic represent paraphyletic taxa in lineages isolated since the Miocene. None of the bipolarities can be attributed to tectonic separations or competitive displacements from the tropics, but the latter situation should be re‐evaluated with comparisons to tropical anchovies of the New World.     

Living in a tilted world: climate change and geography limit speciation in Old World anchovies (Engraulis; Engraulidae)

Historical changes in the distributions of temperate species in response to Milankovitch climate cycles have been well documented in palaeontological studies and recently evaluated with phylogeographical methods. How these cycles influence biological diversity remains a matter of debate. Molecular surveys of terrestrial and freshwater fauna demonstrate glacial refugia in low latitudes and range expansions into high latitudes, but few genetic studies have assessed the corresponding impact on marine fauna. In the present study, mtDNA sequences (N = 84) are surveyed to understand the impact of long‐term climate oscillations on ‘Old World’ anchovies (genus Engraulis), a monophyletic group occurring in north and south temperate zones of the eastern Atlantic and the western Pacific. The analysis of a 521‐bp sequence of mtDNA cytochrome b indicates a late Miocene or Pliocene dispersal from the north‐eastern Pacific (California–Mexico) to the north‐western Pacific (Japan), followed by Pleistocene dispersal from the north‐western Pacific to Europe. Geography mandates that populations in southern Africa and Australia were stepping‐stones for this dispersal. However, neither population occupies an intermediate position in the mtDNA genealogy; both populations are more recently derived from their northern neighbours. Haplotype diversity is high (h = 0.93–0.97) in European, Australian, and Japanese anchovies, but low (h = 0.22) in the southern African population, where all haplotypes are more closely related to European specimens than to each other. These southern populations occupy a precarious position, lacking north–south coastlines that allow range shifts during climatic extremes. Recurring extinctions and episodic recolonizations from northern hemisphere populations are the likely results. In this case, ocean‐climatic changes retard rather than enhance opportunities for evolutionary radiations. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88 , 673–689.     

Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla

Aim Our aims were: (1) to reconstruct a molecular phylogeny of the cephalaspidean opisthobranch genus Bulla, an inhabitant of shallow sedimentary environments; (2) to test if divergence times are consistent with Miocene and later vicariance among the four tropical marine biogeographical provinces; (3) to examine the phylogenetic status of possible Tethyan relict species; and (4) to infer the timing and causes of speciation events. Location Tropical and warm‐temperate regions of the Atlantic, Indo‐West Pacific, Australasia and eastern Pacific. Methods Ten of the 12 nominal species of Bulla were sampled, in a total sample of 65 individuals, together with cephalaspidean outgroups. Phylogenetic relationships were inferred by Bayesian analysis of partial sequences of the mitochondrial cytochrome c oxidase I (COI) and 16S rRNA and nuclear 28S rRNA genes. Divergence times and rates of evolution were estimated using uncorrelated relaxed‐clock Bayesian methods with fossil calibrations (based on literature review and examination of fossil specimens), implemented in beast . The geographical pattern of speciation was assessed by estimating the degree of overlap between sister lineages. Results Four clades were supported: Indo‐West Pacific (four species), Australasia (one species), Atlantic plus eastern Pacific (three species) and Atlantic (two species), with estimated mean ages of 35–46 Ma. Nominal species were monophyletic, but deep divergences were found within one Indo‐West Pacific and one West Atlantic species. Species‐level divergences occurred in the Miocene or earlier. The age of a sister relationship across the Isthmus of Panama was estimated at 7.9–32.1 Ma, and the divergence of a pair of sister species on either side of the Atlantic Ocean occurred 20.4–27.2 Ma. Main conclusions Fossils suggest that Bulla originated in the Tethys realm during the Middle Eocene. Average ages of the four main clades fall in the Eocene, and far pre‐date the 18–19 Ma closure of the Tethys Seaway. This discrepancy could indicate earlier vicariant events, selective extinction or errors of calibration. Similarly, the transisthmian divergence estimate far pre‐dates the uplift of the Panamanian Isthmus at about 3 Ma. Speciation events occurred in the Miocene, consistent with tectonic events in the central Indo‐West Pacific, isolation of the Arabian Sea by upwelling and westward trans‐Atlantic dispersal. Differences in habitat between sister species suggest that ecological speciation may also have played a role. The basal position of the Australasian species supports its interpretation as a Tethyan relict.     

Phylogeography of Emerita analoga (Crustacea,Decapoda, Hippidae), an eastern Pacific Ocean sand crab with long‐lived pelagic larvae

Aim Phylogeographic analyses have confirmed high dispersal in many marine taxa but have also revealed many cryptic lineages and species, raising the question of how population and regional genetic diversity arise and persist in dynamic oceanographic settings. Here we explore the geographic evolution of Emerita analoga, an inter‐tidal sandy beach crab with an exceptionally long pelagic larval phase and wide latitudinal, amphitropical, distribution. We test the hypothesis that eastern Pacific E. analoga constitute a single panmictic population and examine the location(s), timing and cause(s) of phylogeographic differentiation. Location Principally the eastern Pacific Ocean. Methods We sequenced cytochrome c oxidase subunit I (COI) from 742 E. analoga specimens collected between 1997 and 2000 and downloaded homologous sequences of congeners from GenBank. We reconstructed a phylogeny for Emerita species using maximum likelihood and Bayesian methods and estimated times to most recent common ancestors (TMRCAs), using a COI divergence rate of 1% Myr^?1 and timing of closure of the Central American Seaway. We constructed the COI haplotype network of E. analoga using statistical parsimony, calculated population genetic and spatial structure statistics in Arlequin , and estimated the demographic history of E. analoga using Bayesian skyline analysis. Results Population subdivision and allele frequency differences were insignificant among north‐eastern Pacific locations over 2000 km apart (Φ_ST = 0.00, P = 0.70), yet two distinct phylogroups were recovered from the north‐eastern and south‐eastern Pacific (Φ_CT = 0.87, P < 0.001). Amphitropical differentiation of these temperate clades occurred after TMRCA 1.9 ± 0.02 (mean ± SE) Ma and E. analoga has expanded into its present‐day north‐eastern Pacific range since c. 250 ka. Main conclusions Emerita analoga is not panmictic but is very widely dispersed and approaching genetic homogeneity, i.e. ‘eurymixis’, in the north‐eastern Pacific. North‐eastern and south‐eastern Pacific populations of E. analoga probably became isolated c. 1.5 Ma as the tropical eastern Pacific Ocean warmed and expanded, intensifying barriers to gene flow. The fragmentation of a widespread ancestral species previously connected by long‐distance gene flow (‘soft vicariance’) coincident with changing oceanographic conditions may be a common theme in the evolution of Emerita species and in other highly dispersive taxa. Highly dispersive species may differentiate because of, not despite, the dynamic oceanographic setting.     

New insights into halophilic prokaryotes isolated from salting–ripening anchovies (Engraulis anchoita) process focused on histamine-degrading strains

Salted and ripened fish foods are susceptible to cause histamine poisoning. The present study focuses on microbial histamine degradation from high salted fermented fishery products to deepen our understanding about this new and growing field of research. As a result of this first study related to salted–ripened anchovies (Engraulis anchoita), fifty seven moderate and extreme halophilic microbial isolates from salt and salted–ripened anchovy processes were characterized in terms of their phenotype and histamine-degrading capacity. Only 7%—4 isolates—were able to degrade histamine. None of the histamine-degrading isolates presented proteolytic and/or lipolytic activity. One of them designated A18 was chemotactic toward histamine, an interesting property not previously reported for that chemoattractant. However, the S18 and A18 isolates, genotypically identified as Halobacterium sp. and Halomonas sp. respectively, produced indole and/or H₂S, both undesirable characteristics associated to off-flavors occurrence. On the other hand, A28 and S20, identified as Halovibrio sp. and Halobacterium sp. respectively, presented desirable properties, such as cytochrome oxidase and catalase activity, and non-production of H₂S and indole. These strains also showed characteristics previously reported as dominant in the ripened stage. The results are promising, and A28 and S20 may have the desirable features to improve the anchovy salting–ripening process.

     

Phylogeographic patterns of a lower intertidal isopod in the Gulf of California and the Caribbean and comparison with other intertidal isopods

A growing body of knowledge on the diversity and evolution of intertidal isopods across different regions worldwide has enhanced our understanding on biological diversification at the poorly studied, yet vast, sea–land interface. High genetic divergences among numerous allopatric lineages have been identified within presumed single broadly distributed species. Excirolana mayana is an intertidal isopod that is commonly found in sandy beaches throughout the Gulf of California. Its distribution in the Pacific extends from this basin to Colombia and in the Atlantic from Florida to Venezuela. Despite its broad distribution and ecological importance, its evolutionary history has been largely neglected. Herein, we examined phylogeographic patterns of E. mayana in the Gulf of California and the Caribbean, based on maximum‐likelihood and Bayesian phylogenetic analyses of DNA sequences from four mitochondrial genes (16S rDNA, 12S rDNA, cytochrome oxidase I gene, and cytochrome b gene). We compared the phylogeographic patterns of E. mayana with those of the coastal isopods Ligia and Excirolana braziliensis (Gulf of California and Caribbean) and Tylos (Gulf of California). We found highly divergent lineages in both, the Gulf of California and Caribbean, suggesting the presence of multiple species. We identified two instances of Atlantic–Pacific divergences. Some geographical structuring among the major clades found in the Caribbean is observed. Haplotypes from the Gulf of California form a monophyletic group sister to a lineage found in Venezuela. Phylogeographic patterns of E. mayana in the Gulf of California differ from those observed in Ligia and Tylos in this region. Nonetheless, several clades of E. mayana have similar distributions to clades of these two other isopod taxa. The high levels of cryptic diversity detected in E. mayana also pose challenges for the conservation of this isopod and its fragile environment, the sandy shores.     

World travellers: phylogeny and biogeography of the butterfly genus Leptotes (Lepidoptera: Lycaenidae)

Leptotina butterflies (Lycaenidae, Polyommatiinae) are found mostly in tropical and subtropical areas around the globe, marginally penetrating into temperate regions. Here, we investigated phylogenetic and biogeographical relationships of most representatives of the subtribe, using both likelihood and Bayesian approaches. We also estimated the timing of their diversification. And lastly, we studied phylogeographic patterns of the most widespread species, Leptotes pirithous. DNA sequences from two mitochondrial (COI, COII) and two nuclear genes (wingless, Ef1α) were analysed for 13 species of the genus Leptotes Scudder and one species of the genus Cyclyrius Butler. Both genera together form a monophyletic clade, and Cyclyrius is rooted deep inside Leptotes. Therefore, we designate Cyclyrius to be a junior synonym of Leptotes. According to our study, the genus Leptotes originated between the late Eocene and early Oligocene (35–31 Ma). During the Miocene it dispersed to the rest of the southern hemisphere, with further speciation events within the Indo‐Australian region, and separate radiations in the Americas and the Afrotropics. Leptotes webbianus from the Canary Islands turned out to be sister to the American clade from which it split c. 12 Ma. Leptotes pirithous originated in Madagascar c. 4 Ma and invaded the whole of Africa and southern Europe, including numerous surrounding islands. Populations of L. pirithous from Mauritius and Madagascar turned out to represent a distinct species (Leptotes durrelli sp.n. ) and the same applies to the Australasian populations of Leptotes plinius (Leptotes lybas stat. rev. ). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:20308930‐988B‐4327‐A35F‐CC983D46263B .     

Older than New Caledonia emergence? A molecular phylogenetic study of the eneopterine crickets (Orthoptera: Grylloidea)

Aim A New Caledonian insect group was studied in a world‐wide phylogenetic context to test: (1) whether local or regional island clades are older than 37 Ma, the postulated re‐emergence time of New Caledonia; (2) whether these clades show evidence for local radiations or multiple colonizations; and (3) whether there is evidence for relict taxa with long branches in phylogenetic trees that relate New Caledonian species to geographically distant taxa. Location New Caledonia, south‐west Pacific. Methods We sampled 43 cricket species representing all tribes of the subfamily Eneopterinae and 15 of the 17 described genera, focusing on taxa distributed in the South Pacific and around New Caledonia. One nuclear and three mitochondrial genes were analysed using Bayesian and parsimony methods. Phylogenetic divergence times were estimated using a relaxed clock method and several calibration criteria. Results The analyses indicate that, under the most conservative dating scenario, New Caledonian eneopterines are 5–16 million years old. The largest group in the Pacific region dates to 18–29 Ma. New Caledonia has been colonized in two phases: the first around 10.6 Ma, with the subsequent diversification of the endemic genus Agnotecous, and the second with more recent events around 1–4 Ma. The distribution of the sister group of Agnotecous and the lack of phylogenetic long branches in the genus refute an assumption of major extinction events in this clade and the hypothesis of local relicts. Main conclusions Our phylogenetic studies invalidate a simple scenario of local persistence of this group in New Caledonia since 80 Ma, either by survival on the New Caledonian island since its rift from Australia, or, if one accepts the submergence of New Caledonia, by local island‐hopping among other subaerial islands, now drowned, in the region during periods of New Caledonian submergence.     

DNA‐based phylogeny of the marine genus Heterodrilus (Annelida,Clitellata, Naididae)

Heterodrilus is a group of marine Naididae, common worldwide in subtropical and tropical areas, and unique among the oligochaetes by their tridentate chaetae. The phylogenetic relationships within the group are assessed from the nuclear 18S rDNA gene, and the mitochondrial cytochrome c oxidase subunit I (COI) and 16S rDNA genes. Sequence data were obtained from 16 Heterodrilus species and 13 out‐group taxa; 48 sequences are new for this study. The data were analysed by Bayesian inference. Monophyly of the genus is corroborated by the resulting tree, with Heterodrilus ersei (a taxon representing a small group of species with aberrant male genitalia) proposed to be outside all other sampled species. Although earlier regarded as a member of the subfamily Rhyacodrilinae, both molecular and morphological data seem to support that Heterodrilus is closely related to Phallodrilinae. However, the results are not conclusive as to whether the genus is the sister group of, or a group nested inside, or separate from this latter subfamily. The studied sample of species suggests at least two major clades in Heterodrilus with different geographical distributions, in one of the clades, most species are from the Indo‐West Pacific Ocean, while in the other, the majority are from the Western Atlantic Ocean. Morphological characters traditionally used in Heterodrilus taxonomy are optimized on the phylogenetic tree, revealing a high degree of homoplasy.     

Molecular data illuminate cryptic nudibranch species: the evolution of the Scyllaeidae (Nudibranchia: Dendronotina) with a revision of Notobryon

Scyllaeidae represents a small clade of dendronotoid nudibranchs. Notobryon wardi Odhner, 1936, has been reported to occur in tropical oceans from the Indo‐Pacific and eastern Pacific to temperate South Africa. The systematics of Notobryon has not been reviewed using modern systematic tools. Here, specimens of Notobryon were examined from the eastern Pacific, the Indo‐Pacific, and from temperate South Africa. Additionally, representatives of Scyllaea and Crosslandia were studied. Scyllaeidae was found to be monophyletic. Notobryon was also found to be monophyletic and is the sister group to Crosslandia plus Scyllaea. The molecular data also clearly indicate that within Notobryon, at least three distinct species are present, two of which are here described. Genetic distance data indicate that eastern Pacific and South African exemplars are 10–23% divergent from Indo‐Pacific exemplars of Notobryon wardi. Scyllaea pelagica has been regarded as a single, circumtropical species. Our molecular studies clearly indicate that the Atlantic and Indo‐Pacific populations are distinct and we resurrect Scyllaea fulva Quoy & Gaimard, 1824 for the Indo‐Pacific species. Our morphological studies clearly corroborate our molecular findings and differences in morphology distinguish closely related species. Different species clearly have distinct penial morphology. These studies clearly reinforce the view that eastern Pacific, Indo‐Pacific, and temperate biotas consist largely of distinct faunas, with only a minor degree of faunal overlap. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 311–336.     

Freshwater occurrence of the extinct dolphin Parapontoporia (Cetacea: Lipotidae) from the upper Pliocene nonmarine Tulare Formation of California

The diminutive, extinct longirostrine dolphin Parapontoporia is one of the most abundantly represented late Neogene odontocetes from the eastern North Pacific and is widely known from numerous marine strata of late Miocene and Pliocene age in California, Baja California and possibly Japan. Parapontoporia has been identified as the sister taxon of the recently extinct Chinese river dolphin (Lipotes vexillifer); unlike Lipotes, which exclusively inhabited freshwater, the depositional context of Parapontoporia suggests it was marine. A newly identified petrosal of Parapontoporia sp. was preserved alongside terrestrial vertebrates in the nonmarine Tulare Formation (upper Pliocene to Pleistocene, 2.2–0.6 Ma), California, which was deposited under lacustrine and fluviodeltaic conditions. Abundantly preserved freshwater molluscs and rare marine taxa suggest predominantly freshwater settings with intermittent periods of estuarine conditions. This occurrence of Parapontoporia indicates its presence in the San Joaquin basin after the retreat of the inland sea and suggests that this extinct odontocete may have been freshwater tolerant and an inhabitant of marine and freshwater settings, heralding the exclusively freshwater existence of its Recent sister taxon Lipotes vexillifer.     

From Africa via Europe to South America: migrational route of a species‐rich genus of Neotropical lowland rain forest trees (Guatteria,Annonaceae)

Aim Several recent studies have suggested that a substantial portion of today’s plant diversity in the Neotropics has resulted from the dispersal of taxa into that region rather than by vicariance. In general, three routes have been documented for the dispersal of taxa onto the South American continent: (1) via the North Atlantic Land Bridge, (2) via the Bering Land Bridge, or (3) from Africa directly onto the continent. Here a species‐rich genus of Neotropical lowland rain forest trees (Guatteria, Annonaceae) is used as a model to investigate these three hypotheses. Location The Neotropics. Methods The phylogenetic relationships within the long‐branch clade of Annonaceae were reconstructed (using maximum parsimony, maximum likelihood and Bayesian inference) in order to gain insight in the phylogenetic position of Guatteria. Furthermore, Bayesian molecular dating and Bayesian dispersal–vicariance (Bayes‐DIVA) analyses were undertaken. Results Most of the relationships within the long‐branch clade of Annonaceae were reconstructed and had high support. However, the relationship between the Duguetia clade, the Xylopia–Artabotrys clade and Guatteria remained unclear. The stem node age estimate of Guatteria ranged between 49.2 and 51.3 Ma, whereas the crown node age estimate ranged between 11.4 and 17.8 Ma. For the ancestral area of Guatteria and its sister group, the area North America–Africa was reconstructed in 99% of 10,000 DIVA analyses, while South America–North America was found just 1% of the time. Main conclusions The estimated stem to crown node ages of Guatteria in combination with the Bayes‐DIVA analyses imply a scenario congruent with an African origin followed by dispersal across the North Atlantic Land Bridge in the early to middle Eocene and further dispersal into North and Central America (and ultimately South America) in the Miocene. The phylogenetically and morphologically isolated position of the genus is probably due to extinction of the North American and European stem lineages in the Tertiary.     

New Systematic Insights in the Ceramium Sinicolacomplex: Resurrection of C. Interruptum S. & G. (Ceramiaceae,Rhodophyta)

Setchell & Gardner (1924) provided a taxonomic treatment for the genus Ceramium from lower California and the Gulf of California, Mexico, in which they described several new species, including C. sinicola S. & G. described from Ensenada, BCN and C. interruptum S. & G. described from the vicinity of La Paz, BCS. The latter was later reduced to variety rank in C. sinicolaby Dawson (1950), a taxonomic decision that has subsequently been widely adopted. Phylogenetic analyses inferred from three molecular markers (chloroplast‐encoded rbcL, the RUBISCO spacer, and nuclear encoded SSU rDNA) from recent collections from the Pacific North‐west (California, Oregon) and the Gulf of California reveal a well supported assemblage of three corticated taxa: C. codicolaJ. Ag. 1894, C. sinicola and C. sinicola var. interruptum. Sequence divergence values among the three taxa are sufficient to warrant separate species ranking; hence, we reinstate C. interruptum as a widely distributed epiphyte for the region. C. sinicolais more closely related to C. codicolathan to C. interruptum, with the former two restricted to the host Codium. The molecule‐based relationships are congruent with evolutionary trends in cortication pattern and attachment mode.     

Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long‐distance dispersal,but not West Gondwana

Aim The biogeography of the tropical plant family Monimiaceae has long been thought to reflect the break‐up of West and East Gondwana, followed by limited transoceanic dispersal. Location Southern Hemisphere, with fossils in East and West Gondwana. Methods We use phylogenetic analysis of DNA sequences from 67 of the c. 200 species, representing 26 of the 28 genera of Monimiaceae, and a Bayesian relaxed clock model with fossil prior constraints to estimate species relationships and divergence times. Likelihood optimization is used to infer switches between biogeographical regions on the highest likelihood tree. Results Peumus from Chile, Monimia from the Mascarenes and Palmeria from eastern Australia/New Guinea form a clade that is sister to all other Monimiaceae. The next‐deepest split is between the Sri Lankan Hortonia and the remaining genera. The African Monimiaceae, Xymalos monospora, then forms the sister clade to a polytomy of five clades: (I) Mollinedia and allies from South America; (II) Tambourissa and allies from Madagascar and the Mascarenes; (III) Hedycarya, Kibariopsis and Leviera from New Zealand, New Caledonia and Australia; (IV) Wilkiea, Kibara, Kairoa; and (V) Steganthera and allies, all from tropical Australasia. Main conclusions Tree topology, fossils, inferred divergence times and ances‐tral area reconstruction fit with the break‐up of East Gondwana having left a still discernible signature consisting of sister clades in Chile and Australia. There is no support for previous hypotheses that the break‐up of West Gondwana (Africa/South America) explains disjunctions in the Monimiaceae. The South American Mollinedia clade is only 28–16 Myr old, and appears to have arrived via trans‐Pacific dispersal from Australasia. The clade apparently spread in southern South America prior to the Andean orogeny, fitting with its first‐diverging lineage (Hennecartia) having a southern‐temperate range. The crown ages of the other major clades (II–V) range from 20 to 29 Ma, implying over‐water dispersal between Australia, New Caledonia, New Zealand, and across the Indian Ocean to Madagascar and the Mascarenes. The endemic genus Monimia on the Mascarenes provides an interesting example of an island lineage being much older than the islands on which it presently occurs.     

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.     

Two new species of Phyllonema (Rivulariaceae,Cyanobacteria) with an emendation of the genus

Two untapered, heterocytous species were observed and collected from the intertidal and supratidal zones of the Mexican coastline of the Pacific Ocean near Oaxaca and from the Gulf of Mexico. These populations were highly similar in morphology to the freshwater taxon Petalonema incrustans in the Scytonemataceae. However, 16S rRNA sequence data and phylogenetic analysis indicated that they were sister taxa to the epiphyllic, Brazilian species Phyllonema aveceniicola in the Rivulariaceae, described from culture material. While genetic identity between the two new species was high, they differed significantly in morphology, 16S rRNA gene sequence identity, and sequence and structure of the 16S–23S ITS region. Their morphology differed markedly from the generitype of the previously monotypic Phyllonema, which has tapered, heteropolar, single‐false branched trichomes with very thin or absent sheath. The two new species, Phyllonema ansata and Phyllonema tangolundensis, described from both culture and environmental material, have untapered, isopolar, geminately false branched trichomes with thick, lamellated sheaths, differences so significant that the species would not be placed in Phyllonema without molecular corroboration. The morphological differences are so significant that a formal emendation of the genus is required. These taxa provide a challenge to algal taxonomy because the morphological differences are such that one would logically conclude that they represent different genera, but the phylogenetic evidence for including them all in the same genus is conclusive. This conclusion is counter to the current trend in algal taxonomy in which taxa with minor morphological differences have been repeatedly placed in separate genera based primarily upon DNA sequence evidence.     

A genomic evaluation of taxonomic trends through time in coast horned lizards (genus Phrynosoma)

Determining the boundaries between species and deciding when to describe new species are challenging practices that are particularly difficult in groups with high levels of geographic variation. The coast horned lizards (Phrynosoma blainvillii, Phrynosoma cerroense and P. coronatum) have an extensive geographic distribution spanning many distinctive ecological regions ranging from northern California to the Cape Region of Baja California, Mexico, and populations differ substantially with respect to external morphology across much of this range. The number of taxa recognized in the group has been reevaluated by herpetologists over 20 times during the last 180 years, and typically without the aid of explicit species delimitation methods, resulting in a turbulent taxonomy containing anywhere from one to seven taxa. In this study, we evaluate taxonomic trends through time by ranking 15 of these species delimitation models (SDMs) using coalescent analyses of nuclear loci and SNPs in a Bayesian model comparison framework. Species delimitation models containing more species were generally favoured by Bayesian model selection; however, several three‐species models outperformed some four‐ and five‐species SDMs, and the top‐ranked model, which contained five species, outperformed all SDMs containing six species. Model performance peaked in the 1950s based on marginal likelihoods estimated from nuclear loci and SNPs. Not surprisingly, SDMs based on genetic data outperformed morphological taxonomies when using genetic data alone to evaluate models. The de novo estimation of population structure favours a three‐population model that matches the currently recognized integrative taxonomy containing three species. We discuss why Bayesian model selection might favour models containing more species, and why recognizing more than three species might be warranted.     

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.     

Phylogenetic revision of the Hippasterinae (Goniasteridae; Asteroidea): systematics of deep sea corallivores,including one new genus and three new species

The Hippasterinae is a subfamily within the Goniasteridae, consisting of five genera and 26 species, which occur in cold‐water settings ranging from subtidal to abyssal depths. All known genera were included in a cladistic analysis resulting in two most parsimonious trees, supporting the Hippasterinae as monophyletic. Our review supports Sthenaster emmae gen. et sp. nov. as a new genus and species from the tropical Atlantic and two new Evoplosoma species, Evoplosoma claguei sp. nov. and Evoplosoma voratus sp. nov. from seamounts in the North Pacific. Hippasteria caribaea is reassigned to the genus Gilbertaster, which previously contained a single Pacific species. Our analysis supports Evoplosoma as a derived deep water lineage relative to its continental‐shelf, shallow water sister taxa. The genus Hippasteria contains approximately 15 widely distributed, but similar‐looking species, which occur in the northern and southern hemispheres. Except for Gilbertaster, at least one species in each genus has been observed or is inferred to prey on deep‐sea corals, suggesting that this lineage is important to the conservation of deep‐sea coral habitats. The Hippasterinae shares several morphological similarities with Circeaster and Calliaster, suggesting that they may be related. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 266–301.