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.     

Historical biogeography of Iberobathynella (Crustacea, Syncarida, Bathynellacea), an aquatic subterranean genus of Parabathynellids, endemic to the Iberian Peninsula

• 1 The origin and distribution of the species of freshwater interstitial (stygobiont) syncarids of the Iberobathynella genus (Crustacea, Parabathynellidae), endemic to the Iberian Peninsula, are considered in the context of evolutionary biogeography.
• 2 We established the most likely phylogenetic relationships between the 14 known species of the Iberobathynella genus on the basis of morphological characters. We assumed a common marine ancestor and estimated the possible phylogenetic ages of the 14 inland species from the number of cladogram nodes.
• 3 The land ages at the sites where the species are found help to tell us when the landmasses emerged definitively: most species are found in areas that have undergone one or more marine transgressions/regressions since the Triassic and which could therefore have been colonized by the common ancestors.
• 4 We assessed the Boutin–Coineau two‐step model of colonization and evolution in an attempt to elucidate the evolutionary history of Iberobathynella and to offer a plausible palaeogeographical scenario. We looked for concordance between the possible chronology of Iberobathynella taxa colonization and their phylogenetic relationships, but found none of significance. Hence, we cannot reject the possibility that evolutionary clade divergence within the genus was independent of sea‐level changes in the Senonian (88.5–65 mya) and Eocene (56.5–35.4 mya) epochs. We can therefore neither confirm nor rule out a correlation.

     

Repeated vicariance of Eurasian songbird lineages since the Late Miocene

Aim The evolution of avian speciation patterns across much of Eurasia is under‐explored. Excepting phylogeographic patterns of single species, or speciation involving the Himalayas, there has been no attempt to understand the evolution of avian distributional patterns across the rest of the continent. Within many genera there is a pattern of (presumed) sister species occurring in adjacent areas (western, eastern or southern Eurasia), yet this pattern cannot be explained by existing biogeographic barriers. My aim was to examine the possible role of climate‐driven vicariance events in generating avian distributions. Location Eurasia. Methods I constructed a molecular phylogeny of Phoenicurus redstarts, and assembled phylogenetic data from published studies of seven other Eurasian bird genera. On each phylogeny, I assessed the distributional patterns of species and clades relative to refugial areas in western, eastern and southern Eurasia. I also estimated the timing of lineage divergences via a molecular clock, to determine whether distributional patterns can be explained by well‐defined periods of climate change in Eurasia that are recorded from dated sediments in the Chinese Loess Plateau. Results Species relationships in a well‐supported phylogeny of Phoenicurus show a pattern of distributions consistent with repeated speciation in major refugial areas, where one lineage is isolated in a single area of Eurasia relative to its sister lineage. This same pattern is evident in Eurasian Turdus thrushes, and six additional avian genera distributed across Eurasia. Molecular clock dating indicates that divergences within each genus are the result of multiple rounds of speciation in refugia through time, during major climate‐driven episodes of vicariance. Main conclusions Analyses revealed substantial evidence supporting a repeated, non‐random pattern of speciation within and across eight songbird lineages since the Late Miocene. The pattern of speciation supports a model of isolation in refugia during major episodes of vicariance, specifically periods of either intensified desertification of Central Asia or Eurasian glacial cycles. The densely sampled clades used here preclude inter‐continental dispersal as an alternative explanation for distributions. The signature of climate‐driven vicariance across epochs is, given the absence of extant biogeographic barriers, a suitable hypothesis to explain major lineage divergences in widely distributed Eurasian songbird lineages.     

Evolutionary history, biogeography and eco-climatological differentiation of the genus Anthemis L. (Compositae, Anthemideae) in the circum-Mediterranean area

Aim To reconstruct the temporal, geographical and eco‐climatological differentiation of the genus Anthemis (Compositae, Anthemideae) in the circum‐Mediterranean region, in order to evaluate the relative importance of geographical vs. climatological differentiation processes in influencing the actual distribution patterns in this plant group. Location The circum‐Mediterranean region, including the Iberian Peninsula, northern Africa, the Italian and Balkan peninsulas, the Aegean region and Anatolia, the Caucasus, the Arabian Peninsula and western Asia. Methods The phylogeny of the genus Anthemis was obtained from a maximum likelihood analysis based on nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) sequence data, and the chronology of diversification was derived using a penalized likelihood approach. The reconstruction of the spatial diversification of the genus was based on a dispersal/vicariance (DIVA) analysis. Eco‐climatological niche differentiation was inferred by optimizing 19 bioclimatic variables onto the phylogeny. A multi‐dimensional hypervolume, proposed as a representation of the eco‐climatological niche and defined by the combination of ranges for all bioclimatic variables, was calculated for each taxon and each internal node. To identify ‘eco‐climatological vicariance’ events in the phylogeny, the pairwise overlap among hypervolumes of sister groups was calculated. Finally, the temporal and clade‐wise relative importance of geographical vs. eco‐climatological vicariance events was estimated. Results The temporal reconstruction shows a constant increase of lineages through the last 12 Myr. The geographical reconstruction suggests that Anthemis diverged from the rest of the Compositae–Anthemideae in the eastern Mediterranean region, and from there radiated into the whole circum‐Mediterranean region through successive dispersal and vicariance events. The reconstruction of the eco‐climatological niches suggests a progressive adaptation from a montane‐humid climate towards arid environments and the typical mediterranean climate. Main conclusions The results presented here involved phylogenetic, geographical and eco‐climatological reconstructions; joint analyses of all of these aspects have assessed the relative importance of geological vs. climatic forces that have affected the distributional history of the genus Anthemis. Large‐scale differentiation patterns triggered by geological forces appear to have influenced the evolutionary history of the genus in a rather constant manner over the last 12 Myr, whereas climatic forces seem to have played an important role in two phases of the radiation process: at around 9 Ma, when the area experienced the onset of a trend towards aridification, and during the last 3.5 Myr, with the establishment of the typical mediterranean climate and the influence of Pleistocene climate oscillations.     

TERTIARY SPECIATION MODELS IN AUSTRALIAN ANURANS: MOLECULAR DATA CHALLENGE PLEISTOCENE SCENARIO

Similarities between frogs in the faunas of southwestern and southeastern Australia have long been viewed as indicators of close genetic relationships and recent (Pleistocene) divergences. We studied albumin evolution in 16 east-west species pairs of frogs representing six genera to assess the validity of these conclusions. Analysis of albumin evolution in western species of Heleioporus and some species of Litoria suggested recent speciation in these genera, with the closest sister groups occurring in the western and not among the eastern fauna. All divergences measured between eastern and western cognate species point to a Tertiary separation extending from the late Miocene to the early Oligocene. Micro-complement fixation studies provide an independent estimation of both genetic relationships between species pairs and the time of divergence of each species pair, allowing the testing of models of speciation and vicariance biogeography in a way not possible with earlier methodologies.     

Cyclamen: time, sea and speciation biogeography using a temporally calibrated phylogeny

Aim The Mediterranean region is a species‐rich area with a complex geographical history. Geographical barriers have been removed and restored due to sea level changes and local climatic change. Such barriers have been proposed as a plausible mechanism driving the high levels of speciation and endemism in the Mediterranean basin. This raises the fundamental question: is allopatric isolation the mechanism by which speciation occurs? This study explores the potential driving influence of palaeo‐geographical events on the speciation of Cyclamen (Myrsinaceae), a group with most species endemic to the Mediterranean region. Cyclamen species have been shown experimentally to have few genetic barriers to hybridization. Location The Mediterranean region, including northern Africa, extending eastwards to the Black Sea coast. Methods A generic level molecular phylogeny of Myrsinaceae and Primulaceae is constructed, using Bayesian approximation, to produce a secondary age estimate for the stem lineage of Cyclamen. This estimate is used to calibrate temporally an infrageneric phylogeny of Cyclamen, built with nrDNA ITS, cpDNA trnL‐F and cpDNA rps16 sequences. A biogeographical analysis of Cyclamen is performed using dispersal–vicariance analysis. Results The emergence of the Cyclamen stem lineage is estimated at 30.1–29.2 Ma, and the crown divergence at 12.9–12.2 Ma. The average age of Cyclamen species is 3.7 Myr. Every pair of sister species have mutually exclusive, allopatric distributions relative to each other. This pattern appears typical of divergence events throughout the evolutionary history of the genus. Main conclusions Geographical barriers, such as the varying levels of the Mediterranean Sea, are the most plausible explanation for speciation events throughout the phylogenetic history of Cyclamen. The genus demonstrates distributional patterns congruent with the temporally reticulate palaeogeography of the Mediterranean region.     

Ancient vicariance and climate‐driven extinction explain continental‐wide disjunctions in Africa: the case of the Rand Flora genus Canarina (Campanulaceae)

Transoceanic distributions have attracted the interest of scientists for centuries. Less attention has been paid to the evolutionary origins of ‘continent‐wide’ disjunctions, in which related taxa are distributed across isolated regions within the same continent. A prime example is the ‘Rand Flora’ pattern, which shows sister taxa disjunctly distributed in the continental margins of Africa. Here, we explore the evolutionary origins of this pattern using the genus Canarina, with three species: C. canariensis, associated with the Canarian laurisilva, and C. eminii and C. abyssinica, endemic to the Afromontane region in East Africa, as case study. We infer phylogenetic relationships, divergence times and the history of migration events within Canarina using Bayesian inference on a large sample of chloroplast and nuclear sequences. Ecological niche modelling was employed to infer the climatic niche of Canarina through time. Dating was performed with a novel nested approach to solve the problem of using deep time calibration points within a molecular dataset comprising both above‐species and population‐level sampling. Results show C. abyssinica as sister to a clade formed by disjunct C. eminii and C. canariensis. Miocene divergences were inferred among species, whereas infraspecific divergences fell within the Pleistocene–Holocene periods. Although C. eminii and C. canariensis showed a strong genetic geographic structure, among‐population divergences were older in the former than in the latter. Our results suggest that Canarina originated in East Africa and later migrated across North Africa, with vicariance and aridification‐driven extinction explaining the 7000 km/7 million year divergence between the Canarian and East African endemics.     

Biogeography and evolution of Abies (Pinaceae) in the Mediterranean Basin: the roles of long‐term climatic change and glacial refugia

Aim The genus Abies exemplifies plant diversification related to long‐term climatic, geological and evolutionary changes. Today, the Mediterranean firs comprise nine species, one natural hybrid and several varieties. Here I summarize current knowledge concerning the origin and evolution of the genus Abies in the Mediterranean Basin and propose a comprehensive hypothesis to explain the isolation and speciation pattern of Mediterranean firs. Location The Mediterranean Basin. Methods The literature on Abies was reviewed, focusing on the morphology, fossil records, molecular ecology, phytosociology and biogeography of the genus in the Mediterranean Basin. Results Abies fossils from the western Mediterranean indicate a wide Tertiary circum‐Mediterranean distribution of the Abies ancestor. Palaeogeographical data also suggest a single eastern Mediterranean Tertiary ancestor. Following the Miocene to Pliocene climate crisis and marine transgressions, the ancestor of the northern Mediterranean firs is hypothesized to have separated into two eastern groups, one on the Balkan Peninsula and the other in Asia Minor. However, land bridges may have permitted gene flow at times. A southward migration of A. alba to refugia, where older fir species may have remained isolated since the Miocene, could explain recent findings indicating that morphologically distant species are more closely related than expected based on such morphological classification. Main conclusions The Abies genus appears to have undergone significant morphological differentiation that does not necessarily imply reproductive isolation. That is, long‐term Mediterranean Basin dryness along a south‐eastern to north‐western gradient may have caused an initial Miocene–Pliocene speciation sequence. Pleistocene glacial cycles probably forced migrations to occur, leading to repeated contact between fir species in glacial refugia.     

Phylogenetic relationships within the Archiloa genus complex (Proseriata,Monocelididae)

Of the seven genera which we have recognised within the Archiloa genus complex sensu Karling (1966) the cosmopolitan genus Archilina is the most primitive and is characterised only by plesiomorphic characters, and has to be considered paraphyletic. All other species of the Archiloa genus complex are hypothesized to be derived from Archilina-like ancestors through different evolutionary lineages. One lineage led to the genera Archiloa, Inaloa, Archilopsis and Monocelopsis, taxa found in the Atlantic and the Mediterranean. These genera are monophyletic and their relationships are analyzed. The genera Mesoda (Brazil) and Tajikina (Northern Pacific) can be considered as two other separate lineages. Similarly, within what we now consider as the genus Archilina different lineages can be recognized in different regions.     

Phylogenetic analysis of the north‐east Atlantic and Mediterranean species of the genus Stenosoma (Isopoda,Valvifera, Idoteidae)

Xavier, R., Santos, A. M., Harris, D. J., Sezgin, M., Machado, M., Branco, M. (2012). Phylogenetic analysis of the north‐east Atlantic and Mediterranean species of the genus Stenosoma (Isopoda, Valvifera, Idoteidae). —Zoologica Scripta, 41, 386–399. The marine isopod genus Stenosoma is widespread in the northern hemisphere. However, 12 of its 14 known species are found within the Mediterranean basin and adjacent regions of the north‐east Atlantic and the Black Sea. Such a high level of diversity confined to a limited region of a much larger circumglobal distribution suggests that the Mediterranean region may have played a crucial role in the evolutionary history of this genus. In the present work, the phylogeny of the genus Stenosoma was investigated on the basis of DNA sequencing data from one nuclear (28SrRNA) and two mitochondrial (COI, ND4) gene fragments obtained for nine of 12 Atlantic–Mediterranean species. Divergence time estimates point to a Tethyan origin of Stenosoma and suggest that the speciation events from which stem most of the extant species took place well before the Messinian Salinity Crisis. Stenosoma spinosum and Stenosoma appendiculatum are the only exceptions, as they apparently arose within the Mediterranean during the Pleistocene. Phylogenetic reconstruction agrees with current taxonomic status of most species. However, Stenosoma capito clustered in two distinct and well‐supported clades, one composed of eastern Mediterranean and Black Sea specimens and the other by western Mediterranean and Atlantic ones. Such polyphyly suggests the existence of a previously unrecognized species, Stenosoma sp., which so far has been confounded with S. capito.     

Evidence for reticulate palaeogeography: beetle diversity linked to connection-disjunction cycles of the Gibraltar strait

Aim Dispersal barriers between areas within some regions have appeared and disappeared throughout evolutionary time. Here we describe the distributional patterns displayed by three taxa living in such kind of regions. These patterns can be better explained considering a reticulated rather than a hierarchically branched palaeogeography. Location Western Mediterranean. Methods The taxa studied are Misolampus (Coleoptera, Tenebrionidae), Tentyria (Coleoptera, Tenebrionidae) and Thorectes (Coleoptera, Geotrupidae). All them are flightless and show a high degree of endemicity. The individual pattern of area relationships was determined separately for each genus by Brooks Parsimony Analysis (BPA). A theoretical general area cladogram was constructed based on the palaeogeographical history of the region. Finally, the general area cladogram is reconciled with the individual ones. Results The ancestor of Misolampus probably was North African. Land dispersal toward the Iberian Peninsula is proposed. Speciation within Iberia is related to specific vicariance events, and the presence of insular (Balearic Islands) populations is explained by sea‐surface or, more probably, human‐mediated dispersal. The ancestor of Tentyria was Iberic. The proposed hypothesis to explain the current species distribution mainly relies on the occurrence of specific vicariance events. However, the occurrence of some sea‐surface dispersal event is not discarded. Almost all possible vicariance events can be recognized in the first clade of the Thorectes genus. There is evidence for dispersal between Africa and Europe at different dates and in both directions. In spite of some uncertainties, the appearance of the second Thorectes clade can also be explained by the occurrence of specific historical events. An ancient dispersal toward the eastern Mediterranean and several dispersal events during the Messinian seem likely. Main conclusions The same historical events have specific outcomes in every tree (even in every branch within a tree) depending on the ability for dispersal and speciation of each taxon. Connection‐disjunction cycles of dispersal barriers have acted as diversity producers.     

Decapod Crustacea from the Agrio Formation (Lower Cretaceous) of the Neuquén Basin,Argentina

Abstract: Two species of decapod crustacean are recorded from the Agua de la Mula Member of the Agrio Formation (Upper Hauterivian – Lower Barremian) of the Neuquén Basin of west‐central Argentina, namely Astacodes falcifer Bell and a new species of Palaeohomarus, P. pacificus. The preservation of the specimens is exceptional, some showing delicate compound eyes and a stridulatory apparatus, features rarely found in fossil forms. Many specimens are preserved articulated inside calcareous nodules, within dark‐grey shales. The lobster‐bearing sediments accumulated in a low‐energy marine environment and diagenetic mineralization occurred very rapidly, prior to significant decay, thus allowing exceptional preservation of specimens. Palaeohomarus was a rare genus in the Cretaceous with a palaeogeographic distribution restricted to the Mediterranean Tethys, the eastern USA and Madagascar, while Astacodes falcifer has been recorded only from Speeton (eastern England) and Neuquén.     

Deep-sea squat lobster biogeography (Munidopsidae: Leiogalathea) unveils Tethyan vicariance and evolutionary patterns shared by shallow-water relatives

The ecology, abundance and diversity of galatheoid squat lobsters make them an ideal group to study deep-sea diversification processes. Here, we reconstructed the evolutionary and biogeographic history of Leiogalathea, a genus of circum-tropical deep-sea squat lobsters, in order to compare patterns and processes that have affected shallow-water and deep-sea squat lobster species. We first built a multilocus phylogeny and a calibrated species tree with a relaxed clock using StarBEAST2 to reconstruct evolutionary relationships and divergence times among Leiogalathea species. We used BioGeoBEARS and a DEC model, implemented in RevBayes, to reconstruct ancestral distribution ranges and the biogeographic history of the genus. Our results showed that Leiogalathea is monophyletic and comprises four main lineages; morphological homogeneity is common within and between clades, except in one; the reconstructed ancestral range of the genus is in the Atlantic and Indian oceans (Tethys). They also revealed the divergence of the Atlantic species around 25 million years ago (Ma), intense cladogenesis 15–25 Ma and low levels of speciation over the last 5 million years (Myr). The four Leiogalathea lineages showed similar patterns of speciation: allopatric speciation followed by range expansion and subsequent stasis. Leiogalathea started diversifying during the Oligocene, likely in the Tethyan. The Atlantic lineage then split from its Indo-Pacific sister group due to vicariance driven by closure of the Tethys Seaway. The Atlantic lineage is less speciose compared with the Indo-Pacific lineages, with the Tropical Southwestern Pacific being the current centre of diversity. Leiogalathea diversification coincided with cladogenetic peaks in shallow-water genera, indicating that historical biogeographic events similarly shaped the diversification and distribution of both deep-sea and shallow-water squat lobsters.     

Conchospore production and seasonal occurrence of some Porphyra species (Bangiales,Rhodophyta) in Washington State

The leafy thalli of species of the marine red algal genus Porphyra grow rapidly but persist for a relatively short time on rocky intertidal or subtidal substrata or as epiphytes on other marine plants. In most species, the large, short-lived leafy thalli alternate with small, presumably perennial, filamentous conchocelis plants. Depending on the species of northeastern Pacific Porphyra, photoperiod and temperature are important regulators of conchospore formation and release. Data from laboratory studies of conchospore formation and release in five Washington species of Porphyra (P. abottae, P. nereocystis, P. perforata, P. pseudolanceolata and P. torta) indicate that conchospores are most likely to be released at a time that precedes the appearance of the leafy thalli in the field.     

Life history strategies and habitat templets of tropical butterflies in north-eastern Australia

Three multivoltine species of satyrine butterflies in the genus Mycalesis (Lepidoptera: Nymphalidae) are narrowly sympatric in the wet–dry tropics of north-eastern Australia. They show a range of ecological strategies and adaptations associated with contrasting habitats and varying selective pressures. Two abiotic factors, namely favorability (the reciprocal of seasonal adversity) and predictability (broadly the reciprocal of disturbance), were examined as potential environmental selective forces in shaping their life histories. Comparison of several key life history traits of the wet-season form revealed that the life histories of each species corresponded well with their habitat characteristics. M. perseus, which lives in habitats which are less favorable (i.e. adverse) and more unpredictable (i.e. temporary), shows many traits of an r-type strategy: smaller size, faster development, earlier maturation, higher fecundity, smaller egg size, and rapid population increase. By contrast, M. sirius and M. terminus, which live in more favorable and predictable (i.e. permanent) habitats, have many life history attributes and other characteristics in common which link them closer to K-type strategies. The only discrepancy is lower potential reproductive effort of M. perseus, which may be accounted for in terms of an evolutionary trade-off, such as with dispersal or dormancy. Other correlates associated with the M. perseus life history tactic include higher sex-size dimorphism, greater dispersal ability, better tolerance to adverse conditions, stronger phenotypic variation, greater degree of polyandry, and a more flexible breeding strategy. The life history patterns of these species are discussed in the context of evolutionary life history models, particularly the Southwood–Greenslade habitat templet.     

Pillars of Hercules: is the Atlantic–Mediterranean transition a phylogeographical break?

The geological history of the Mediterranean Sea, its hydrography and connection with the Atlantic Ocean have been well documented. Despite a wealth of historical and oceanographic data, the Atlantic-Mediterranean transition remains controversial at the biological level as there are discordant results regarding the biogeographical separation between the Atlantic and Mediterranean biota. The opening of the Strait of Gibraltar at the end of the Messinian Salinity Crisis (some 5.33 million years ago), removed the land barrier that impeded the marine biota allowing it to disperse freely into the Mediterranean Sea. However, present day genetic patterns suggest a limitation to gene flow for some marine species, preventing population admixture. In the last few years, a large number of studies have challenged the hypothesis of the Strait of Gibraltar representing a phylogeographical break. A review of more than 70 papers reveals no obvious relationship between either dispersal ability or life history, and observed patterns of partial or complete genetic isolation between Atlantic and Mediterranean populations. We re-analysed a selection of this large body of data (20 studies in total) in order to provide a homogeneous and coherent view on the generality of the phylogeographical patterns and the presence of a phylogeographical barrier. This offered the opportunity to summarize the state of the art on this matter and reach some general conclusions on the evolutionary history across the Atlantic-Mediterranean range. Geographically, some species in the transition zone showed step changes of allele frequencies associated with the Almeria-Oran Front rather than with the Strait of Gibraltar itself. A major part of the data describe evolutionary events well within the time frame of the Quaternary age as very few taxa pre-date closure of the Tethys Sea. Results point to a combined signature of vicariance, palaeoclimate fluctuation and life-history traits on the Atlantic-Mediterranean phylogeographical patterns. Principal component analysis failed to show any particular association between biological traits and genetic variables. It would argue that organismal determinism may play a far less significant role than marine biogeographers have generally believed.     

Ribosomal gene variation in soybean (Glycine) and its relatives

Summary The genes encoding the 18S25S ribosomal RNA gene repeat in soybean (Glycine max) and its relatives in the genus Glycine are surveyed for variation in repeat length and restriction enzyme site locations. Within the wild species of subgenus Glycine, considerable differences in repeat size occur, with a maximum observed in G. falcata. Repeat length and site polymorphisms occur in several species, but within individual plants only single repeat types are observed. The rDNA of the cultivated soybean and its wild progenitor, G. soja are identical at the level of this study, and no variation is found in over 40 accessions of the two species. Data from rDNA mapping studies are congruent with those of previous biosystematic studies, and in some instances give evidence of divergences not seen with other approaches.     

Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina

A phylogenetic approach to the origin and maintenance of species diversity ideally requires the sampling of all species within a clade, confirmation that they are evolutionarily distinct entities, and knowledge of their geographical distributions. In the marine tropics such studies have mostly been of fish and reef-associated organisms, usually with high dispersal. In contrast, snails of the genus Echinolittorina (Littorinidae) are restricted to rocky shores, have a four-week pelagic development (and recorded dispersal up to 1400 km), and show different evolutionary patterns. We present a complete molecular phylogeny of Echinolittorina, derived from Bayesian analysis of sequences from nuclear 28S rRNA and mitochondrial 12S rRNA and COI genes (nodal support indicated by posterior probabilities, maximum likelihood, and neighbor-joining bootstrap). This consists of 59 evolutionarily significant units (ESUs), including all 50 known taxonomic species. The 26 ESUs found in the Indo-West Pacific region form a single clade, whereas the eastern Pacific and Atlantic species are basal. The earliest fossil occurred in the Tethys during the middle Eocene and we suggest that the Indo-West Pacific clade has been isolated since closure of the Tethyan seaway in the early Miocene. The geographical distributions of all species (based on more than 3700 locality records) appear to be circumscribed by barriers of low temperature, unsuitable sedimentary habitat, stretches of open water exceeding about 1400 km, and differences in oceanographic conditions on the continuum between oceanic and continental. The geographical ranges of sister species show little or no overlap, indicating that the speciation mode is predominantly allopatric. Furthermore, range expansion following speciation appears to have been limited, because a high degree of allopatry is maintained through three to five branching points of the phylogeny. This may be explained by infrequent long-distance colonization, habitat specialization on the oceanic/continental gradient, and perhaps by interspecific competition. In the eastern Pacific plus Atlantic we identify five cases of divergence on either side of the Isthmus of Panama, but our estimates of their ages pre-date the emergence of the Isthmus. There are three examples of sister relationships between species in the western Atlantic and eastern Atlantic, all resulting from dispersal to the east. Within the Indo-West Pacific, we find no geographical pattern of speciation events; narrowly endemic species of recent origin are present in both peripheral and central parts of the region. Evidence from estimated divergence times of sister species, and from a plot of the number of lineages over time, suggest that there has been no acceleration of diversification during the glacio-eustatic cycles of the Plio-Pleistocene. In comparison with reefal organisms, species of Echinolittorina on rocky shores may be less susceptible to extinction or isolation during sea-level fluctuations. The species richness of Echinolittorina in the classical biogeographic provinces conforms to the common pattern of highest diversity (11 species) in the central "East Indies Triangle" of the Indo-West Pacific, with a subsidiary focus in the eastern Pacific and western Atlantic, and lowest diversity in the eastern Atlantic. The diversity focus in the East Indies Triangle is produced by a mosaic of restricted allopatric species and overlap of a few widespread ones, and is the result of habitat specialization rather than historical vicariance. This study emphasizes the plurality of biogeographic histories and speciation patterns in the marine tropics.     

Methanogenium,a new genus of marine methanogenic bacteria,and characterization ofMethanogenium cariaci sp. nov. andMethanogenium marisnigri sp. nov.

A new genus of marine methanogenic bacteria and two species within this genus are described.Methanogenium is the proposed genus andMethanogenium cariaci the type species. Cells of the type species are Gram-negative, peritrichously flagellated, irregular cocci with a periodic wall surface pattern. Colonies formed by these bacteria are yellow, circular and umbonate with entire edges. The DNA base composition is 52 mol% guanine plus cytosine. Formate or hydrogen and carbon dioxide serve as substrates for growth. Cells ofMethanogenium marisnigri are of similar shape but smaller diameter thanM. cariaci. The colonies ofM. marisnigri are convex, and the DNA base composition is 61 mol % G+C. Formate or hydrogen and carbon dioxide are growth substrates. Sodium chloride is required for growth of both methanogens.Abbreviations SDS sodium dodecylsulfate - PIPES piperazine-N,N-bis (2 ethanesulfonic acid) - HS-CoM coenzyme M, 2-mercaptoethanesulfonic acid     

Influence du substrat sur la répartition des Crustacés stygobies interstitiels du Haut-Atlas occidental

The interstitial stygobiont distributions of two rivers in the western High Atlas is analyzed in relation with sediment granulometry. Sixteen stations were sampled along the rivers N'Fis and Zat and their tributaries. Granulometry analyses were performed. Three grain size sediment types constitute the subterranean biotopes and reflect the local hydrodynamical conditions. The main rivers stations, which undergo frequent natural or human disturbances, are characterized by a high content of fine sediment; in contrast, tributaries stations show coarse and well-classified sediments due to more stable biotopes and more regular hydrological conditions. The 92 taxa group 28 stygobiontic species with 11 crustaceans species. PCA. based on 16 stations, three faunistic richness indexes and three granulometric factors display a high correlation between coarse grain-size content and the richness of stygobiontic taxa. A high silt content both limits the interstitial taxa and excludes crustaceans that occur only in coarse gravels. The granulometric preferenda in the species of the amphipod genus Metacrangonyx and isopods Microcerberus. Microcharon and Typhlocirolana are shown on Shepard triangular diagrams. Cohabitation of several species of the same genus could be explained by the occupancy of different ecological niches resulting from the grain-size proportion.