Sampling at appropriate spatial scales in the Southern Ocean is logistically challenging and may influence estimates of diversity by missing intermediate representatives. With the assistance of sampling efforts especially influenced by the International Polar Year 2007-2008, we gathered nearly 1500 specimens of the crinoid species Promachocrinus kerguelensis from around Antarctica. We used phylogeographic and phylogenetic tools to assess its genetic diversity, demographic history and evolutionary relationships. Six phylogroups (A-F) identified in an earlier study are corroborated here, with the addition of one new phylogroup (E2). All phylogroups are circumpolar, sympatric and eurybathic. The phylogeny of Promachocrinus phylogroups reveals two principal clades that may represent two different cryptic species with contrasting demographic histories. Genetic diversity indices vary dramatically within phylogroups, and within populations, suggesting multiple glacial refugia in the Southern Ocean: on the Kerguelen Plateau, in the East Weddell Sea and the South Shetland Islands (Atlantic sector), and on the East Antarctic continental shelf in the Dumont d'Urville Sea and Ross Sea. The inferences of gene flow vary among the phylogroups, showing discordant spatial patterns. Phylogroup A is the only one found in the Sub-Antarctic region, although without evident connectivity between Bouvet and Kerguelen populations. The Scotia Arc region shows high levels of connectivity between populations in most of the phylogroups, and barriers to gene flow are evident in East Antarctica.
There has been a significant body of literature on species flock definition but not so much about practical means to appraise them. We here apply the five criteria of Eastman and McCune for detecting species flocks in four taxonomic components of the benthic fauna of the Antarctic shelf: teleost fishes, crinoids (feather stars), echinoids (sea urchins) and crustacean arthropods. Practical limitations led us to prioritize the three historical criteria (endemicity, monophyly, species richness) over the two ecological ones (ecological diversity and habitat dominance). We propose a new protocol which includes an iterative fine-tuning of the monophyly and endemicity criteria in order to discover unsuspected flocks. As a result nine « full » species flocks (fulfilling the five criteria) are briefly described. Eight other flocks fit the three historical criteria but need to be further investigated from the ecological point of view (here called « core flocks »). The approach also shows that some candidate taxonomic components are no species flocks at all. The present study contradicts the paradigm that marine species flocks are rare. The hypothesis according to which the Antarctic shelf acts as a species flocks generator is supported, and the approach indicates paths for further ecological studies and may serve as a starting point to investigate the processes leading to flock-like patterning of biodiversity.
A revision of the stalked crinoid species attributed to the genus Endoxocrinus A.H. Clark, 1908 (Diplocrininae, Pentacrinitidae, Crinoidea, Echinodermata) is conducted using studies on phenotype variation and its relation with environment. Specimens collected via submersible at five sites in the Bahamas exhibit distinct phenotypes that correlate with different apparent ecological niches and serve as references for interpreting specimens dredged in Atlantic and Pacific Oceans where detailed information on their benthic environment is unknown. Documentation of ecophenotypic convergences or divergences allows us to distinguish between adaptive characters and those revealing genetic affinities, and to discuss allopatric evolution and bathymetric zonation. The results suggest the following taxonomy: the genus Endoxocrinus is subdivided into two subgenera, i.e., Endoxocrinus A.H. Clark, 1908 and Diplocrinus Döderlein, 1912 (Annacrinus A. H. Clark, 1923 becomes a junior synonym of Diplocrinus); the subgenus Endoxocrinus is monospecific with E. (E.) parrae [Gervais (in Guérin, 1835)] from the western tropical Atlantic; the subgenus Diplocrinus includes E. (D.) alternicirrus (Carpenter, 1882) from the western and central Pacific, E. (D.) maclearanus (Thomson, 1872) from the western tropical Atlantic, and E. (D.) wyvillethomsoni (Jeffreys, 1870) from the northeastern Atlantic. Endoxocrinus (E.) parrae includes three subspecies adapted to different habitats and depths: E. (E.) parrae parrae usually in 154–518 m with moderate to high current velocity and moderate turbulence to laminar flow, E. (E.) parrae carolinae (A.H. Clark, 1934) in 504–724 m with moderate current velocity and high turbulence, and E. (E.) parrae prionodes H.L. Clark, 1941 in 402–832 m with high current velocity in laminar flow. E. (D.) alternicirrus includes two subspecies, E. (D.) alternicirrus alternicirrus in 625–1476 m and E. (D.) alternicirrus sibogae (Döderlein, 1907) usually in 364–800 m. E. (D.) maclearanus has a depth range of 432–878 m and occurs as a dwarf variety minimus n. var. in high current velocities and high turbulence. E. (D.) wyvillethomsoni from depths of 1214–2070 m lives on various substrates under a variety of hydrodynamic conditions.
We describe the first molecular and morphological analysis of extant crinoid highlevel inter-relationships. Nuclear and mitochondrial gene sequences and a cladistically coded matrix of 30 morphological characters are presented, and analysed by phylogenetic methods.The molecular data were compiled from concatenated nuclear-encoded 18S rDNA, internal transcribed spacer 1, 5.8S rDNA, and internal transcribed spacer 2, together with part of mitochondrial 16S rDNA, and comprised 3593 sites, of which 313 were parsimonyinformative. The molecular and morphological analyses include data from the bourgueticrinid, Bathycrinus; the antedonid comatulids, Dorometra and Florometra; the cyrtocrinids Cyathidium, Gymnocrinus, and Holopus; the isocrinids Endoxocrinus, and two species of Metacrinus; as well as from Guillecrinus and Caledonicrinus, whose ordinal relationships are uncertain, together with morphological data from Proisocrinus. Because the molecular data include indel-rich regions, special attention was given to alignment procedure, and it was found that relatively low, gene-specific, gap penalties gave alignments from which congruent phylogenetic information was obtained from both well-aligned, indel-poor and potentially misaligned, indel-rich regions. The different sequence data partitions also gave essentially congruent results. The overall direction of evolution in the gene trees remains uncertain: an asteroid outgroup places the root on the branch adjacent to the slowly-evolving isocrinids (consistent with palaeontological order of first appearances), but maximum likelihood analysis with a molecular clock places it elsewhere. Despite lineage-specific rate differences, the clock Only an unusual, divergent 18S rDNA sequence was obtained from the morphologically strange cyrtocrinid Cyathidium. Although not analysed in detail, features of this sequence suggested that it may be a PCR artefact, so that the apparenly basal position of this taxon requires confirmation. If not an artefact, Cyathidium either diverged from the crinoid stem much earlier than has been reognized hitherto (i.e., it may be a Palaeozoic relic), or it has an atypically high rate of molecular evolution.
The genus Conocrinus d’Orbigny, 1850 (Crinoidea, Bourgueticrinina) was established on the basis of two aboral cups that had previously been described as Bourgueticrinus thorenti d’Archiac, 1846. One of these (now considered lost) came from the “Rocher du Goulet” at the base of the Biarritz section (Bartonian, Côte des Basques, southwest France). D’Archiac figured only the second cup; this belongs to the d’Orbigny Collection and is still housed in the palaeontological collection of the Muséum national d’Histoire naturelle (Paris) as the lectotype of the species, C. thorenti. It appears that it was collected from Priabonian levels exposed near Castellane (Alpes de Haute Provence, southeast France). New observations on this cup, as well as a detailed study of the characters of aboral cups, columnals and proximal brachials in a few extant and fossil species classically attributed to Conocrinus or to closely related genera such as Democrinus, Rhizocrinus and Tormocrinus, have yielded arguments for a revision of the taxonomy and interrelationships of extant and fossil taxa in the family Bourgueticrinidae. Conocrinus (= Tormocrinus), as here interpreted, includes six Eocene species: C. thorenti, C. archiaci, C. cahuzaci n. sp., C. duperrieri, C. cf. suessi and C. veronensis. Numerous extinct species previously attributed to Conocrinus or Democrinus are here transferred to two new genera which first occur in the lower Paleocene: Paraconocrinus n. gen. (type species: P. pyriformis) and Pseudoconocrinus n. gen. (type species: P. doncieuxi). Aboral cups from the “Rocher du Goulet” (Biarritz) are here assigned to Paraconocrinus pellati n. gen., n. sp., while the Danian species Democrinus maximus is transferred to Pseudoconocrinus n. gen. A new genus, Cherbonniericrinus, is created to accommodate a single extant species, Ch. cherbonnieri, previously attributed to Conocrinus, while the extant genus Rhizocrinus, closely related to Democrinus, is resurrected. Conocrinus and closely related genera are derived from a bourgueticrinine lineage the first record of which is from the lower Campanian, with the new genus Carstenicrinus. These are all attributed to the family Rhizocrinidae which is here considered distinct from the family Bourgueticrinidae. Rhizocrinids rapidly diversified immediately after the Cretaceous-Paleogene (K/Pg) event. Cretaceous taxa previously placed within the family Bourgueticrinidae now appear to be polyphyletic. Some of them do not belong to Bourgueticrinina, such as those of the Dunnicrinus lineage. Interrelationships of Rhizocrinidae and other post-Palaeozoic families having a xenomorphic stalk are discussed.
A large collection of ascidians was made during the CEAMARC Aurora Australis V3 cruise off Terre Adélie and George V Land a region rarely investigated before at these depths. Sampling was performed by beam trawls and a dredge between 138°– 146° latitude East and from 150 to 1700 m depth, on the Antarctic shelf and slope. Three of the 33 ascidian species identified are new and belong to the Stolidobranchia. Half of the species have an exclusive Antarctic distribution, others also occur in Sub-Antarctic areas, but none are common with the southern temperate fauna. The CEAMARC collection does not contain the whole range of already known species from this region. Moreover, brittle and very small specimens were not collected. COI sequences were obtained for 37 specimens, including two of the new species.
Zimmermann, G., Bosc, P., Valade, P., Cornette, R., Améziane, N. and Debat, V. 2012. Geometric morphometrics of carapace of Macrobrachium australe (Crustacea: Palaemonidae) from Reunion Island. -Acta Zoologica (Stockholm) 93: 492-500.We investigated the structure of carapace shape variation in six populations of Macrobrachium australe Guérin-Méneville 1838 (Crustacea: Decapoda: Palaemonidae) from Reunion Island (Indian Ocean) freshwaters. The morphometric analysis revealed the occurrence of two morphotypes corresponding to two different types of habitats. Individuals living in lotic habitats present a thick carapace armed with a short, robust and straight rostrum, while individuals from lentic habitats have a slender carapace armed with a thin long rostrum orientated upward. This difference suggests an adaptation to lotic disturbances and is tentatively interpreted as adaptive phenotypic plasticity. In such amphidromous organisms regressing to freshwaters after a marine larval phase, selection for physiological and developmental flexibility might facilitate further adaptation and allows the colonisation of a wide panel of environmentally different and sometimes geographically distant insular streams.
Species flocks (SFs) fascinate evolutionary biologists who wonder whether such striking diversification can be driven by normal evolutionary processes. Multiple definitions of SFs have hindered the study of their origins. Previous studies identified a monophyletic taxon as a SF if it displays high speciosity in an area in which it is endemic (criterion 1), high ecological diversity among species (criterion 2), and if it dominates the habitat in terms of biomass (criterion 3); we used these criteria in our analyses. Our starting hypothesis is that normal evolutionary processes may provide a sufficient explanation for most SFs. We thus clearly separate each criterion and identify which biological (intrinsic) and environmental (extrinsic) traits are most favourable to their realization.The first part focuses on evolutionary processes. We highlight that some popular putative causes of SFs, such as key innovations or ecological speciation, are neither necessary nor sufficient to fulfill some or all of the three criteria. Initial differentiation mechanisms are diverse and difficult to identify a posteriori because a primary differentiation of one type (genetic, ecological or geographical) often promotes other types of differentiation. Furthermore, the criteria are not independent: positive feedbacks between speciosity and ecological diversity among species are expected whatever the initial cause of differentiation, and ecological diversity should enhance habitat dominance at the clade level. We then identify intrinsic and extrinsic factors that favour each criterion. Low dispersal emerges as a convincing driver of speciosity. Except for a genomic architecture favouring ecological speciation, for which assessment is difficult, high effective population sizes are the single intrinsic factor that directly enhances speciosity, ecological diversity and habitat dominance. No extrinsic factor appeared to enhance all criteria simultaneously but a combination of factors (insularity, fragmentation and environmental stability) may favour the three criteria, although the effect is indirect for habitat dominance.We then apply this analytical framework to Antarctic marine environments by analysing data from 18 speciose clades belonging to echinoderms (five unrelated clades), notothenioid fishes (five clades) and peracarid crustaceans (eight clades). Antarctic shelf environments and history appear favourable to endemicity and speciosity, but not to ecological specialization. Two main patterns are distinguished among taxa. (i) In echinoderms, many brooding, species-rich and
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