Allopatric and Sympatric Drivers of Speciation in Alviniconcha Hydrothermal Vent Snails Corinna Breusing ,*‡,1 Shannon B

Total Page:16

File Type:pdf, Size:1020Kb

Allopatric and Sympatric Drivers of Speciation in Alviniconcha Hydrothermal Vent Snails Corinna Breusing ,*‡,1 Shannon B Allopatric and Sympatric Drivers of Speciation in Alviniconcha Hydrothermal Vent Snails Corinna Breusing ,*‡,1 Shannon B. Johnson,‡,2 Verena Tunnicliffe,3 David A. Clague,2 Robert C. Vrijenhoek,2 and Roxanne A. Beinart1 1Graduate School of Oceanography, University of Rhode Island, Narragansett, RI 2Monterey Bay Aquarium Research Institute, Moss Landing, CA 3Department of Biology and School of Earth and Ocean Sciences, University of Victoria, Victoria, BC, Canada Downloaded from https://academic.oup.com/mbe/advance-article/doi/10.1093/molbev/msaa177/5870836 by guest on 24 September 2020 †These authors contributed equally to this work. *Corresponding author: E-mail: [email protected]. Associate editor: Corinna Breusing New Sanger sequences for Alviniconcha and Ifremeria hosts and symbionts are available in GenBank under accession numbers MT131487–MT131781 (COI), MT130994–MT131149 (12S), MT146898–MT147385 (H3), MT148092–MT148633 (ATPSa), MT148634–MT149209 (ATPSb), MT147386–MT148091 (EF1a), and MT137388–MT137420 (16S). 16S amplicon sequences have been deposited in the Sequence Read Archive under BioProject numbers PRJNA473256, PRJNA473257, PRJNA610289, and PRJNA610290. Abstract Despite significant advances in our understanding of speciation in the marine environment, the mechanisms un- derlying evolutionary diversification in deep-sea habitats remain poorly investigated. Here, we used multigene molecular clocks and population genetic inferences to examine processes that led to the emergence of the six extant lineages of Alviniconcha snails, a key taxon inhabiting deep-sea hydrothermal vents in the Indo-Pacific Ocean. We show that both allopatric divergence through historical vicariance and ecological isolation due to niche segregation contributed to speciation in this genus. The split between the two major Alviniconcha clades (separating A. boucheti and A. marisindica from A. kojimai, A. hessleri, and A. strummeri) probably resulted from tectonic processes leading to geographic separation, whereas the splits between co-occurring species might have been influenced by ecological factors, such as the availability of specific chemosynthetic symbionts. Phylogenetic origin of the sixth species, Alviniconcha adamantis, remains uncertain, although its sister position to other extant Alviniconcha lineages indicates a possible ancestral relationship. This study lays a foundation for future genomic studies aimed at deciphering the roles of local adaptation, reproductive biology, and host–symbiont compatibility in speciation of these vent-restricted snails. Key words: Alviniconcha, speciation, allopatric divergence, ecological isolation, deep-sea hydrothermal vents, chemo- Article synthetic symbionts. Introduction ecological niches (Rundle and Nosil 2005; Schluter 2009). Habitat isolation can occur at different spatial scales that How novel species evolve in response to environmental range from “macrospatial” (a form of allopatric speciation, pressures remains a fundamental question in evolutionary where gene flow between populations is absent due geo- biology (Schluter 1998). New species can originate through a graphic separation) to “microspatial” (where distinct popu- number of mechanisms, including geographic isolation (al- lations co-occur geographically but rarely interbreed lopatric speciation) and formation of reproductive barriers because of adaptations to different ecological niches) due to localized natural selection (sympatric speciation) (Coyne and Orr 2004). By contrast, mutation-order specia- (Darwin 1859; Mayr 1942; Dobzhansky 1951; Coyne and tion happens when reproductive isolation evolves due to Orr 2004; Schuler et al. 2016; Yang et al. 2017). The evolution random fixation of alleles in two populations that face the of new species commonly occurs as a result of ecological or same selective pressures (Mani and Clarke 1990; Schluter mutation-order speciation (Schluter 2009). Ecological speci- 2009; Nosil and Flaxman 2011). Evolutionary divergence un- ation theory predicts that reproductive isolation should der this scenario is unlikely (though not impossible) if gene evolve between populations adapting to different environ- flow is present (Schluter 2009; Nosil and Flaxman 2011), ments (Mayr 1942) and is a consequence of divergent selec- whereas ecological speciation is common with or without tion on functional traits enabling specialization for distinct gene flow (Schluter 2009; Nosil and Flaxman 2011; Feder ß The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/ licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. Open Access Mol. Biol. Evol. doi:10.1093/molbev/msaa177 Advance Access publication July 13, 2020 1 Breusing et al. doi:10.1093/molbev/msaa177 MBE et al. 2012; Campbell et al. 2018). Additional processes in- Alviniconcha and Ifremeria are nutritionally dependent on cluding inter-specific hybridization and host–microbe asso- horizontally transmitted, chemosynthetic sulfide-oxidizing ciations have received less attention but are now accepted endosymbionts that are acquired from a diverse community as significant drivers of speciation (Schuler et al. 2016). of environmental bacteria and are harbored within vacuoles Our current understanding of speciation mechanisms in the gill filaments (Windoffer and Giere 1997; Suzuki, Sasaki, in animals stems mostly from evolutionary research on ter- Suzuki, Nogi, et al. 2006; Trembath-Reichertetal.2019). restrial and freshwater organisms, whereas less is known Except for A. boucheti and A. marisindica,whicharecon- about speciation in oceanic environments, in particular strained to symbionts of the class Campylobacteria (formerly Downloaded from https://academic.oup.com/mbe/advance-article/doi/10.1093/molbev/msaa177/5870836 by guest on 24 September 2020 deep ocean basins (Miglietta et al. 2011). Processes that Epsilonproteobacteria; Waite et al. 2017), all other lead to evolutionary divergence in marine systems may differ Alviniconcha species and Ifremeria nautilei typically host from other environments given that physical barriers are less Gammaproteobacteria symbionts (Urakawa et al. 2005; pronounced and deep-sea habitats might have been less af- Suzuki, Sasaki, Suzuki, Nogi, et al. 2005;Suzuki,Sasaki, fected by past climatic fluctuations (Miglietta et al. 2011;but Suzuki, Tsuchida, et al. 2005; Suzuki, Kojima, Sasaki, et al. see Vrijenhoek 2013), at least over much of the Cenozoic. 2006; Suzuki, Kojima, Watanabe, et al. 2006; Beinartetal. Nonetheless, deep-sea hydrothermal vents provide an excep- 2012). In zones of sympatry, the two snail genera occupy tion to other deep-sea environments due to their global but relatively narrow ranges of chemical and thermal conditions that are well within their physiological tolerance limits and island-like distribution and their relative instability on ecolog- that do not support maximal rates for chemoautotrophic ical and geological time scales (Vrijenhoek 2013). The dense growth (Sen et al. 2013). This observation suggests that inter- animal communities living in these habitats are exposed to species competition is an important factor determining the highly variable and extreme environmental conditions, and realized niche of these snails (Sen et al. 2013). Beinart et al. many of the invertebrates are sustained through obligate (2012) proposed that host niche utilization is strongly influ- symbiosis with chemosynthetic bacteria (Dubilier et al. enced by the availability and metabolic capacity of locally dom- 2008). The phylogenetically and ecologically closely related inant symbiont phylotypes. In their study of co-occurring snail genera Alviniconcha and Ifremeria (Gastropoda: Alviniconcha species from the Lau Basin, they showed that Abyssochrysoidea) are among the dominant inhabitants of the distribution and abundance of host–symbiont combina- Indo-Pacific hydrothermal vents. The genus Ifremeria is tions (holobionts) followed a latitudinal gradient in vent geo- monotypic and apparently restricted to Western Pacific vents chemistry, wherein A. boucheti holobionts dominated at intheManus,Lau,andNorthFijibasins.Incontrast,the northern localities characterized by high end-member con- broadly distributed genus Alviniconcha comprises six species centrations of sulfide and hydrogen (3.5–7 mM and 100– (Johnson et al. 2015): Alviniconcha kojimai and Alviniconcha 500 mM, respectively), A. kojimai holobionts dominated at boucheti co-occur in the Manus, Lau, and North Fiji basins; midlatitude localities characterized by medium end- Alviniconcha strummeri is known from the southern Lau member concentrations (2.5–4 mM and 50–100 mM, re- Basin; Alviniconcha hessleri and Alviniconcha adamantis ap- spectively), and A. strummeri holobionts dominated at south- pear to be endemic to the Mariana Back-Arc and Volcanic ern localities characterized by low end-member Arc, respectively; and Alviniconcha marisindica occurs along concentrations of these reductants (1–3 mM and 35– the Central Indian Ridge (fig. 1 and table 1). 135 mM, respectively). These broad-scale patterns of FIG.1.Map of sampling localities for the six Alviniconcha species analyzed in this study. 2 Speciation in Hydrothermal Vent Snails . doi:10.1093/molbev/msaa177 MBE Table 1. Geographic Coordinates, Depths,
Recommended publications
  • A Problematic Zygopleuroid Gastropod Acanthostrophia Revisited 21
    50 /Reihe A Series A Zitteliana An International Journal of Palaeontology and Geobiology Series A /Reihe A Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Geologie 50Jubilee Volume An International Journal of Palaeontology and Geobiology München 2010 Zitteliana Zitteliana An International Journal of Palaeontology and Geobiology Series A/Reihe A Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Geologie 50 CONTENTS/INHALT BABA SENOWBARI-DARYAN & MICHAELA BERNECKER Amblysiphonella agahensis nov. sp., and Musandamia omanica nov. gen., nov. sp. (Porifera) from the Upper Triassic of Oman 3 ALEXANDER NÜTZEL A review of the Triassic gastropod genus Kittliconcha BONARELLI, 1927 – implications for the phylogeny of Caenogastropoda 9 ANDRZEJ KAIM & MARIA ALESSANDRA CONTI A problematic zygopleuroid gastropod Acanthostrophia revisited 21 GERNOT ARP Ammonitenfauna und Stratigraphie des Grenzbereichs Jurensismergel/Opalinuston- Formation bei Neumarkt i.d. Opf. (oberstes Toarcium, Fränkische Alb) 25 VOLKER DIETZE Über Ammonites Humphriesianus umbilicus QUENSTEDT, 1886 an seiner Typus-Lokalität (östliche Schwäbische Alb, Südwestdeutschland) 55 VOLKER DIETZE, GÜNTER SCHWEIGERT, GERD DIETL, WOLFGANG AUER, WOLFGANG DANGELMAIER, ROGER FURZE, STEFAN GRÄBENSTEIN, MICHAEL KUTZ, ELMAR NEISSER, ERICH SCHNEIDER & DIETMAR SCHREIBER Rare Middle Jurassic ammonites of the families Erycitidae, Otoitidae and Stephanoceratidae from southern Germany 71 WOLFGANG WITT Late Miocene non-marine ostracods from the Lake Küçükçekmece region, Thrace (Turkey) 89 JÉRÔME PRIETO Note on the morphological variability of Keramidomys thaleri (Eomyidae, Mammalia) from Puttenhausen (North Alpine Foreland Basin, Germany) 103 MARTIN PICKFORD Additions to the DEHM collection of Siwalik hominoids, Pakistan: descriptions and interpretations 111 MICHAEL KRINGS, NORA DOTZLER, THOMAS N. TAYLOR & JEAN GALTIER Microfungi from the upper Visean (Mississippian) of central France: Structure and development of the sporocarp Mycocarpon cinctum nov.
    [Show full text]
  • A New Fossil Provannid Gastropod from Miocene Hydrocarbon Seep Deposits, East Coast Basin, North Island, New Zealand
    A new fossil provannid gastropod from Miocene hydrocarbon seep deposits, East Coast Basin, North Island, New Zealand KRISTIAN P. SAETHER, CRISPIN T.S. LITTLE, and KATHLEEN A. CAMPBELL Saether, K.P., Little, C.T.S., and Campbell, K.A. 2010. A new fossil provannid gastropod from Miocene hydrocarbon seep deposits, East Coast Basin, North Island, New Zealand. Acta Palaeontologica Polonica 55 (3): 507–517. Provanna marshalli sp. nov. is described from Early to Middle Miocene−age fossil hydrocarbon seep localities in the East Coast Basin, North Island, New Zealand, adding to 18 modern and three fossil species of the genus described. Modern species are well represented at hydrothermal vent sites as well as at hydrocarbon seeps and on other organic substrates in the deep sea, including sunken wood and whale falls. Described fossil Provanna species have been almost exclusively re− ported from hydrocarbon seep deposits, with a few reports of suspected fossil specimens of the genus from other chemosynthetic environments such as sunken wood and large vertebrate (whale and plesiosaurid) carcasses, and the old− est occurrences are dated to the Middle Cenomanian (early Late Cretaceous). The New Zealand fossil species is the most variable species of the genus described to date, and its shell microstructure is reported and found to be comparable to the fossil species Provanna antiqua and some modern species of the genus. Key words: Mollusca, Gastropoda, Provannidae, Provanna, hydrocarbon seeps, Miocene, East Coast Basin, New Zealand. Kristian P. Saether [[email protected]] and Kathleen A. Campbell [[email protected]], School of Environment, Faculty of Science, University of Auckland, Private Bag 92019, Auckland Mail Centre, Auckland 1142, New Zealand; Crispin T.S.
    [Show full text]
  • Caenogastropoda
    13 Caenogastropoda Winston F. Ponder, Donald J. Colgan, John M. Healy, Alexander Nützel, Luiz R. L. Simone, and Ellen E. Strong Caenogastropods comprise about 60% of living Many caenogastropods are well-known gastropod species and include a large number marine snails and include the Littorinidae (peri- of ecologically and commercially important winkles), Cypraeidae (cowries), Cerithiidae (creep- marine families. They have undergone an ers), Calyptraeidae (slipper limpets), Tonnidae extraordinary adaptive radiation, resulting in (tuns), Cassidae (helmet shells), Ranellidae (tri- considerable morphological, ecological, physi- tons), Strombidae (strombs), Naticidae (moon ological, and behavioral diversity. There is a snails), Muricidae (rock shells, oyster drills, etc.), wide array of often convergent shell morpholo- Volutidae (balers, etc.), Mitridae (miters), Buccin- gies (Figure 13.1), with the typically coiled shell idae (whelks), Terebridae (augers), and Conidae being tall-spired to globose or fl attened, with (cones). There are also well-known freshwater some uncoiled or limpet-like and others with families such as the Viviparidae, Thiaridae, and the shells reduced or, rarely, lost. There are Hydrobiidae and a few terrestrial groups, nota- also considerable modifi cations to the head- bly the Cyclophoroidea. foot and mantle through the group (Figure 13.2) Although there are no reliable estimates and major dietary specializations. It is our aim of named species, living caenogastropods are in this chapter to review the phylogeny of this one of the most diverse metazoan clades. Most group, with emphasis on the areas of expertise families are marine, and many (e.g., Strombidae, of the authors. Cypraeidae, Ovulidae, Cerithiopsidae, Triphori- The fi rst records of undisputed caenogastro- dae, Olividae, Mitridae, Costellariidae, Tereb- pods are from the middle and upper Paleozoic, ridae, Turridae, Conidae) have large numbers and there were signifi cant radiations during the of tropical taxa.
    [Show full text]
  • Intracellular Oceanospirillales Inhabit the Gills of the Hydrothermal Vent Snail Alviniconcha with Chemosynthetic, Proteobacteri
    bs_bs_banner Environmental Microbiology Reports (2014) doi:10.1111/1758-2229.12183 Intracellular Oceanospirillales inhabit the gills of the hydrothermal vent snail Alviniconcha with chemosynthetic, γ-Proteobacterial symbionts R. A. Beinart,1 S. V. Nyholm,2 N. Dubilier3 and could play a significant ecological role either as a P. R. Girguis1* host parasite or as an additional symbiont with 1Department of Organismic and Evolutionary Biology, unknown physiological capacities. Harvard University, Cambridge, MA 02138, USA. 2Department of Molecular and Cell Biology, University of Introduction Connecticut, Storrs, CT 06269, USA. In recent years, lineages from the γ-Proteobacterial order 3Symbiosis Group, Max Planck Institute for Marine Oceanospirillales have emerged as widespread associ- Microbiology, Bremen 28359, Germany. ates of marine invertebrates. In shallow-water habitats, Oceanospirillales are common and even dominant Summary members of the tissue and mucus-associated microbiota of temperate and tropical corals (Sunagawa et al., 2010; Associations between bacteria from the γ- Bayer et al., 2013a,b; Bourne et al., 2013; Chen et al., Proteobacterial order Oceanospirillales and marine 2013; La Rivière et al., 2013) and sponges (Kennedy invertebrates are quite common. Members of the et al., 2008; Sunagawa et al., 2010; Flemer et al., 2011; Oceanospirillales exhibit a diversity of interac- Bayer et al., 2013a,b; Bourne et al., 2013; Chen et al., tions with their various hosts, ranging from the 2013; La Rivière et al., 2013; Nishijima et al., 2013), and catabolism of complex compounds that benefit host they have been detected in the gills of commercially growth to attacking and bursting host nuclei. Here, important shellfish (Costa et al., 2012), as well as invasive we describe the association between a novel oysters (Zurel et al., 2011).
    [Show full text]
  • A New Late Pliocene Large Provannid Gastropod Associated with Hydrothermal Venting at Kane Megamullion, Mid-Atlantic Ridge
    A new Late Pliocene large provannid gastropod associated with hydrothermal venting at Kane Megamullion, Mid-Atlantic Ridge Andrzej Kaima*, Brian E. Tucholkeb, and Anders Warénc aBayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Str. 10, 80333 München, Germany and Instytut Paleobiologii PAN, ul. Twarda 51/55, 00-818 Warszawa, Poland. bDepartment of Geology and Geophysics, Woods Hole Oceanographic Institution, Woods Hole, MA 02543 USA. cDepartment of Invertebrate Zoology, Swedish Museum of Natural History, Box 50007, SE-10405 Stockholm, Sweden. *[email protected] Abstract A new gastropod, Kaneconcha knorri gen et sp. nov., was found in marlstone dredged from the surface of Adam Dome at Kane Megamullion on the flank of the Mid-Atlantic Ridge in an area of former hydrothermal activity. The snail is interpreted as a large provannid similar to the chemosymbiotic genera Ifremeria and Alviniconcha. This is the first record of presumably chemosymbiotic provannids from the Atlantic Ocean and also the first fossil record of such large provannids associated with hydrothermal venting. Extant Alviniconcha and Ifremeria are endemic to hydrothermal vents in the Pacific and Indian oceans. Kaneconcha differs from Ifremeria in having no umbilicus and a posterior notch, and it differs from Alviniconcha in having the profile of the whorl slightly flattened and having no callus on the inner lip. A dark layer covering the Kaneconcha shell is interpreted here as a fossilized periostracum. The shell/periostracum interface shows fungal traces attributed to the ichnospecies Saccomorpha clava. We hypothesize that large chemosymbiotic provannids (i.e., Kaneconcha, Ifremeria, and Alviniconcha) form a clade that possibly diverged from remaining provannids in the Late Jurassic, with the Late Jurassic/Early Cretaceous Paskentana being an early member.
    [Show full text]
  • Earliest Known (Campanian) Members of the Vermetidae, Provannidae and Litiopidae (Cerithioidea, Gastropoda), and a Discussion of Their Possible Relationships
    Mitt. Geol.-Palaont. Inst. Univ. Hamburg Earliest known (Campanian) members of the Vermetidae, Provannidae and Litiopidae (Cerithioidea, Gastropoda), and a discussion of their possible relationships KLAUS BANDEL & STEFFENKIEL, Hamburg *) With 7 Figures Abstract 209 Zusammenfassung 2W I. Introduction 210 II. Material and methods 211 III. Systematic descriptions 212 IV. Discussion 215 Acknowledgements 217 References 217 The newly discovered Campanian species Vermetus nielseni n. sp., Desbruyeresia antigua n. sp. and Litiopella schoeningi n. gen. n. sp. are described and the taxonomy of these gastropod groups is reassessed. Based on their protoconch morphology and radula characters, the Dendropominae, Provannidae, Litiopidae and Sculptifer are considered as related taxa within the Cerithioidea. They are interpreted to have arisen from a common ancestor that lived during the Cretaceous, apparently parallel to the radiation of the Vermetidae. Die neuen campanischen Arten Vermetus nielseni n. sp., Desbruyeresia antigua n. sp. und Litiopella schoeningi n. gen., n. sp. werden beschrieben und die Taxonomie dieser Gastropoden- *) Authors addresses: Prof. Dr. Klaus BANDEL& Steffen KIEL,Geologisch-Palaontologisches lnstitut und Museum, Universitat Hamburg, BundesstraBe 55,20146 Hamburg, Germany. e-mails:[email protected]@grnx.de Gruppen neu bewertet. Basierend auf der Morphologie ihrer Protoconche und Radulae werden die Dendropominae, Provannidae, Litiopidae und Sculptifer als verwandte Taxa innerhalb der Cerithioidea angesehen, die sich wahrscheinlich aus einem gemeinsamen kretazischen Vorfahren entwickelten. Die Entwicklung dieser Gruppe verlief offensichtlich parallel zur Radiation der Vermetidae, deren Vertre- ter jedoch eine andere Protoconchmorphologie zeigen. Vermetids are sessile marine gastropods with a tubular shell that is irregularly coiled and totally or partly cemented to hard substrates.
    [Show full text]
  • (Gastropoda: Abyssochrysoidea) Discovered from the Shinkai Seep
    Journal of the Marine Biological Association of the United Kingdom, page 1 of 9. # Marine Biological Association of the United Kingdom, 2016 doi:10.1017/S0025315416001648 A very deep Provanna (Gastropoda: Abyssochrysoidea) discovered from the Shinkai Seep Field, Southern Mariana Forearc chong chen1, hiromi kayama watanabe2 and yasuhiko ohara1,3 1Department of Subsurface Geobiological Analysis and Research (D-SUGAR), Japan Agency for Marine-Earth Science and Technology (JAMSTEC), 2-15 Natsushima-cho, Yokosuka, Kanagawa, 237-0061, Japan, 2Department of Marine Biodiversity Research (BIO-DIVE), Japan Agency for Marine-Earth Science and Technology (JAMSTEC), 2-15 Natsushima-cho, Yokosuka, Kanagawa, 237-0061, Japan, 3Hydrographic and Oceanographic Department of Japan, Building 4, 3-1-1 Kasumigaseki, Chiyoda-ku, Tokyo 100-8932, Japan The ‘Shinkai Seep Field’ is a serpentinite-hosted chemosynthetic ecosystem in the Southern Mariana Forearc. In June 2015 the site was revisited and a number of rissoiform gastropods were collected. Taxonomic investigations revealed that these speci- mens represent a hitherto undescribed species of Provanna (Gastropoda: Abyssochrysoidea), described herein as Provanna cingulata n. sp. This new species is characterized by numerous spiral keels, lack of significant axial sculpture, rounded and inflated whorls, and large size for the genus. With the shell height exceeding 16.5 mm (may reach 20 mm), it is the largest Provanna species known thus far. Phylogenetic analysis using 411 bp of the cytochrome oxidase c subunit I (COI) gene confirmed its systematic placement within the genus Provanna. This is the only gastropod from a family endemic to chemosynthetic ecosystems thus far known from the ‘Shinkai Seep Field’.
    [Show full text]
  • Mollusks and a Crustacean from Early Oligocene Methane-Seep Deposits in the Talara Basin, Northern Peru
    Mollusks and a crustacean from early Oligocene methane-seep deposits in the Talara Basin, northern Peru STEFFEN KIEL, FRIDA HYBERTSEN, MATÚŠ HYŽNÝ, and ADIËL A. KLOMPMAKER Kiel, S., Hybertsen, F., Hyžný, M., and Klompmaker, A.A. 2020. Mollusks and a crustacean from early Oligocene methane- seep deposits in the Talara Basin, northern Peru. Acta Palaeontologica Polonica 65 (1): 109–138. A total of 25 species of mollusks and crustaceans are reported from Oligocene seep deposits in the Talara Basin in north- ern Peru. Among these, 12 are identified to the species-level, including one new genus, six new species, and three new combinations. Pseudophopsis is introduced for medium-sized, elongate-oval kalenterid bivalves with a strong hinge plate and largely reduced hinge teeth, rough surface sculpture and lacking a pallial sinus. The new species include two bivalves, three gastropods, and one decapod crustacean: the protobranch bivalve Neilo altamirano and the vesicomyid bivalve Pleurophopsis talarensis; among the gastropods, the pyropeltid Pyropelta seca, the provannid Provanna pelada, and the hokkaidoconchid Ascheria salina; the new crustacean is the callianassid Eucalliax capsulasetaea. New combina- tions include the bivalves Conchocele tessaria, Lucinoma zapotalensis, and Pseudophopsis peruviana. Two species are shared with late Eocene to Oligocene seep faunas in Washington state, USA: Provanna antiqua and Colus sekiuensis; the Talara Basin fauna shares only genera, but no species with Oligocene seep fauna in other regions. Further noteworthy aspects of the molluscan fauna include the remarkable diversity of four limpet species, the oldest record of the cocculinid Coccopigya, and the youngest record of the largely seep-restricted genus Ascheria.
    [Show full text]
  • 1 Expanding Dispersal Studies at Hydrothermal Vents Through Species Identification Of
    *Manuscript Click here to download Manuscript: Adams_revised2.doc Click here to view linked References 1 Expanding dispersal studies at hydrothermal vents through species identification of 2 cryptic larval forms 3 4 D. K. Adams* 5 S. W. Mills 6 T. M. Shank 7 L. S. Mullineaux 8 9 Biology Department, Woods Hole Oceanographic Institution, Woods Hole, MA 02543 10 USA 11 12 * Corresponding author; current address: National Institutes of Health, NIDCR, Bethesda 13 MD 20982 USA; email: [email protected]; Tel: (301) 496-1392; Fax: (301) 480-5353 14 15 Abstract 16 The rapid identification of hydrothermal vent-endemic larvae to the species level is a key 17 limitation to understanding the dynamic processes that control the abundance and 18 distribution of fauna in such a patchy and ephemeral environment. Many larval forms 19 collected near vents, even those in groups such as gastropods that often form a 20 morphologically distinct larval shell, have not been identified to species. We present a 21 staged approach that combines morphological and molecular identification to optimize 22 the capability, efficiency, and economy of identifying vent gastropod larvae from the 23 northern East Pacific Rise (NEPR). With this approach, 15 new larval forms can be 1 1 identified to species. A total of 33 of the 41 gastropod species inhabiting the NEPR, and 2 26 of the 27 gastropod species known to occur specifically in the 9° 50‟ N region, can be 3 identified to species. Morphological identification efforts are improved by new 4 protoconch descriptions for Gorgoleptis spiralis, Lepetodrilus pustulosus, Nodopelta 5 subnoda, and Echinopelta fistulosa.
    [Show full text]
  • Variations in Seep Mussel Bed Communities Along Physical and Chemical Environmental Gradients
    MARINE ECOLOGY PROGRESS SERIES Vol. 293: 99–108, 2005 Published June 2 Mar Ecol Prog Ser Variations in seep mussel bed communities along physical and chemical environmental gradients Derk C. Bergquist1, 3,*, Clint Fleckenstein1, Julie Knisel1, Brett Begley1, Ian R. MacDonald2, Charles R. Fisher1 1Department of Biology, Pennsylvania State University, 208 Mueller Laboratory, University Park, Pennsylvania 16802, USA 2Physical and Life Sciences Department, Texas A&M University — Corpus Christi, 6300 Ocean Dr. ST320, Corpus Christi, Texas 78412, USA 3Present address: Department of Zoology, University of Florida, 223 Bartram Hall, Gainesville, Florida 32611-8525, USA ABSTRACT: This study examined the faunal communities associated with beds of the hydrocarbon seep mussel Bathymodiolus childressi in the Gulf of Mexico and the relationships between these communities and local environmental characteristics. Mussels, their associated fauna and accompa- nying water chemistry parameters were collected at 17 locations across 4 hydrocarbon seep sites. A total of 19 species, mostly seep endemics, were identified. Species richness and fauna density were significantly and negatively correlated with methane concentration and were significantly and posi- tively correlated with oxygen concentration. Canonical Correspondence Analysis (CCA) separated polychaetes, alvinocarid shrimp and gastropods along gradients of methane and oxygen concentra- tions. Polychaetes and alvinocarid shrimp tended towards lower oxygen and higher methane concen- trations and gastropods tended towards higher oxygen and lower methane. The larger-sized preda- ceous gastropods and decapods were associated with lower sulfide and higher oxygen concentrations. The proportion of the community comprising of endemic species was significantly and positively correlated with mussel biomass and shell surface area. Other measures of habitat com- plexity (mussel density, shell surface area and biomass) explained little of the variation in the associ- ated faunal communities.
    [Show full text]
  • Four New Species of Provanna (Gastropoda: Provannidae) from Vents and a Seep Off Nansei-Shoto Area, Southwestern Japan
    VENUS 74 (1–2): 1–17, 2016 New Species of Provanna from SW Japan ©Malacological Society of Japan1 Four New Species of Provanna (Gastropoda: Provannidae) from Vents and a Seep off Nansei-shoto Area, Southwestern Japan Takenori Sasaki1*, Tomomi Ogura2,3, Hiromi Kayama Watanabe3 and Katsunori Fujikura2,3 1The University Museum, The University and Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan 2Graduate School of Marine Science and Technology, Tokyo University of Marine Science and Technology, 4-5-7 Konan, Minato-ku, Tokyo 108-8477, Japan 3Japan Agency for Marine-Earth Science and Technology, 2-15 Natsushima-cho, Yokosuka-shi, Kanagawa 237-0061, Japan Abstract: Members of the genus Provanna are abundant and widely distributed in deep-sea chemosynthetic environments. Around Japan provannids inhabit a wide range of depths and substrates. In this study, we report additional species of Provanna from vents and a seep in the Nansei-shoto area. Shell morphological and molecular phylogenetic analyses showed that there are four new species: (1) P. subglabra is characterized by an inated smooth shell and is most abundant in vent elds in the Okinawa Trough; (2) P. clathrata has a roughly sculptured shell and is distributed in vents in the southern Okinawa Trough; (3) P. lucida possesses a thin smooth shell with a deep suture and is limited to vents on the Minami-Ensei Knoll in the northern Okinawa Trough; and (4) P. kuroshimensis, which is distinguished by an olive-colored periostracum and is endemic to a seep site on the Kuroshima Knoll. In contrast to shell and molecular characters, radula morphology does not show clear differences among these species.
    [Show full text]
  • Miocene Abyssochrysoid Gastropod Provanna from Japanese Seep and Whale-Fall Sites
    Miocene abyssochrysoid gastropod Provanna from Japanese seep and whale-fall sites KAZUTAKA AMANO and CRISPIN T.S. LITTLE Amano, K. and Little, C.T.S. 2014. Miocene abyssochrysoid gastropod Provanna from Japanese seep and whale-fall sites. Acta Palaeontologica Polonica 59 (1): 163–172. We describe three Miocene species of Provanna from Japan, two new and one in open nomenclature, that represent the only known fossil examples from whale-falls and a considerable increase in the Miocene diversity of the genus. Provanna hirokoae sp. nov. comes from the latest Middle Miocene Kuroiwa seep site in central Honshu. The shells of this species are mostly recrystallized, but contain relict crossed lamellar microstructures. Provanna alexi sp. nov. is from the early Middle Miocene Shosanbetsu whale-fall site in northwestern Hokkaido, and has well preserved shells comprising an outer simple prismatic layer and an inner crossed lamellar layer. The two Provanna specimens from the Middle Miocene Rekifune whale-fall site, in eastern Hokkaido, are preserved as external moulds only, so are left in open nomenclature. Based on current knowledge, the presence of an outer prismatic layer and an underlying crossed lamellar layer seems to be a common feature in the shells of Provanna, as well as in other genera belonging to the family Provan- nidae and the superfamily Abyssochrysoidea. Although the oldest occurrence of Provanna was in the Late Cretaceous, the genus did not spread geographically and ecologically until the Miocene (with four, or possibly five species), a date concordant with some molecular estimates. However, this could be an artefact of the fossil record because the known pre-Miocene seep and whale-falls are more geographically restricted than those from the Miocene.
    [Show full text]