Marine Mussels. Elizabeth Gosling

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1.6 Vent mussels and associated fauna are bathed in hydrothermal fluids at the Wideawake vent field on the Mid‐Atlantic Ridge. Photo taken by the remotely operated, deep‐sea submersible MARUM‐QUEST.

      Source: From Orphan & Hoehler (2011). Reproduced with permission from Nature Publishing Group.

      Although these mussels were initially regarded as archaic, molecular age estimates and fossil records suggest that most of the modern vent and seep animals appeared during a short time interval between the Late Mesozoic and the Early Cenozoic, less than 100 mya (Van Dover et al. 2002). The evolutionary history of deep‐sea symbiotic mussels has been investigated over the past two decades using a range of mitochondrial and nuclear DNA markers. Distel et al. (2000) were the first to propose the stepping stone hypothesis (i.e. that vent and seep mussels derived from ancestors associated with wood and whale fall ecosystems). Results from subsequent phylogenetic studies have strongly supported this proposal (Samadi et al. 2007; Miyazaki et al. 2010; Lorion et al. 2013; Thubaut et al. 2013). Rather than a single step toward colonisation of vents and seeps, there were multiple habitat shifts from organic substrates to vents and cold seeps, always in the one direction (Thubaut et al. 2013). Cold seeps seem to be an intermediate habitat for the colonisation of deep vents (Thubaut et al. 2013). Ancestors of Bathymodiolus mussels were shallow species that acquired the ability to associate with bacteria, most likely sulphur oxidisers (Duperron 2010). For mussels, the acquisition of such symbionts is seen as a prerequisite for their adaptation to, and successful radiation within, chemosynthetic environments (Lorion et al. 2013). Subsequent acquisition of methanotrophic symbionts allowed the colonisation of new niches within the vent and seep habitats, resulting in a second wave of diversification (Lorion et al. 2013). To date, there is no general consensus on the phylogenetic relationships of deep‐sea Bathymodiolus mussels and their mytilid relatives (see Jones et al. 2006; Fujita et al. 2009; Miyazaki et al. 2010; Thubaut et al. 2013; Oliver 2015).

      Notes

      1 1 A system of classification based on the phylogenetic relationships and evolutionary history of groups of organisms, rather than purely on shared features.

      2 2 A monophyletic group of organisms believed to comprise all the evolutionary descendants of a common ancestor.

      3 3 The classification of organisms and the evolutionary relationships among them.

      4 4 A polyphyletic taxon is defined as one that does not include the common ancestor of all members of the taxon.

      5 5 A paraphyletic taxon is one that includes the most recent common ancestor but not all of its descendants.

      1 Bieler, R. & Mikkelsen, P.A. (2006) Bivalvia – a look at the branches. Zoological Journal of the Linnean Society, 148, 223–235.

      2 Bieler, R., Carter, J.G. & Coan, E.V. (2010) Classification of bivalve families. In: Nomenclator of Bivalve Families (eds P. Bouchet & J.‐P. Rocroi), pp. 113–133. Conch Books, Hackenheim.

      3 Bieler, R., Mikkelsen, P.M., Collins, T.M., Glover, E.A., González, V.L., Graf, D.L. et al. (2014) Investigating the bivalve tree of life – an exemplar‐based approach combining molecular and novel morphological characters. Invertebrate Systematics, 28, 32–115.

      4 Boss, K.J. (1982) Mollusca. In: Synopsis and Classification of Living Organisms, 1st edn (ed S.P. Parker), pp. 945–1166. McGraw‐Hill, New York.

      5 Cannuel, R., Beninger, P.G., McCombie, H. & Boudry, P. (2009) Gill development and its functional and evolutionary implications in the blue mussel Mytilus edulis (Bivalvia: Mytilidae). Biological Bulletin, 217, 173–188.

      6 Carter, J.G., Campbell, D.C. & Campbell, M.R. (2000) Cladistic perspectives on early bivalve evolution. In: The Evolutionary Biology of the Bivalvia (eds E.M. Harper, J.D. Taylor & J.A. Crame), pp. 47–79. Geological Society, London.

      7 Carter, J.G., Altaba, C.R., Anderson, L.C., Araujo, R., Biakov, A.S., Bogan, A.E. et al. (2011) A synoptical classification of the Bivalvia (Mollusca). Paleontological Contributions, 4, 1–47.

      8 Coan, E.V. & Valentich‐Scott, P. (2012) Bivalve Seashells of Tropical West America. Marine Bivalve Mollusks from Baja California to Northern Peru. Santa Barbara Museum of Natural History, Santa Barbara, CA.

      9 Coan, E.V., Valentich‐Scott, P. & Bernard, F.R. (2000) Bivalve Seashells of Western North America. Bivalve Mollusks from Arctic Alaska to Baja California. Santa Barbara Museum of Natural History, Santa Barbara, CA.

      10 Cope, J.C.W. (2000) A new look at early bivalve phylogeny. In: The Evolutionary Biology of the Bivalvia (eds E.M. Harper, J.D. Taylor & J.A. Crame), pp. 81–95. Geological Society, London.

      11 Distel, D.L. (2000) Phylogenetic relationships among Mytilidae (Bivalvia): 18S rRNA data suggest convergence in mytilid body plans. Molecular Phylogenetics and Evolution, 15, 25–33.

      12 Distel, D.L., Baco, A.R., Chuang, E., Morrill, W., Cavanaugh, C. & Smith, C.R. (2000) Marine ecology: do mussels take wooden steps to deep‐sea vents? Nature, 403, 725–726.

      13 Duperron, S. (2010) The diversity of deep‐sea mussels and their bacterial symbioses. In: The Vent and Seep Biota (ed S. Kiel), pp. 137–167. Springer, New York.

      14 Duperron, S., Halary, S., Lorion, J., Sibuet, M. & Gaill F. (2008) Unexpected co‐occurrence of six bacterial symbionts in the gills of the cold seep mussel Idas sp. (Bivalvia: Mytilidae). Environmental Microbiology, 10, 433–445.

      15 Duperron, S., Lorion, J., Samadi, S., Gros, O. & Gaill, F. (2009) Symbioses between deep‐sea mussels (Mytilidae: Bathymodiolinae) and chemosynthetic bacteria: diversity, function and evolution. Comptes Rendus Biologies, 332, 298–310.

      16 Fujita, Y., Matsumoto, H., Fujiwara, Y., Hashimoto, J., Galkin, S.V., Ueshima, R. et al. (2009) Phylogenetic relationships of deep‐sea Bathymodiolus mussels to their mytilid relatives from sunken whale carcasses and wood. Venus, 67, 123–134.

      17 Giribet, G. (2008) Bivalvia. In: Phylogeny and Evolution of the Mollusca (eds W. Ponder & D.R. Lindberg), pp. 105–141. University of California Press, Berkeley, CA.

      18 Giribet, G. & Wheeler, W.C. (2002) On bivalve phylogeny: a high‐level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data. Invertebrate Biology, 121, 271–324.

      19 Glynn, P.W. & Manzello, D.P. (2015) Bioerosion and coral reef growth: a dynamic balance. In: Coral Reefs in the Anthropocene (ed C. Birkeland), pp. 67–97. Springer, New York.

      20 González, V.L. & Giribet, G. (2014) A multilocus phylogeny of archiheterodont bivalves (Mollusca, Bivalvia, Archiheterodonta). Zoologica Scripta, 44, 41–58.

      21 González, V.L., Andrade, S.C.S., Bieler, R., Collins, T.M., Dunn, C.W., Mikkelsen, P.M. et al. (2015) A phylogenetic backbone for Bivalvia: an RNA‐seq approach.

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