Skip to main content

Functional morphology of Cardiomya cleryana (d'Orbigny, 1842) (Bivalvia: Anomalodesmata: Cuspidariidae) from Brazilian waters: new insights into the lifestyle of carnivorous bivalves

  • Fabrizio Marcondes Machado (a1), Brian Morton (a2) and Flávio Dias Passos (a3)

Of the more than 800 Recent species of Anomalodesmata only 16 have been studied alive. The Septibranchia, comprising a number of carnivorous bivalve superfamilies, are no exception to this generalization and, until recently, no living member of this group has been studied since the 1980s. Collected from the shallow, shelf seabed off Brazil, Cardiomya cleryana is one of only a few species to have its morphology described in recent years and the first ever anomalodesmatan to be filmed alive. Important anatomical features such as a greatly extensible foot to secure itself in the sediment with a single byssal thread, exhalant siphon inter-tentacular projections and micro-papillae on the surface of the siphonal tentacles are described. Observations on the species’ behaviour have revealed a lifestyle hitherto not recorded for any septibranch, indeed any anomalodesmatan. The anchoring mechanism of C. cleryana using an extraordinarily long byssal thread is described for the first time. The life position of C. cleryana is at 45° to the sediment-water interface whereas other cuspidariids have been considered to orient themselves vertically. An anatomical comparison between Cardiomya and Bathyneaera has revealed close affinities. This study thus provides new insights into the morphology and behaviour of the Cuspidariidae.

Corresponding author
Correspondence should be addressed to: F. M. Machado, Programa de Pós-Graduação em Biologia Animal, Universidade Estadual de Campinas (UNICAMP), CEP 13083-970, Campinas, SP, Brasil email:
Hide All
Absalão, R.S., Caetano, C.H.S. and Pimenta, A.D. (2003) New records of marine gastropods and bivalves from Brazil (Mollusca). Revista Brasileira de Zoologia 20, 323328. [In Portuguese]
Absalão, R.S. and Oliveira, C.D.C. (2011) The genus Cuspidaria (Pelecypoda: Septibranchia: Cuspidariidae) from the deep sea of Campos Basin, Brazil, with descriptions of two new species. Malacologia 54, 119138.
Absalão, R.S. and Pimenta, D.A. (2005) Moluscos marinhos da APA do Arquipélago de Santana. In Macaé, R.J. (ed.) Chave ilustrada para identificação das espécies do substrato inconsolidado. Rio de Janeiro: Editora Ciência Moderna. [In Portuguese]
Allen, J.A. (2008) Bivalvia of the deep Atlantic. Malacologia 50, 57173.
Allen, J.A. (2011) Descriptions of a new deep-water species of the family Cuspidariidae (Mollusca: Bivalvia) and including a bibliography of the known species from the Atlantic. Journal of Conchology 40, 428445.
Allen, J.A. and Morgan, R.E. (1981) The functional morphology of Atlantic deep water species of the families Cuspidariidae and Poromyidae (Bivalvia): an analysis of the evolution of the septibranch condition. Philosophical Transactions of the Royal Society of London, Series B 294, 413546.
Allen, J.A. and Turner, J.F. (1974) On the functional morphology of the family Verticordiidae (Bivalvia) with descriptions of new species from the abyssal Atlantic. Philosophical Transactions of the Royal Society of London, Series B 268, 401520.
Altena, C.O. Van, Regteren (1971) On six species of marine Mollusca from Suriname, four of which are new. Zoologische Mededelingen 45, 7586.
Bernard, F.R. (1989) Six new species of the order Septibranchia (Bivalvia) from the northeastern Pacific Ocean. Venus 48, 6166.
Bieler, R., Carter, J.G. and Coan, E.V. (2010) Classification of bivalve families. In Bouchet, P. and Rocroi, J.P. (eds) Nomenclator of bivalve families, pp. 113133. Malacologia 52, 1–184.
Bieler, R. and Gofas, S. (2015) Cuspidarioidea Dall, 1886. In MolluscaBase (2015). Accessed through: World Register of Marine Species. Available at (accessed 19 November 2015).
Bieler, R., Mikkelsen, P.M., Collins, T.M., Glover, E.A., Gonzaléz, V.L., Graf, D.L., Harper, E.M., Healy, J., Kawauchi, G.Y., Sharma, P.P., Staubach, S., Strong, J., Taylor, J.D., Temkin, I., Zardus, J.D., Clark, S., Guzmán, A., Mcintyre, E., Sharp, P. and Giribet, G. (2014) Investigating the bivalve tree of life–an exemplar-based approach combining molecular and novel morphological characters. Invertebrate Systematics 28, 32115.
Boehls, G., Absher, T.M. and Cruz-Kaled, A.D. (2004) Composition and distribution of benthic mollusks on intertidal flats of Paranaguá Bay (Paraná, Brazil). Scientia Marina 68, 537543.
Carcelles, A.R. (1950) Catálogo de los moluscos marinos de la Patagonia. Anales del Museu Nahuel Huapi 2, 4199. [In Spanish]
Coan, E.V. and Valentich-Scott, P. (2012) Bivalve seashells of tropical West America: marine bivalve mollusks from Baja California to Northern Peru. Santa Barbara, CA: Santa Barbara Museum of Natural History Monographs No. 6, Studies in Biodiversity No. 4.
Coan, E.V., Valentich-Scott, P. and Bernard, F.R. (2000) Bivalve seashells of western North America: marine bivalve mollusks from Arctic Alaska to Baja California. Santa Barbara, CA: Santa Barbara Museum of Natural History Monographs No. 2, Studies in Biodiversity 2.
Domaneschi, O. (1979) Aspectos da biologia de Diplodonta punctata (Say, 1822) (Bivalvia: Lucinacea: Ungulinidae). Revista Nordestina de Biologia 2, 2125. [In Portuguese]
d'Orbigny, A. (1834–1847) Voyage dans l'Amérique Méridionale […] Exécute Pendent les Années 1826, […] 1833. Volume 5(3), Mollusques, pp. 529–600 [1845]; Paris Bertrand, Paris & Levrault, Strasbourg [dates teste C. D. Sherborn & F. J. Griffin. 1934. On the dates of publications of the natural history portions of Alcide d'Orbigny's ‘Voyage Amérique Méridionale’. Annals and Magazine of Natural History, Series 10, 13(73), 130–134].
d'Orbigny, A. (1842–1853) Mollusques. In de la Sagra, R. (ed.) Histoire physique, politique, et naturelle de I'lle de Cuba. Volume 2, pp. [iv] + 225–380, pls. 26–28? [1853]. Paris: Bertrand.
Dottori, M., Siegle, E. and Castro, M.B. (2015) Hydrodynamics and water properties at the entrance of Araçá Bay, Brazil. Ocean Dynamics 65, 17311741.
Dufour, S.C. and Felbeck, H. (2003) Sulphide mining by the superextensile foot of symbiotic thyasirid bivalves. Nature (London) 426, 6567.
Gofas, S. (2015) Anomalodesmata. In MolluscaBase (2015). Accessed through: World Register of Marine Species. Available at (accessed 19 November 2015).
Gofas, S. and Bouchet, P. (2015a) Cuspidariidae. In MolluscaBase (2015). Accessed through: World Register of Marine Species. Available at (accessed 19 November 2015).
Gofas, S. and Bouchet, P. (2015b) Cardiomya A. Adams, 1864. In MolluscaBase (2015). Accessed through: World Register of Marine Species. Available at (accessed 21 October 2015).
Grillo, M.C.G., Ventura, C.R.R. and Silva, S.H.G. (1998) Spatial distribution of bivalvia (Mollusca) in the soft-bottoms of Ilha Grande Bay, Rio de Janeiro, Brazil. Revista Brasileira de Oceanografia 46, 1931.
Harper, E.M., Hide, E.A. and Morton, B. (2000) Relationship between the extant Anomalodesmata: a cladistic test. In Harper, E.M., Taylor, J.D. and Crame, J.A. (eds) The evolutionary biology of the Bivalvia. Volume 177. London: Geological Society, pp. 129143.
Knudsen, J. (1970) The systematics and biology of abyssal and hadal Bivalvia. Galathea 11, 7241.
Krylova, H.M. (1993) Bivalve molluscs of the genus Bathyneaera (Septibranchia, Cuspidariidae) of the World Ocean. Ruthenica 3, 5159.
Krylova, H.M. (1995) Clams of the family Protocuspidariidae (Septibranchia, Cuspidarioidea): taxonomy and distribution. Zoologicheskii Zhurnal 74, 2038. [In Russian]
Leal, J.H. (2008) A remarkable new genus of carnivorous, sessile bivalves (Mollusca: Anomalodesmata: Poromyidae) with descriptions of two new species. Zootaxa 1764, 118.
Machado, F.M. and Passos, F.D. (2015) Spheniopsidae Gardner, 1928 (Bivalvia): conchological characters of two new species from off Brazil, Southwestern Atlantic. American Malacological Bulletin 33, 212220.
Martins, A.M.F., Ávila, S., Borges, J.P., Madeira, P. and Morton, B. (2009) Illustrated checklist of the infralittoral molluscs off Vila Franco do Campo. In Martins, A.M.F. (ed.) The marine fauna and flora of the Azores (Proceedings of the Third International Workshop of Malacology and Marine Biology, São Miguel, Portugal 2006). Açoreana, Supplement 6, pp. 15103.
Massemin, D., Lamy, D., Pointier, J.P. and Gargominy, O. (2009) Coquillages et escargots de Guyane, Seashells and snails from Guiana. Paris: Méze (Collection Parthénope): Muséum national d’ Historie naturelle. [In France]
Migotto, A.E., Tiago, C.G. and Magalhaens, A.R.M. (1993) Malacofauna marinha da região costeira do Canal de São Sebastião, SP, Brasil: Gastropoda, Bivalvia, Polyplacophora e Scaphopoda. Boletim do Instituto Oceanográfico 41, 1327. [In Portuguese]
Mikkelsen, P.M. and Bieler, R. (2008) Seashells of Southern Florida: living marine mollusks of the Florida Keys and adjacent regions: Bivalves. Princeton, NJ: Princeton University Press.
Morgan, R.E. and Allen, J.A. (1976) On the functional morphology and adaptations of Entodesma saxicola (Bivalvia: Anamolodesmacea). Malacologia 15, 233240.
Morton, B. (1973) The biology and functional morphology of Laternula truncata (Lamarck 1818) (Bivalvia: Anomalodesmata: Pandoracea). Biological Bulletin 145, 509531.
Morton, B. (1977) Some aspects of the biology and functional morphology of Myadora striata (Quoy & Gaimard) (Bivalvia: Anomalodesmata: Pandoracea). Journal of Molluscan Studies 43, 141154.
Morton, B. (1980) Anatomy of the “living fossil” Pholadomya candida Sowerby 1823 (Bivalvia: Anomalodesmata: Pholadomyacea). Videnskabelige Meddelelser fra dansk naturhistorisk Forening i Kjøbenhavn 142, 7101.
Morton, B. (1981a) The biology and functional morphology of Periploma (Offadesma) angasai (Bivalvia: Anomalodesmata: Periplomatidae). Journal of Zoology, London 193, 3970.
Morton, B. (1981b) Prey capture in the carnivorous “septibranch” Poromya granulata (Bivalvia: Anomalodesmata: Poromyacea). Sarsia 66, 241256.
Morton, B. (1981c) The Anomalodesmata. Malacologia 21, 3560.
Morton, B. (1982) The functional morphology of Parilimya fragilis (Grieg, 1920) (Bivalvia: Parilimyidae nov. fam.) with a discussion of the origin and evolution of the carnivorous septibranchs and a reclassification of the Anomalodesmata. Transactions of the Zoological Society of London 36, 153216.
Morton, B. (1984a) Prey capture in Lyonsiella formosa (Bivalvia: Anomalodesmata: Verticordiacea). Pacific Science 38, 283297.
Morton, B. (1984b) The biology and functional morphology of Clavagella australis (Bivalvia: Anomalodesmata). Journal of Zoology, London 202, 489511.
Morton, B. (1984c) The adaptations of Frenamya ceylanica (Bivalvia: Anomalodesmata: Pandoracea) to life on the surface of soft muds. Journal of Conchology 31, 359371.
Morton, B. (1985a) Adaptive radiation in the Anomalodesmata. In Wilbur, K.M., Trueman, E.R. and Clarke, M. (eds) The Mollusca. Volume 10. Evolution. New York, NY: Academic Press, pp. 405459.
Morton, B. (1985b) Statocyst structure in the Anomalodesmata (Bivalvia). Journal of Zoology, London 206, 2334.
Morton, B. (1987) Siphon structure and prey capture as a guide to affinities in the abyssal septibranch Anomalodesmata (Bivalvia). Sarsia 72, 4969.
Morton, B. (1995) The ecology and functional morphology of Trigonothracia jinxingae (Bivalvia: Anomalodesmata: Thracioidea) from Xiamen, China. Journal of Zoology, London 237, 445468.
Morton, B. (2002a) The biology and functional morphology of Humphreyia strangei (Bivalvia: Anomalodesmata: Clavagellidae): an Australian cemented ‘watering pot’ shell. Journal of Zoology, London 258, 1125.
Morton, B. (2002b) Biology and functional morphology of the watering pot shell Brechites vaginiferus (Bivalvia: Anomalodesmata: Clavagelloidea). Journal of Zoology, London 257, 545562.
Morton, B. (2003) The functional morphology of Bentholyonsia teramachii (Bivalvia: Lyonsiellidae): clues to the origin of predation in the deep water Anomalodesmata. Journal of Zoology, London 261, 363380.
Morton, B. (2004) The biology and functional morphology of Foegia novaezelandiae (Bivalvia: Anomalodesmata: Clavagelloidea) from Western Australia. Malacologia 46, 3755.
Morton, B. (2005) Biology and functional morphology of a new species of endolithic Bryopa (Bivalvia: Anomalodesmata: Clavagelloidea) from Japan and a comparison with fossil species of Stirpulina and other Clavagellidae. Invertebrate Biology 124, 202219.
Morton, B. (2006) The functional morphology of Penicillus philippinensis (Anomalodesmata: Clavagelloidea: Penicillidae) and the evolution of an unique muscular system in the Bivalvia. Records of the Western Australian Museum 23, 175192.
Morton, B. (2012) The functional morphology and inferred biology of the enigmatic South African ‘quadrivalve’ bivalve Clistoconcha insignis Smith, 1910 (Thracioidea: Clistoconchidae fam. nov.): another anomalodesmatan evolutionary eccentric. Transactions of the Royal Society of South Africa 67, 5989.
Morton, B. (2015) The biology and functional morphology of the predatory septibranch Cardiomya costellata (Deshayes, 1833) (Bivalvia: Anomalodesmata: Cuspidariidae) from the Açores: survival at the edge. Journal of the Marine Biological Association of the United Kingdom. doi: 10.1017/S0025315415001848.
Morton, B., Dinesen, E.G. and Ockelmann, W.K. (2015a) Functional morphology, biology and sexual strategy of the circumboreal, adventitious crypt-building, Crenella decussata (Bivalvia: Mytiloidea: Crenellidae). Journal of the Marine Biological Association of the United Kingdom. doi: 10.1017/S0025315415001927.
Morton, B. and Dinesen, G.E. (2011) The biology and functional morphology of Modiolarca subpicta (Bivalvia: Mytilidae: Musculinae), epizoically symbiotic with Ascidiella aspersa (Urochordata: Ascidiacea), from the Kattegat, northern Jutland, Denmark. Journal of the Marine Biological Association of the United Kingdom 91, 16371649.
Morton, B., Machado, F.M. and Passos, F.D. (2015b) The smallest carnivorous bivalve? Biology, morphology and behaviour of Grippina coronata (Anomalodesmata: Cuspidarioidea: Spheniopsidae) predating epipsammic microcrustaceans in the southwestern Atlantic off Brazil. Journal of Molluscan Studies. doi: 10.1093/mollus/eyv055.
Morton, B., Machado, F.M. and Passos, F.D. (2015c) The organs of prey capture and digestion in the miniature predatory bivalve Spheniopsis brasiliensis (Anomalodesmata: Cuspidarioidea: Spheniopsidae) expose a novel life history trait. Journal of Natural History. doi: 10.1080/00222933.2016.1155780.
Narchi, W. (1968) The functional morphology of Lyonsia californica Conrad, 1837 (Bivalvia). The Veliger 10, 305313.
Narchi, W. and Assis, R.C.F. (1980) Anatomia funcional de Lucina pectinata (Gmelin, 1791) (Bivalvia-Lucinidae). Boletim de Zoologia da Universidade de São Paulo, São Paulo 5, 79110. [In Portuguese]
Oliveira, C.D.C. and Absalão, R.S. (2009) The genera Myonera, Octoporia, and Protocuspidaria (Pelecypoda, Cuspidariidae) from deep waters of Campos Basin, Rio de Janeiro, Brazil with descriptions of two new species. American Malacological Bulletin 27, 141156.
Oliveira, C.D.C. and Sartori, A.F. (2013) Discovery and anatomy of the arenophilic system of cuspidariid clams (Bivalvia: Anomalodesmata). Journal of Morphology 275, 916.
Passos, F.D., Meserani, G.L.C. and Gros, O. (2007) Structural and ultrastructural analysis of the gills in the bacterial-bearing species Thyasira falklandica (Bivalvia, Mollusca). Zoomorphology 126, 153162.
Payne, C.M. and Allen, J.A. (1991) The morphology of deep sea Tyasiridae (Mollusca: Bivalvia) from the Atlantic Ocean. Philosophical Transactions of the Royal Society, Series B 334, 481562.
Poutiers, J.M. and Bernard, F.R. (1995) Carnivorous bivalve molluscs (Anomalodesmata) from the tropical western Pacific Ocean, with a proposed classification and a catalogue of Recent species. In Bouchet, P. (ed.) Résultats des Campagnes MUSORSTOM. Volume 14. Mémoires Muséum national Histoire naturelle 167, 107187.
Purchon, R.D. (1955) A note on the biology of Brechites penis (L.) (Lamellibranchia). Zoological Journal of the Linnean Society 43, 4354.
Purchon, R.D. (1956) The stomach in the Protobranchia and Septibranchia (Lamellibranchia). Proceedings of the Zoological Society of London 127, 511525.
Purchon, R.D. (1960) A further note on the biology of Brechites penis (L.) (Lamellibranchia). Proceedings of the Malacological Society of London 34, 1923.
Reid, R.G.B. and Crosby, S.P. (1980) The raptorial siphonal apparatus of the carnivorous septibranch Cardiomya planetica Dall (Mollusca: Bivalvia), with notes on feeding and digestion. Canadian Journal of Zoology 58, 670679.
Reid, R.G.B. and Reid, A.M. (1974) The carnivorous habit of members of the septibranch genus Cuspidaria (Mollusca: Bivalvia). Sarsia 56, 4756.
Rios, E.C. (1994) Seashells of Brazil, 2nd edition. Rio Grande: Museu Oceanográfico Prof. E. C. Rios da Fundação Universidade do Rio Grande.
Rios, E.C. (2009) Compendium of Brazilian sea shells, 1st edition. Rio Grande: Evangraf.
Runnegar, B. (1974) Evolutionary history of the bivalve subclass Anomalodesmata. Journal of Paleontology 48, 904939.
Sartori, A.F. and Domaneschi, O. (2005) The functional morphology of the antarctic bivalve Thracia meridionalis Smith, 1885 (Anomalodesmata: Thraciidae). Journal of Molluscan Studies 71, 199210.
Scarabino, F. (2003) Lista sistemática de los Bivalvia marinos y estuarinos vivientes de Uruguay. Comunicaciones de la Sociedad Malacológica del Uruguay 8, 229259. [In Spanish]
Scarlato, O.A. and Starogobatov, Ya.I. (1983) System of the bivalve molluscs Lamelibranchia of the superorder Septibranchia. Seventeen Meeting on the Investigation of Molluscs, Leningrad, “Nauka”, 7–14. [In Russian]
Simone, L. and Cunha, C.M. (2008) Revision of the genus Spinosipella (Bivalvia: Verticordiidae), with descriptions of two new species from Brazil. The Nautilus 122, 5778.
Temkin, I. and Strong, E.E. (2013) New insights on stomach anatomy of carnivorous bivalves. Journal of Molluscan Studies 79, 332339.
Thomas, K.A. (1994) The functional morphology and biology of Pandora filosa (Carpenter, 1864) (Bivalvia: Anomalodesmata: Pandoracea). The Veliger 37, 2329.
Taylor, J.D. and Glover, E.A. (2006) Lucinidae (Bivalvia) – the most diverse group of chemosymbiotic molluscs. Zoological Journal of the Linnean Society 148, 421438.
Yonge, C.M. (1928) Structure and function of the organs of feeding and digestion in the septibranchs, Cuspidaria and Poromya . Philosophical Transactions of the Royal Society, Series B 216, 221263.
Yonge, C.M. (1948) Formation of siphons in Lamellibranchia. Proceedings of the United States National Museum 20, 775901.
Yonge, C.M. (1952) Structure and adaptation in Entodesma saxicola (Baird) and Mytilimeria nuttallii Conrad. Volume 55. Los Angeles, CA: University of California Press, pp. 439450.
Yonge, C.M. (1957) Mantle fusion in the Lamellibranchia. Pubblicazioni della Stazione Zoologica di Napoli 29, 151171.
Yonge, C.M. (1982) Mantle margins with a revision of siphonal types in the Bivalvia. Journal of Molluscan Studies 48, 102103.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

Journal of the Marine Biological Association of the United Kingdom
  • ISSN: 0025-3154
  • EISSN: 1469-7769
  • URL: /core/journals/journal-of-the-marine-biological-association-of-the-united-kingdom
Please enter your name
Please enter a valid email address
Who would you like to send this to? *


Type Description Title
Supplementary materials

Machado supplementary material

 Video (25.1 MB)
25.1 MB


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed