Pholadoidea


Drawing of a shipworm Teredo without its burrow, from here.


Belongs within: Neoheterodontei.
Contains: Pholas.

The Pholadoidea are a clade of two families of bivalves, Pholadidae and Teredinidae, which are adapted for boring into hard substrates (rock or wood in the case of Pholadidae, wood for Teredinidae) using their shells to grind out a burrow. Teredinids, commonly known as shipworms, are particularly derived, with the shells reduced to cover only the tip of the long worm-like animal. A calcareous lining is usually secreted to cover the living tube. The Pholadidae have a more complete shell which nevertheless bears a distinct pedal gape (though this may be secondarily closed by a callum). Fossils of Pholadidae are known from the mid-Jurassic onwards with fossil Teredinidae recorded from the mid-Cretaceous (Skelton & Benton 1993).

Characters (from Cox et al. 1969): Closed mantle and truncate, more or less circular foot developed as suction disc. Shells inequilateral, dorsal margin reflected anterior to beak, forming attachment area for anterior adductor muscle which works counter to posterior adductor when animal is boring. Hinge teeth lacking, but small chondrophore and internal ligament usually present. Ventral adductor muscle or at least thickening of muscles of ventral margin of mantle located at union of anterior slope and disc, forming pivotal point opposite umbones on which valves rock when boring. Gills usually elongate, with two demibranchs except in Xylophagainae and Teredinidae. Apophyses found in all Pholadoidea except in some Pholadidae.

Pholadoidea [Pholadacea]
    |  i. s.: Nototeredo C96
    |         Xylophaga dorsalis C96, PP64
    |         Dactylina C64
    |           |--D. campechensis [incl. Pholas oblongata] C64
    |           |--D. (Gitocentrum) chiloensis King 1832 [incl. Pholas laqueata Sby. 1849, D. laquiata] C64
    |           `--D. dactylus C64
    |         Navea subglobosa Gray 1851 C64
    |--Teredinidae GW02
    |    |--Teredora malleolus (Turton 1822) [incl. Teredo thomsonii] FH89
    |    |--Lyrodus pedicellatus (Quatrefages 1849) FH89
    |    |--Psiloteredo healdi G88
    |    |--Terebrimya SB93
    |    |--Xylotrya C64
    |    |    |--X. fimbriata Jeffr. 1860 [incl. X. palmulata] C64
    |    |    `--X. pennatifera Blainv. 1860 C64
    |    |--Bankia Gray 1842 P61
    |    |    |--*B. bipalmulata [=Teredo bipalmulata] P61
    |    |    |--B. australis Calman 1920 P61
    |    |    |--B. carinata GW02
    |    |    |--B. gouldi (Bartsch 1908) FH89
    |    |    |--B. martensi (Stempell 1899) FH89
    |    |    `--B. setacea (Tryon 1863) FH89
    |    |--Teredo Linnaeus 1758 P61
    |    |    |--*T. navalis Linnaeus 1758 P61
    |    |    |--T. antarctica Hutton 1873 P61
    |    |    |--T. bipennata N79
    |    |    |--T. heaphyi [incl. Ciadopoda directa] H86
    |    |    |--T. lapidaria Linnaeus 1758 L58
    |    |    |--T. megotara N79
    |    |    |--T. nana [incl. T. denticulata, T. dilatata, Pholas teredo] J64
    |    |    |--T. norvegica Spengler 1792 [incl. T. bruguieri Delle Chiaje 1828] F26
    |    |    |--T. poculifer RD73
    |    |    `--T. vastisas Etheridge 1902 F71
    |    `--Nausitora RD73
    |         |--N. dunlopei Wright 1864 (see below for synonymy) RD73
    |         |--N. hedleyi RD73
    |         `--N. lanceolata Rajagopal 1964 RD73
    `--Pholadidae GW02
         |--Aspidopholas JB12
         |--Turnus SB93
         |--Netastoma darwinii [incl. Pholas rostrata] C64
         |--Zirphaea crispata C64 [=Pholas crispata K91]
         |--Pholas L58
         |--Anchomasa Leach 1852 P61
         |    |--*A. parva [=Pholas parva] P61
         |    `--A. similis (Gray 1835) P61
         |--Jouannetia Des Moulins 1828 JB12, H75
         |    |--J. cumingi Sowerby 1849 H09
         |    |--J. globosa Q. & G. 1832 H09
         |    `--J. (Pholadopsis) pectinata [incl. Triomphalia pulcherrima] C64
         |--Parapholas C64
         |    |--P. bisulcata C64
         |    |--P. californica [=Pholas californica; incl. Ph. janellii] C64
         |    `--P. calva [incl. P. acuminata] C64
         |--Penitella JB12
         |    |--P. conradi [incl. P. tubigera] C64
         |    |--P. dolichothyra Tchang, Tsi & Li 1960 XZ10
         |    |--P. spelaea Conrad 1855 C64
         |    `--P. xilophaga C64
         |--Martesia G88
         |    |--M. (Particoma) australis Moore 1870 SB93
         |    |--M. intercalata C64
         |    |--M. obtecta (Sowerby 1849) [=Pholas obtecta] H09
         |    |--M. rivicola G88
         |    `--M. striata (Linnaeus 1758) [=Pholas striatus] H09
         |--Barnea TW07
         |    |--B. candida (Linnaeus 1758) TW07
         |    |--B. davidi (Deshayes 1874) XZ10
         |    |--B. fragilis (Sowerbyy 1844) XZ10
         |    |--B. parva (Pennant 1777) TW07
         |    `--B. similis HS01
         `--Pholadidea Goodall in Turton 1819 P61
              |--*P. loscombiana P61
              |--P. (Hatasia) melanura [=Penitella melanura; incl. Pe. wilsoni] C64
              |--P. ovoidea [=Penitella ovoidea] C64
              |--P. penita [=Parapholas penita, Penitella penita, Pholas penita; incl. Pholas concamerata] C64
              |--P. spathulata (Sowerby 1850) P61
              |--P. suteri HS01
              `--P. tridens (Gray 1843) P61

Nausitora dunlopei Wright 1864 [incl. Calobates fluviatilis Hedley 1898, Bankia globosa Sivickis 1928, Nausitora madagassica Roch 1935, B. (Nausitora) madrasensis Nair 1954, N. messeli Iredale 1932, B. pennamseris Roch 1935, B. quadrangularis Sivickis 1928, Nausitora queenslandica Iredale 1936, N. schneideri Moll 1935, Bankia (Nausitora) smithi Bartsch 1927, B. triangularis Sivickis 1928] RD73

*Type species of generic name indicated

REFERENCES

[C64] Carpenter, P. P. 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Report of the British Association for the Advancement of Science 33: 517–686.

[C96] Cragg, S. M. 1996. The phylogenetic significance of some anatomical features of bivalve veliger larvae. In: Taylor, J. D. (ed.) Origin and Evolutionary Radiation of the Mollusca pp. 371–380. Oxford University Press: Oxford.

[F26] Finlay, H. J. 1926. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[FH89] Fuller, S. C., Y.-P. Hu, R. A. Lutz & M. Castagna. 1989. Shell and pallet morphology in early developmental stages of Teredo navalis Linn√© (Bivalvia: Teredinidae). Nautilus 103 (1): 24–35.

[GW02] Giribet, G., & W. Wheeler. 2002. On bivalve phylogeny: a high-level analysis of the Bivalvia (Mollusca) based on combined morphology and DNA sequence data. Invertebrate Biology 121 (4): 271–324.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H75] Häntzschel, W. 1975. Treatise on Invertebrate Paleontology pt W. Miscellanea Suppl. 1. Trace Fossils and Problematica 2nd ed. The Geological Society of America: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[HS01] Hayward, B. W., A. B. Stephenson, M. S. Morley, W. M. Blom, H. R. Grenfell, F. J. Brook, J. L. Riley, F. Thompson & J. J. Hayward. 2001. Marine biota of Parengarenga Harbour, Northland, New Zealand. Records of the Auckland Museum 37: 45–80.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[H86] Hutton, F. W. 1886. The Mollusca of the Pareora and Oamaru systems of New Zealand. Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 205–237.

[J64] Jeffreys, J. G. 1864. The Upper Tertiary fossils at Uddevalla, in Sweden. Report of the British Association for the Advancement of Science 33 (Notices and Abstracts): 73–79.

[JB12] Johnson, M. E., & B. G. Baarli. 2012. Development of intertidal biotas through Phanerozoic time. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 63–128. Springer.

[K91] Kozloff, E. N. 1991. Malacobdella siliquae sp. nov. and Malacobdella macomae sp. nov., commensal nemerteans from bivalve molluscs on the Pacific coast of North America. Canadian Journal of Zoology 69: 1612–1618.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[N79] Norman, A. M. 1879. The Mollusca of the fiords near Bergen, Norway. Journal of Conchology 2: 8–77.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d'Endoume, Bulletin 31 (27): 5–137.

[P61] Powell, A. W. B. 1961. Shells of New Zealand: An illustrated handbook 4th ed. Whitcombe and Tombs Limited: Christchurch.

[RD73] Rajagopal, A. S., & A. Daniel. 1973. Boring organisms of the Great Nicobar Island. Mollusca: Teredinidae. Journal of the Bombay Natural History Society 69 (3): 676–678.

[SB93] Skelton, P. W., & M. J. Benton. 1993. Mollusca: Rostroconchia, Scaphopoda and Bivalvia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 237–263. Chapman & Hall: London.

[TW07] Taylor, J. D., S. T. Williams, E. A. Glover & P. Dyal. 2007. A molecular phylogeny of heterodont bivalves (Mollusca: Bivalvia: Heterodonta): new analyses of 18S and 28S rRNA genes. Zoologica Scripta 36 (6): 587–606.

[XZ10] Xu, F., & J. Zhang. 2010. Study on the bivalve faunal composition of the Yellow Sea and Bohai Sea. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 7–12. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

Last updated: 5 March 2022.

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