Belongs within: Neoheterodontei.
Contains: Teredininae, Martesiinae, 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).
Most Pholadidae are filter-feeders with the exception of the deep-sea wood-boring Xylophagainae (Beesley et al. 1998).
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.: DactylinaC64
| |--D. campechensis [incl. Pholas oblongata]C64
| |--D. (Gitocentrum) chiloensis King 1832 [incl. Pholas laqueata Sby. 1849, D. laquiata]C64
| `--D. dactylusC64
| Navea subglobosa Gray 1851C64
|--TeredinidaeGW02
| | i. s.: Psiloteredo healdiG88
| | TerebrimyaSB93
| | XylotryaC64
| | |--X. fimbriata Jeffr. 1860 [incl. X. palmulata]C64
| | `--X. pennatifera Blainv. 1860C64
| | Zachsia zenkewitschiBRW98
| |--TeredininaeBRW98
| |--Kuphus [Kuphinae]BRW98
| | |--K. dubia Sivickis 1928BRW98
| | `--K. polythalamia [incl. K. arenarius]BRW98
| `--BankiinaeBRW98
| |--Nototeredo edaxBRW98
| |--Spathoteredo obtusaBRW98
| |--NausitoraBRW98
| | |--N. dunlopei Wright 1864 (see below for synonymy)RD73
| | |--N. hedleyiRD73
| | `--N. lanceolata Rajagopal 1964RD73
| `--Bankia Gray 1842P61
| |--*B. bipalmulata [=Teredo bipalmulata]P61
| |--B. australis Calman 1920P61
| |--B. barthelowiBRW98
| |--B. campanellataBRW98
| |--B. carinataGW02
| |--B. gouldi (Bartsch 1908)FH89
| |--B. gracilisBRW98
| |--B. johnsoniBRW98
| |--B. martensi (Stempell 1899)FH89
| |--B. nestaliaBRW98
| |--B. nordiBRW98
| |--B. rochiBRW98
| `--B. setacea (Tryon 1863)FH89
`--Pholadidae [Pholadaria]GW02
| i. s.: TurnusSB93
| Netastoma darwinii [incl. Pholas rostrata]C64
| Zirphaea crispataC64 [=Pholas crispataK91]
| Anchomasa Leach 1852P61
| |--*A. parva [=Pholas parva]P61
| `--A. similis (Gray 1835)P61
| PenitellaJB12
| |--P. conradi [incl. P. tubigera]C64
| |--P. dolichothyra Tchang, Tsi & Li 1960XZ10
| |--P. spelaea Conrad 1855C64
| `--P. xilophagaC64
| Pholadidea Goodall in Turton 1819P61
| |--*P. loscombianaP61
| |--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. suteriHS01
| `--P. tridens (Gray 1843)P61
|--MartesiinaeBRW98
|--XylophagainaeBRW98
| |--Xylophaga dorsalisPP64
| `--XyloredoBRW98
|--Jouannetia Des Moulins 1828JB12, H75 [Jouannetiinae]
| |--J. cumingi Sowerby 1849H09
| |--J. globosa Q. & G. 1832H09
| `--J. (Pholadopsis) pectinata [incl. Triomphalia pulcherrima]C64
`--PholadinaeBRW98
|--PholasL58
`--BarneaBRW98
|--B. australasiaeBRW98
|--B. candida (Linnaeus 1758)TW07
|--B. davidi (Deshayes 1874)XZ10
|--B. dilatataBRW98
|--B. fragilis (Sowerbyy 1844)XZ10
|--B. obturamentumBRW98
|--B. parva (Pennant 1777)TW07
`--B. similisHS01
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
[BRW98] Beesley, P. L., G. J. B. Ross & A. Wells (eds) 1998. Fauna of Australia vol. 5. Mollusca: The Southern Synthesis. Part A. Australian Biological Resources Study: Canberra.
[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.
Cox, L. R., N. D. Newell et al. 1969. Treatise on Invertebrate Paleontology pt N. Bivalvia vol. 2. The Geological Society of America, Inc. and The University of Kansas.
[FH89] Fuller, S. C., Y.-P. Hu, R. A. Lutz & M. Castagna. 1989. Shell and pallet morphology in early developmental stages of Teredo navalis Linné (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.
[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.
[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.