Trichotropis bicarinatus, photographed by Mingzhu Gao.

Belongs within: Hypsogastropoda.

The Capulidae are a morphologically diverse group of gastropods sharing a distinctive larval form called the echinospira in which a second larval shell is present outside what eventually becomes the adult shell. The adult shell bears a thick (often spiky) periostracum. A unique feeding structure called the pseudoproboscis is present that differs from a true gastropod proboscis in the presence of a slit along the lower side.

Some authors have divided this group between the limpet-shaped Capulidae sensu stricto and the high-spired and coiled Trichotropidae. The Late Cretaceous Lysinae are low turbiniform to almost haliotiform shells with the columella and inner lip flattened and expanded to form a shelf inside the shell aperture. Within the modern Capulinae, members of the genus Capulus have a conical shell with a spirally recurved apex (Oldroyd 1927).

<==Capulidae [Capulusidae]
| i. s.: Piliscus Loven 1856O27
| `--P. commodus Middendorff 1851O27 [=Pilidium commodumJ64; incl. Capulus fallaxJ64, C. obliquusJ64]
| Neojanacus Suter 1907P61
| `--*N. perplexus Suter 1907P61
| Icuncula Iredale 1924W93
| |--*I. torcularis (Tenison Woods 1878) [=Lippistes torcularis, Cingulina torcularis]W93
| |--I. consobrina (May 1915)W93
| `--I. zodiacus Hedley 1907W93
|--Lysis Gabb 1864 [Lysinae]BR17
| `--*L. duplicosta Gabb 1864BR17
`--Capulinae [Trichotropidae, Trichotropinae]BR17
|--Turbinopsis Conrad 1860TTE93
|--‘Verena’ Gray 1857 non Adams & Adams 1854BR17, BR05 [Verenadae, Verenidae]
| `--*V. borealis (Broderip & Sowerby 1829) [=Trichotropis borealis]BR17
|--Cerithioderma Conrad 1860BR17, BR05 [Cerithiodermatidae, Cerithiodermidae]
| `--*C. prima Conrad 1860BR17
|--Lirpsa Stephenson 1952BD12
| |--*L. cornuta Stephenson 1952BD12
| `--L. garudamangalami Bandel 2000BD12
|--Astandes Wade 1917BD12
| |--*A. dentatus Wade 1917BD12
| `--A. nodosus (Dockery 1993) [=Cerithioderma nodosa]BD12
| |--A. cornuelianus (Orbigny 1843)TTE93
| `--A. lallierianus (Orbigny 1843)TTE93
|--Zelippistes Finlay 1926P61, W93
| |--*Z. benhami (Suter 1902) [=Separatista benhami]P61
| |--Z. pehuensis (Marwick 1926) [=Lippistes pehuensis]F27a
| `--Z. perornatus (Marshall & Murdoch 1923) [=Lippistes perornatus]F27a
|--Torellia Jeffreys 1867PH90 [incl. Trachysma Sars 1878BR17; Trachysmatidae]
| |--*T. vestita Jeffreys 1867PH90
| |--T. ammonia Dall 1919O27
| |--T. delicata (Philippi 1844) (see below for synonymy)BR17
| `--T. vallonia Dall 1919O27
|--Sirius Hedley 1900BR17, BR05 [incl. DolichosiriusW93, OpposiriusW93; Siriidae]
| |--*S. badius (Tenison-Woods 1876) [=Raulinia badia]BR17
| |--S. cupiens Iredale 1931 [=S. (*Dolichosirius) cupiens]W93
| |--S. desponsus Iredale 1936W93
| |--S. idoneus (Iredale 1931) [=S. (*Opposirius) idoneus]W93
| `--S. meracus Iredale 1936W93
|--Trichosirius Finlay 1927P61
| |--*T. inornatus (Hutton 1873) [=Trichotropis inornata]P61
| | |--T. i. inornatusP61
| | `--T. i. chathamensis Finlay 1928P61
| |--T. cavatocarinatus (Laws 1940)P61 [=Cerithioderma cavatocarinatusD56]
| `--T. octocarinatus (Powell 1931)P61
|--Lippistes Montfort 1810BR17, BR05 [incl. Separatista Gray 1847W93; Lippistidae]
| |--*L. cornu (Gmelin 1791) [=Argonauta cornu]BR17
| |--L. blainvilleanus (Petit 1851) [=Trichotropis blainvilleana]H09
| |--L. gabrieli Pritchard & Gatliff 1898 [incl. L. meridionalis Verco 1906]W93
| |--L. gracilenta Brazier 1877W93 [=Trichotropis gracilentaH09]
| |--L. helicoides (Gmelin 1791) (see below for synonymy)W93
| `--L. zodiacus Hedley 1907H09
|--Trichotropis Broderip & Sowerby 1829BR05
| | i. s.: T. acuminatusN79
| | T. cancellata Hinds 1843S11
| | T. carinata [=Murex carinatus; incl. T. atlantica, Fusus umbilicatus]J64
| | T. ciliataC64
| | T. conica Möller 1842O27
| | T. costellata Couthouy 1838O27
| | T. flavida Hinds 1843H43
| | T. inermis Hinds 1843H43
| | T. insignis Middendorff 1849O27
| | T. mississippiensis Sohl 1960BD12
| |--*T. (Trichotropis) bicarinatus (Sowerby 1825)BR17 [=Turbo bicarinatusBR17; incl. Tr. sowerbiensisC64]
| `--T. (Iphinoe Adams & Adams 1854)O27
| `--T. (I.) coronata Gould 1860O27 [incl. T. multicaudataC64]
`--Capulus Montfort 1810BR05 (see below for synonymy)
|--*C. ungaricus (Linnaeus 1758)P61, L58 [=Patella ungaricaP61, Pileopsis ungaricusJ64]
|--C. ‘angustus’ Wanner 1922 non Pileopsis angustus Philipps 1836F27b
|--‘Pileopsis’ angustus Philipps 1836F27b
|--C. australisH08
|--C. calcareus Suter 1909P61
|--C. californicus Dall 1900O27
|--C. danieli (Crosse 1858)W93 (see below for synonymy)
|--C. devexus May 1916W93
|--C. devotus Hedley 1904 [incl. C. banksi Cotton 1939]W93
|--C. dilatatusW93
|--C. liberatus Pease 1868 [=*Tenpetasus liberatus]W93
|--C. nutatus Hedley 1908H08
|--C. sycophanta Garrard 1961S11
|--C. tenellus (Dana 1849) [=Pileopsis tenella]W63
|--C. verus (Böhm 1885)TTE93
`--C. violaceus Angas 1867W93

Capulus Montfort 1810BR05 [=Pileopsis Lamarck 1822BR05; incl. Capulonix Iredale 1929W93, Tenpetasus Iredale 1929W93; Pileopsidae]

Capulus danieli (Crosse 1858)W93 [=Capulonix danieliW93; incl. Patella calyptra Martyn 1784 (nom. inv.)H09, *Capulonix calyptraW93, Capulus calyptraH09]

Lippistes helicoides (Gmelin 1791) [incl. L. fraterna Iredale 1936, *Separatista separatista Dillwyn 1817, L. tricarinata Brazier 1877]W93

Torellia delicata (Philippi 1844) [=Cyclostoma delicatum, *Trachysma delicatum; incl. Tr. sarsianum Thiele 1910]BR17

*Type species of generic name indicated


[BD12] Bandel, K., & D. T. Dockery III. 2012. Protoconch characters of Late Cretaceous Latrogastropoda (Neogastropoda and Neomesogastropoda) as an aid in the reconstruction of the phylogeny of the Neogastropoda. Freiberger Forschungshefte C542 (psf 20): 93–128.

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[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.

[D56] Dell, R. K. 1956. The archibenthal Mollusca of New Zealand. Dominion Museum Bulletin 18: 1–235.

[F27a] Finlay, H. J. 1927a. A further commentary on New Zealand molluscan systematics. Transactions and Proceedings of the New Zealand Institute 57: 320–485.

[F27b] Finlay, H. J. 1927b. New specific names for austral Mollusca. Transactions and Proceedings of the New Zealand Institute 57: 488–533.

[H08] Hedley, C. 1908. Studies on Australian Mollusca. Part X. Proceedings of the Linnean Society of New South Wales 33: 456–489, pls 7–10.

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

[H43] Hinds, R. B. 1843. Descriptions of new shells from the collection of Captain Belcher, R.N., C.B., &c. Annals and Magazine of Natural History 11: 16–21.

[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.

[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.

[O27] Oldroyd, I. S. 1927. The Marine Shells of the West Coast of North America vol. 2 pt 3. Stanford University Press: Stanford University (California).

[PH90] Petit, R. E., & M. G. Harasewych. 1990. Catalogue of the superfamily Cancellarioidea Forbes and Hanley, 1851 (Gastropoda: Prosobranchia). Nautilus 103 (Suppl. 1): 1–69.

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

[S11] Simone, L. R. L. 2011. Phylogeny of the Caenogastropoda (Mollusca), based on comparative morphology. Arquivos de Zoologia 42 (4): 161–323.

[TTE93] Tracey, S., J. A. Todd & D. H. Erwin. 1993. Mollusca: Gastropoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 131–167. Chapman & Hall: London.

[W63] Waterhouse, J. B. 1963. Permian gastropods of New Zealand. Part 4—Platyceratacea, Anomphalacea, Neritacea, and correlations. New Zealand Journal of Geology and Geophysics 6 (5): 817–842.

[W93] Wilson, B. 1993. Australian Marine Shells vol. 1. Prosobranch Gastropods. Part One. Odyssey Publishing: Kallaroo (Western Australia).


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