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 and a unique feeding structure called the pseudoproboscis. The pseudoproboscis differs from a true gastropod proboscis in the presence of a slit along the lower side. The shell bears a thick (often spiky) periostracum. 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 [Capulacea, Capulusidae]
    |  i. s.: Piliscus Loven 1856 O27
    |           `--P. commodus Middendorff 1851 O27 [=Pilidium commodum J64; incl. Capulus fallax J64, C. obliquus J64]
    |         Neojanacus Suter 1907 P61
    |           `--*N. perplexus Suter 1907 P61
    |--Lysis Gabb 1864 [Lysinae] BR17
    |    `--*L. duplicosta Gabb 1864 BR17
    `--Capulinae [Trichotropidae, Trichotropinae] BR17
         |--Lepistes P08
         |--Turbinopsis Conrad 1860 TTE93
         |--‘Verena’ Gray 1857 non Adams & Adams 1854 BR17, BR05 [Verenadae, Verenidae]
         |    `--*V. borealis (Broderip & Sowerby 1829) [=Trichotropis borealis] BR17
         |--Sirius Hedley 1900 BR17, BR05 [Siriidae]
         |    `--*S. badius (Tenison-Woods 1876) [=Raulinia badia] BR17
         |--Cerithioderma Conrad 1860 BR17, BR05 [Cerithiodermatidae, Cerithiodermidae]
         |    `--*C. prima Conrad 1860 BR17
         |--Lirpsa Stephenson 1952 BD12
         |    |--*L. cornuta Stephenson 1952 BD12
         |    `--L. garudamangalami Bandel 2000 BD12
         |--Astandes Wade 1917 BD12
         |    |--*A. dentatus Wade 1917 BD12
         |    `--A. nodosus (Dockery 1993) [=Cerithioderma nodosa] BD12
         |--Atresius TTE93
         |    |--A. cornuelianus (Orbigny 1843) TTE93
         |    `--A. lallierianus (Orbigny 1843) TTE93
         |--Zelippistes Finlay 1927 P61
         |    |--*Z. benhami (Suter 1902) [=Separatista benhami] P61
         |    |--Z. pehuensis (Marwick 1926) [=Lippistes pehuensis] F27a
         |    `--Z. perornatus (Marshall & Murdoch 1923) [=Lippistes perornatus] F27a
         |--Torellia Jeffreys 1867 PH90 [incl. Trachysma Sars 1878 BR17; Trachysmatidae]
         |    |--*T. vestita Jeffreys 1867 PH90
         |    |--T. ammonia Dall 1919 O27
         |    |--T. delicata (Philippi 1844) (see below for synonymy) BR17
         |    `--T. vallonia Dall 1919 O27
         |--Lippistes Montfort 1810 BR17, BR05 [Lippistidae]
         |    |--*L. cornu (Gmelin 1791) [=Argonauta cornu] BR17
         |    |--L. blainvilleanus (Petit 1851) [=Trichotropis blainvilleana] H09
         |    |--L. gracilentus (Brazier 1877) [=Trichotropis gracilenta] H09
         |    `--L. zodiacus Hedley 1907 H09
         |--Trichosirius Finlay 1927 P61
         |    |--*T. inornatus (Hutton 1873) [=Trichotropis inornata] P61
         |    |    |--T. i. inornatus P61
         |    |    `--T. i. chathamensis Finlay 1928 P61
         |    |--T. cavatocarinatus (Laws 1940) P61
         |    `--T. octocarinatus (Powell 1931) P61
         |--Trichotropis Broderip & Sowerby 1829 BR05
         |    |  i. s.: T. acuminatus N79
         |    |         T. cancellata Hinds 1843 S11
         |    |         T. carinata [=Murex carinatus; incl. T. atlantica, Fusus umbilicatus] J64
         |    |         T. ciliata C64
         |    |         T. conica Möller 1842 O27
         |    |         T. costellata Couthouy 1838 O27
         |    |         T. inermis C64
         |    |         T. insignis Middendorff 1849 O27
         |    |         T. mississippiensis Sohl 1960 BD12
         |    |--*T. (Trichotropis) bicarinatus (Sowerby 1825) BR17 [=Turbo bicarinatus BR17; incl. Tr. sowerbiensis C64]
         |    `--T. (Iphinoe Adams & Adams 1854) O27
         |         `--T. (I.) coronata Gould 1860 O27 [incl. T. multicaudata C64]
         `--Capulus Montfort 1810 [=Pileopsis Lamarck 1822; Pileopsidae] BR05
              |--*C. ungaricus (Linnaeus 1758) P61, L58 [=Patella ungarica P61, Pileopsis ungaricus J64]
              |--C. ‘angustus’ Wanner 1922 non Pileopsis angustus Philipps 1836 F27b
              |--‘Pileopsis’ angustus Philipps 1836 F27b
              |--C. australis H08
              |--C. calcareus Suter 1909 P61
              |--C. californicus Dall 1900 O27
              |--C. calyptra (Martyn 1784) [=Patella calyptra] H09
              |--C. devotus M54
              |--C. nutatus Hedley 1908 H08
              |--C. sycophanta Garrard 1961 S11
              |--C. tenellus (Dana 1849) [=Pileopsis tenella] W63
              `--C. verus (Böhm 1885) TTE93

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.

[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.

[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.

[M54] Macpherson, J. H. 1954. Molluscs (sea shells and snails). In: Willis, J. M. (ed.) The Archipelago of the Recherche pp. 55–63. Australian Geographical Society: Melbourne.

[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).

[P08] Parkhaev, P. Y. 2008. The Early Cambrian radiation of Mollusca. In: Ponder, W. F., & D. R. Lindberg (eds) Phylogeny and Evolution of the Mollusca pp. 33–69. University of California Press: Berkeley.

[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.

Last updated: 16 September 2021.

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