Galeodea echinophora, copyright H. Zell.

Belongs within: Tonnoidea.
Contains: Cassis, Phalium, Semicassis.

The Cassidae, helmet shells, are medium-sized to large marine gastropods found burrowing in sand. They are predatory on echinoids which they attack by squirting a paralysing salivary juice. Females lay large egg-masses containing hundreds of gelatinous capsules; in some cases large irregular masses may be produced seemingly by several females spawning together (Wilson 1993).

Within the Cassidae, the subfamily Phaliinae is distinguished by possession of a slender, fan-shaped operculum with the nucleus halfway down the columellar edge, and a strongly twisted siphonal canal. Cassinae have an oval operculum with the nucleus towards the anterior end, and the siphonal canal is straight or only weakly twisted (Wilson 1993).

Characters (from Wilson & Gillett 1971): Shell usually oval or globose, aperture rather wide and long; shell may be glossy-smooth or heavily sculptured with ribs and nodules. Outer lip thickened to form a varix, with inner edge often toothed; other varices representing outer lip of earlier growth stages may be present. Left side of aperture with flat or inclined shield-like callus, either confined to columellar area (as columellar shield), or covering both columella and parietal wall and forming flange along whole left side. Spiral channel present running under columellar shield into true umbilicus; second smaller cavity (false umbilicus) often present under columellar shield at extreme anterior end. Operculum horny, semicircular, not filling aperture.

<==Cassidae [Cassidites]S11
    |  i. s.: Pseudogaleodea tricarinata Nagao 1932TTE93
    |--Oocorys Fischer 1884BR05 [Oocorythidae, OocorythinaeBR17]
    |    |--*O. sulcata Fischer 1884BR17
    |    `--O. tosaensis Habe & Azuma 1959HA59
    |    |--CassisBR05
    |    |--Cypraecassis Stutchbury 1837 [=Cypraeicassis Cossmann 1903]W93
    |    |    |--*C. rufa (Linnaeus 1758)W93 (see below for synonymy)
    |    |    |--C. tenuis [incl. C. marsenae]C64
    |    |    `--C. testiculus (Linné 1758)S11
    |    `--Galeodea Link 1807DK08
    |         |  i. s.: G. apodemetesM49
    |         |         G. modestaK95
    |         |         G. triganceae Dell 1953P61
    |         |--G. (Galeodea) [incl. Cassidaria Lamarck 1812, Echinora Schumacher 1817, Morio Montfort 1810]W93
    |         |    |--*G. (G.) echinophora (Linnaeus 1758)P61, L58 [=Buccinum echinophorumP61]
    |         |    |--G. (G.) echinophorella Habe 1961W93
    |         |    |--G. (G.) maccamleyi Ponder 1983W93
    |         |    |--‘Morio’ rugosaPP64
    |         |    |--G. (G.) senexA27 [=Cassidaria senexH86, Struthiolaria senexH86]
    |         |    `--‘Cassidaria’ sulcataH86
    |         |--G. (Galeoocorys Kuroda & Habe 1957)W93
    |         |    |--G. (*G.) leucodomaW93
    |         |    `--G. (G.) nipponica (Sakurai & Habe 1961)W93
    |         `--G. (Taieria) klingeri Kiel & Bandel 2003BD12
         |--Echinophoria Sacco 1890 (see below for synonymy)W93
         |    |--*E. intermediaW93 [=Buccinum intermediumW93, Bezoardica intermediaF27a, Cassis intermediaF27a]
         |    |--E. carnosa (Kuroda & Habe 1961)W93
         |    |--*Bathygalea’ coronadoi (Crosse 1867)W93
         |    |--*Trachydolium’ dalli [=*Miogalea dalli]W93
         |    |--*Shichiheia’ etchuensisW93
         |    |--‘Phalium’ grangei (Marwick 1926) [=Euspinacassis grangei]F27a
         |    |--E. kurodai (Abbott 1968)W93
         |    |--‘Euspinacassis’ muricata [=Cassis muricata]F27a
         |    |--*Euspinacassis’ pollens Finlay 1926W93, F27a
         |    `--E. wyvillei (Watson 1886)W93
         |--Casmaria Adams & Adams 1853W93
         |    |--C. erinaceus (Linnaeus 1758)BW09 (see below for synonymy)
         |    |    |--C. e. erinaceusW93
         |    |    |--C. e. kalosmodix (Melvill 1883)W93
         |    |    `--C. e. vibexmexicana (Stearns 1894)W93 (see below for synonymy)
         |    |--C. perryi Iredale 1912 [=C. ponderosa perryi]W93
         |    |--C. ponderosa (Gmelin 1791) (see below for synonymy)W93
         |    |    |--C. p. ponderosaW93
         |    |    |--C. p. atlantica Clench 1944W93
         |    |    |--C. p. nipponensis Abbott 1968W93
         |    |    `--C. p. unicolor (Dautzenberg 1926)W93
         |    |--C. pyrumF27a
         |    `--C. radiata Tate 1889C54

Casmaria erinaceus (Linnaeus 1758)BW09 [incl. Casmaria denticulata Röding 1798W93, Casmaria edentulum Link 1807W93, Casmaria glabra Bolten 1798C54, Casmaria meles Dillwyn 1817C54, Casmaria panthera Solander 1797C54, Casmaria ventricosa Rigacci 1866W93, Buccinum vibex Linnaeus 1758W93, *Casmaria vibexW93, Cassidea vibexH09, Cassis (Phalium) vibexEO63]

Casmaria erinaceus vibexmexicana (Stearns 1894)W93 [=Cassis (Phalium) vibexmexicanaEO63, Cassis vibex mexicana (l. c.)EO63]

Casmaria ponderosa (Gmelin 1791) [incl. C. biarmatum Dillwyn 1817, C. cernica Sowerby 1888, C. nodulosum Gmelin 1791, C. pantherina Solander in Dillwyn 1817, C. quadratum Link 1807, C. tenuilabris Menke 1828, C. torquata Reeve 1848, C. turgida Reeve 1848]W93

*Cypraecassis rufa (Linnaeus 1758)W93 [=Buccinum rufumW93; incl. C. labiata Perry 1811W93, C. ostauri Humphrey 1797C54, C. pennatum Gmelin 1791W93, C. pullum Born 1778W93, C. rufescens Röding 1798W93, C. tuberosa Röding 1798W93, C. ventricosum Gmelin 1791W93]

Echinophoria Sacco 1890 [incl. Bathygalea Woodring & Olsson 1957, Euspinacassis Finlay 1926, Miogalea Woodring & Olsson 1957, Shichiheia Hatai & Nisiyama 1949, Trachydolium Howe 1926]W93

*Type species of generic name indicated


[A27] Allan, R. S. 1927. The geology and palaeontology of the Lower Waihao Basin, south Canterbury, New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 265–309.

[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.

[BW09] Bryce, C., & C. Whisson. 2009. The macromolluscs of Mermaid (Rowley Shoals), Scott and Seringapatam Reefs, Western Australia. Records of the Western Australian Museum Supplement 77: 177–208.

[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.

[C54] Cotton, B. C. 1954. Family Cassididae. Royal Society of Australia, Malacological Section 4.

[DK08] Darragh, T. A., & G. W. Kendrick. 2008. Silicified Eocene molluscs from the Lower Murchison district, Southern Carnarvon Basin, Western Australia. Records of the Western Australian Museum 24 (3): 217–246.

[EO63] Emerson, W. K., & W. E. Old Jr. 1963. Remarks on Cassis (Casmaria) vibexmexicana. Nautilus 76 (4): 143–145.

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

[HA59] Habe, T., & M. Azuma. 1959. A new Oocorys from Japan. Nautilus 72 (4): 116.

[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.

[K95] Köhler, R. 1995. A new species of the fossil turtle Psephophorus (order Testudines) from the Eocene of the South Island, New Zealand. Journal of the Royal Society of New Zealand 25: 371–384.

[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.

[M49] Mason, B. 1949. The geology of Mandamus-Pahau district, north Canterbury. Transactions and Proceedings of the Royal Society of New Zealand 77 (3): 403–428.

[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.

[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.

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

Wilson, B. R., & K. Gillett. 1971. Australian Shells: illustrating and describing 600 species of marine gastropods found in Australian waters. A. H. & A. W. Reed: Sydney.

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