Belongs within: Tonnoidea.
Contains: Phalium, Semicassis, Echinophoria, Cassis.
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). Oocorys is a deepwater genus with a thin, pale shell and lacking obvious pigmented eyes.
Characters (Abbott 1968): Shell usually ovate, body whorl large; one or more varices; well-developed parietal or columellar shield; small, bulimoid, shelly nucleus. Operculum corneous, thin, semi-circular or fan-shaped, brown. Radula taenioglossate, about 80 to 220 rows of teeth, each row bearing seven rather delicate, minutely-cusped teeth. Proboscis pleurembolic, retracted within head when not in use. Eyes at base of moderately long, thin tentacles. Males with long, prong-like penis attached to right side of head, sperm travelling to top of penis along open groove. Females laying eggs within small capsules built into tower-like structures, capsules are formed by round sphincter in sole of foot.
<==Cassidae [Cassidites]S11
| i. s.: Pseudogaleodea tricarinata Nagao 1932TTE93
| Morionella Dall 1909A68
| `--M. cancellataA68
| ‘Buccinum’ plicatum Linnaeus 1758 (n. d.) [=Cassis plicata, Phalium plicatum]A68
|--Oocorys Fischer 1884BR05 [Oocorythidae, OocorythinaeBR17]
| |--*O. sulcata Fischer 1884BR17
| `--O. tosaensis Habe & Azuma 1959HA59
|--PhaliinaeBR05
| | i. s.: ‘Cassis’ bilineata Fleming 1828 [=Buccinum porcatum Pulteney 1799 non Gmelin 1791]A68
| |--PhaliumBR05
| |--SemicassisW93
| |--EchinophoriaW93
| `--Casmaria Adams & Adams 1853W93 [incl. Casmeria Jousseaume 1888A68]
| |--C. erinaceus (Linnaeus 1758)BW09 (see below for synonymy)
| | |--C. e. erinaceusW93
| | |--C. e. kalosmodix (Melvill 1883)W93 [=Cassis kalosmodixA68]
| | `--C. e. vibexmexicana (Stearns 1894)W93 [=Cassis (Phalium) vibexmexicanaEO63, Cassis vibex mexicana (l. c.)EO63]
| |--C. perryi (Iredale 1912)W93 [=Cassidea perryiA68, Casmaria ponderosa perryiW93]
| |--C. ponderosa (Gmelin 1791)W93 (see below for synonymy)
| | |--C. p. ponderosaW93
| | |--C. p. atlantica Clench 1944W93 [=C. erinaceus atlanticaA68]
| | |--C. p. nipponensis Abbott 1968W93 [=C. erinaceus nipponensisA68]
| | `--C. p. unicolor (Dautzenberg 1926)W93 (see below for synonymy)
| |--C. pyrumF26
| `--C. radiata Tate 1889C54
`--CassinaeBR05
|--CassisBR05
|--SconsiaP94
| |--S. ambigua [incl. S. tessellata Wrigley 1934]A68
| |--‘Cassis’ beyrichi Michelotti 1861A68
| |--S. hodgii (Conrad 1841)P94
| |--S. metae Petuch 1994P94
| |--S. prolongata Petuch 1994P94
| |--S. striataA68
| `--S. sublaevigata (Guppy 1966)P94
|--Cypraecassis Stutchbury 1837 [=Cypraeicassis Cossmann 1903]W93
| |--C. (Cypraecassis)A68
| | |--*C. (C.) rufa (Linnaeus 1758)W93 (see below for synonymy)
| | |--C. (C.) cypraeiformis (Borson 1820) (see below for synonymy)A68
| | |--C. (C.) pseudocrumena (Sacco 1890) (see below for synonymy)A68
| | |--C. (C.) pustulata (Cox 1927) [=Cassis (C.) pustulata; incl. Cassis (C.) morus Cox 1927]A68
| | |--C. (C.) subcrumena (Orbigny 1852) [=Cassis subcrumena]A68
| | |--C. (C.) subtesticulus (Orbigny 1852) (see below for synonymy)A68
| | |--C. (C.) tenuis (Wood 1828) [=Cassis tenuis; incl. Cassis massenae Kiener 1835]A68
| | `--C. (C.) testiculus (Linnaeus 1758)A68 (see below for synonymy)
| | |--C. t. testiculus [incl. Cassis testiculus var. bicincta Bayer 1935, Cassis pileolus Röding 1798]A68
| | `--C. t. senegalica (Gmelin 1791) (see below for synonymy)A68
| `--C. (Levenia Gray 1847)A68
| `--C. (L.) coarctata (Sowerby 1825) [=Cassis coarctata, Buccinum coarctatum, Cassis coarctatum]A68
`--Galeodea Link 1807DK08 [=Echinophora Latreille 1825 non Linnaeus 1753 (ICBN)A68]
| i. s.: ‘Cassidea (Semicassis)’ anteniana Martin 1931A68
| G. apodemetesM49
| G. arntzenii (Martin 1914) [=Cassidaria arntzenii]A68
| ‘Cassis’ bicatenatus Sowerby 1817A68
| G. bituminataFG19
| ‘Cassis’ brunnea Röding 1798A68
| G. bullata Brown 1839 [incl. Cassis gigantica Edwards in Newton 1891 (n. n.)]A68
| ‘Cassis’ caelata Conrad 1830A68
| G. ‘coronata’ (Deshayes 1830) (see below for synonymy)A68
| G. gallica Wrigley 1934 (see below for synonymy)A68
| G. geminata Wrigley 1934 [=Cassis geminata Edwards in Newton 1891 (n. n.)]A68
| ‘Cassis’ mississippiensis Conrad 1847A68
| G. modestaK95
| G. nodosa Solander 1766 [incl. Cassidea carinata Bruguière 1792, Cassis carinata Lamarck 1803]A68
| 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, *Echinophora echinophoraA68]
| |--‘Cassidaria’ desori d’Archiac & Haime 1854A68
| |--G. (G.) echinophorella Habe 1961W93
| |--G. (G.) maccamleyi Ponder 1983W93
| |--‘Cassidaria’ orbiculata Risso 1826A68
| |--‘Morio’ rugosaPP64
| |--G. (G.) senexA26 [=Cassidaria senexH86, Struthiolaria senexH86]
| |--‘Cassidaria’ sulcataH86
| `--‘Cassidaria’ thyrrenaP71
|--G. (Galeoocorys Kuroda & Habe 1957)W93
| |--G. (*G.) leucodoma Dall 1907W93, A68
| `--G. (G.) nipponica (Sakurai & Habe 1961)W93
|--G. (Galeodaria)A68
| |--G. (G.) petersoniA68
| `--G. (G.) shubutensis (Aldrich 1885) [=Cassis (Semicassis) shubutensis]A68
`--G. (Taieria) klingeri Kiel & Bandel 2003BD12
Casmaria erinaceus (Linnaeus 1758)BW09 [=Buccinum erinaceusA68, Cassidea erinaceusA68, Phalium (Casmaria) erinaceumA68; incl. Cassis denticulata Röding 1798A68, Phalium edentulum Link 1807A68, Casmaria glabra Bolten 1798C54, Buccinum meles Dillwyn 1817 (nom. inv.)A68, Casmaria melesC54, Casmaria panthera Solander 1797C54, Cassis vibex var. ventricosa Rigacci 1866 (n. n.)A68, Buccinum vibex Linnaeus 1758W93, *Casmaria vibexW93, Cassidea vibexH09, Cassis (Phalium) vibexEO63, Phalium erinaceum var. vibexA68]
Casmaria ponderosa (Gmelin 1791)W93 [=Buccinum ponderosumA68; incl. Buccinum biarmatum Dillwyn 1817A68, Cassis cernica Sowerby 1888A68, Cassidea cernicaA68, Buccinum nodulosum Gmelin 1791A68, Cassis nodulosaA68, Buccinum pantherina Solander in Dillwyn 1817A68, W93, Phalium quadratum Link 1807A68, Cassis quadrataA68, Cassis tenuilabris Menke 1828A68, Cassis torquata Reeve 1848A68, W93, *Casmeria torquataA68, Cassidea nodulosa var. torquataA68, Cassis turgida Reeve 1848A68]
Casmaria ponderosa unicolor (Dautzenberg 1926)W93 [=Cassis (Casmaria) turgida var. unicolorA68, Phalium turgidum var. unicolorA68]
Cypraecassis (Cypraecassis) cypraeiformis (Borson 1820) [=Cassis cypraeiformis, Cassidea cypraeiformis; incl. Cassidea cypraeiformis var. crassilabiata Sacco 1890, Cassidea cypraeiformis var. reticulosa Sacco 1890, Cassidea cypraeiformis var. tuberculosa Sacco 1890]A68
Cypraecassis (Cypraecassis) pseudocrumena (Sacco 1890) [=Cassidea crumena var. pseudocrumena, Cassis pseudocrumena Orbigny 1852 (n. n.); incl. Cassidea crumena var. pseudocrumena subvar. miocenica Sacco 1890, Cassidea crumena var. pseudocrumena subvar. multinodosa Sacco 1890, Cassidea crumena var. pseudocrumena subvar. ornata Sacco 1890, Cassidea crumena var. pseudocrumena subvar. rarituberculata Sacco 1890, Cassidea crumena var. pseudocrumena subvar. testiculoides Sacco 1890, Cassidea crumena var. pseudocrumena subvar. venusta Sacco 1890]A68
*Cypraecassis (Cypaecassis) rufa (Linnaeus 1758)W93 [=Buccinum rufumW93, Cassidea rufaA68, Cassis rufaA68, Cassis rufumA68; incl. Cassis chartoprates Meuschen 1786 (nom. inv.)A68, Cassis labiata Perry 1811W93, Cassida ostauri Humphrey 1797 (nom. inv.)A68, Buccinum pennatum Gmelin 1791W93, Cassidea pennataA68, Cassis pennataA68, Buccinum pullum Born 1778A68, Cassis rufescens Röding 1798A68, Cassis tuberosa Röding 1798 non Buccinum tuberosum Linnaeus 1758W93, Buccinum ventricosum Gmelin 1791A68]
Cypraecassis (Cypraecassis) subtesticulus (Orbigny 1852) [=Cassis subtesticulus; incl. Cassidea crumena var. atuberculata Sacco 1891, Cassis marginata var. initalis Sacco 1890, Cypraeicassis marginata var. initalis, Cassis marginatus de Serres 1829 non Borson 1820, Cassis marginata, Cypraeicassis marginata, Cassis marginata var. ornata Sacco 1890, Cypraeicassis marginata var. ornata, Cassis plicata Defrance 1817 nec Buccinum plicatum Linnaeus 1758 nec Scopoli 1786, Cassis (Cassidea) protesticulus Sacco 1890, Cassis ruscinensis, Cassis marginata var. savonensis Sacco 1890, Cypraeicassis marginata var. savonensis]A68
Cypraecassis (Cypraecassis) testiculus (Linnaeus 1758)A68 [=Buccinum testiculusW62, Cassis testiculusW62; incl. Cassis mitella polonica Röding 1798A68]
Cypraecassis (Cypraecassis) testiculus senegalica (Gmelin 1791) [=Buccinum senegalicum, Cassis testiculus var. senegalica; incl. Cassidea crumena Bruguière 1792, Cassis testiculus var. crumena, Cypraecassis testiculus f. crumena, Cassis (Cypraecassis) senegalica var. mamillata Salmon 1948, Buccinum plicatum Gmelin 1791 nec Linnaeus 1758 nec Martyn 1784]A68
Galeodea ‘coronata’ (Deshayes 1830) [=Cassis coronata non Röding 1798; incl. C. venatrix Edwards in Newton 1891 (n. n.)]A68
Galeodea gallica Wrigley 1934 [=Cassis carianta Sowerby 1812 non Lamarck 1803; incl. G. clarendonensis Wrigley 1934, C. substriata Edwards in Newton 1891 (n. n.)]A68
*Type species of generic name indicated
References
[A68] Abbott, R. T. 1968. The helmet shells of the world (Cassidae). Part 1. Indo-Pacific Mollusca 2 (9): 15–201.
[A26] Allan, R. S. 1926. 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.
[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.
[FG19] Fedosov, A. E., M. C. Gutierrez, B. Buge, P. V. Sorokin, N. Puillandre & P. Bouchet. 2019. Mapping the missing branch on the neogastropod tree of life: molecular phylogeny of marginelliform gastropods. Journal of Molluscan Studies 85: 439–451.
[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 revised 10th ed. vol. 1. 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.
[P71] Panetta, P. 1971. Molluschi del Golfo di Taranto raccolti durante le crociere dell’Albatros. Annali del Museo Civico di Storia Naturale “Giacomo Doria” 78: 149–180.
[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.
[P94] Petuch, E. J. 1994. Atlas of Florida Fossil Shells (Pliocene and Pleistocene Marine Gastropods). Chicago Spectrum Press: Evanston (Illinois).
[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.
[W62] Weisbord, N. E. 1962. Late Cenozoic gastropods from northern Venezuela. Bulletins of American Paleontology 62: 7–672.
[W93] Wilson, B. 1993. Australian Marine Shells vol. 1. Prosobranch Gastropods. Part One. Odyssey Publishing: Kallaroo (Western Australia).