4 
T.A. Darragh 
taxon, such as Protocypraea Schilder, 1927, in the earliest 
Tertiary or even in the late Cretaceous. Species of Cypraeorbis 
(recorded as Bernaya ) and Protocypraea have been recorded 
from the upper Cretaceous of North America by Groves (1990, 
2004) and Protocypraea is also known from the upper 
Cretaceous of India and the Paleocene of Pakistan (Cox, 1930). 
Schilder (1927) erected Barycypraea as a sub genus of Zoila 
with type species Cypraea (Aricia) caputviperae Martin, 1899 
(probably a synonym of Cypraea murisimilis Martin, 1879) for 
a group of cowries known from the upper Tertiary of Indonesia 
and living in the Indian Ocean. Molecular biological studies of 
cowries by Meyer (2004) showed that the nearest living relatives 
of the Australian living Zoila species are the western Indian 
Ocean species of Barycypraea and he grouped both genera in 
the subfamily Bemayinae. He included no other living taxa in 
this subfamily. Species of Barycypraea have a similar fossula 
(fig. 3L) to Zoila species and they are known to be spongivores 
like Zoila (Liltved, 1989). Species of Barycypraea occur in 
South Africa, Pakistan, India and Indonesia, and range in age 
from Miocene to Recent (Kay, 1990). The morphology and 
biogeography of Barycypraea suggests an origin similar to that 
of Zoila and a probable common ancestry. 
Terminology and repositories 
Measurements are given as follows: L = total length of shell; 
W = width of shell; H = height of shell. Tooth counts are cited 
as LT = labral teeth and CT = columellar teeth. The terms left 
and right refer to the animal’s true left and right sides, 
respectively. 
All material used in this study, unless otherwise stated, is 
held in the collections of Museum Victoria, registration 
numbers with prefixes P (invertebrate palaeontology 
collection) and F (living mollusca collection). Localities are 
cited where possible using the Museum Victoria fossil locality 
register with prefix PL (listed in Beu and Darragh, 2001). 
Museum acronyms used for other material are as follows: 
BM(NH) = Natural History Museum, London; SAM T = Tate 
Collection, South Australian Museum, Adelaide; AMNZ = 
Auckland Museum, New Zealand; AM C = Australian 
Museum, Sydney, New South Wales; WAM = Western 
Australian Museum; USNM = Smithsonian Institution, 
National Museum of Natural History; MNHN = Museum 
national d’Histoire naturelle, Paris; GNS TM = GNS Science 
(New Zealand). 
All specimens figured were coated with ammonium 
chloride for photography, unless stated otherwise. 
Class Gastropoda 
Family Cypraeidae Rafinesque, 1815 
Subfamily Cypraeorbinae Schilder, 1927 
Zoila Jousseaume, 1884 
Zoila Jousseaume, 1884a: 14.— Jousseaume, 1884b:89.— 
Cossmann, 1903:146,149.- Thiele, 1929:275.- Schilder, 1935:336.- 
Schilder and Schilder, 1939:173.—Wenz, 1941:971.— Dolin, 1991:11 
(synonomised Gigantocypraea Schilder, 1927). — Wilson and 
Clarkson, 2004:44. 
Cypraeorbis (Zoila) Jousseaume. Schilder, 1926:378. 
Umbilia (Gigantocypraea) Schilder, 1927:86 (type species, 
original designation, Cypraea gigas McCoy, 1867, Miocene, Victoria). 
Zoila (Zoila) Jousseaume. Schilder, 1935:337.— Schilder, 
1939:177. 
Zoila (Gigantocypraea) Schilder. Schilder, 1935:337.— Schilder, 
1939:177.- Wenz, 1941:971. 
Cypraea (Zoila) Jousseaume. Wilson and McComb, 1967:469. — 
Wilson, 1993:192. 
Type species. Cypraea scottii Broderip, 1832 (= Cypraea 
friendii Gray, 1831) by subsequent designation Jousseaume 
(1884b, p. 89), western and southern Australia. 
Diagnosis. Shell small (35 mm) to very large (247 mm) in size, 
highly glazed, varying in shape, elongate-ovate or pyriform or 
globose, ventrally flattened. Spire whorls usually covered in 
callus, rarely protruding. Protoconch, where known, consisting 
of one to two whorls, the first whorl large, somewhat irregular 
in shape, deviated slightly from shell axis. Anterior and 
posterior canals deeply incised, usually short, but on some 
species produced and bounded by weak to prominent lateral 
flanges. Aperture narrow, sinuous with weakly to strongly 
developed short to elongate teeth not extending into aperture or 
onto fossula. Fossula weakly to strongly developed, broad, 
slightly to deeply concave and bounded anteriorly by a weak to 
strong terminal ridge; on some species with very weak sulcus 
parallel and immediately posterior to terminal ridge. 
Time range. Late Paleocene/early Eocene-Recent. 
Distribution. India (early Miocene), Indonesia (Pliocene- 
Pleistocene), Western Australia (late Eocene, Miocene- 
Recent), South Australia (Miocene, Recent), Victoria (late 
Oligocene-late Miocene, Recent), Tasmania (early Miocene), 
New Zealand (late Paleocene/early Eocene). 
Remarks. Specimens of Zoila itoigawa Tomida, 1989 from the 
late Miocene of Japan are poorly preserved and have not been 
prepared sufficiently to enable generic assignment. 
As I have recognised two groups of species, an eastern and 
a western group, it could be argued that these should be 
accorded taxonomic status if there are significant 
morphological differences to separate them, in which case 
Gigantocypraea could be used for the eastern group. However, 
I regard these groups as more geographic entities. There are 
some morphological differences, but I regard them as minor 
and not of sufficient importance to use as generic characters. 
Such variations in morphology are to be expected in species 
that have no pelagic larval stages. 
Some species of the eastern group have considerably 
produced anterior and posterior canals — for example, Zoila 
platypyga (McCoy) and Z. mulderi (Tate) — but other species 
have canals similar to the living species of the western group 
— for example, Z. glomerabilis n. sp. is similar to Z. venusta 
(Sowerby, 1846) or Z. gendinganensis (Martin, 1899). 
Columellar dentition varies considerably from species to 
species. In the living species, columellar dentition can vary 
from strong to weak even within the one species; for example, 
