B. M. Landau & L. T. Groves 
NOVAPEX 12(1-2): 1-38, 10 mars 2011 
axially striped colour pattern or blotching of dull 
reddish brown or yellowish orange. Additionally, the 
marginal callus is extremely thin and poorly delimited, 
and does not extend onto the dorsum, and the shell 
periphery of the Gatun shells feel corrugated, whereas 
they are smooth in M. henekeni and M. hyaena. We 
separate two distinct taxa within the Gatun 
assemblages (see below). The early Pliocène 
Muracypraea grahami (Ingram, 1947) t'rom Cubagua, 
Venezuela (Figs 29-32) is again different (Landau & 
Silva, 2010). Ingram (1947) described two 
Muracypraea species from Cubagua; Cypraea nigosa 
[junior homonym of C. nigosa Broderip, 1827] and 
Cypraea grahami. Although the holotypes of the two 
shells look quite different, Landau & Silva (2010) 
illustrated intermediate forms and recognised a single 
species in the Cubagua deposits. The shell is very 
large and solid (63-73 mm in length) and their outline 
is more triangular and their dorsum even higher than 
M hyaena (Landau & Silva 2010). The tubercles are 
staggered on the dorsum in a similar fashion to M. 
hyaena. The margins are corrugated, as in 
Muracypraea isthmica Schilder, 1927, but more 
coarsely so. In the best preserved specimen from 
Cubagua an indication of a striped colour pattern is 
présent, akin to that seen in the Gatun Formation 
shells. On the Pacific side of the Gatunian Province 
Muracypraea cayapa (Pilsbry & Olsson, 1941) from 
the early Pliocène Jama Formation of Ecuador is a 
large poorly known species represented by a single 
broken specimen. Many other taxa of Muracypraea 
hâve been described from the Caribbean Neogene, 
which awail validation. Species of the extant 
Muracypraea mus complex are known to hâve an 
intracapsular larval development (Ranson, 1967; 
Hoeblich, 1979). This assumption cannot be validated 
with the fossil taxa, however, this type of development 
would favour a restricted distribution of species and a 
high degree of variability between populations. This 
helps explain the difficulty encountered with the 
Caribbean Neogene Muracypraea populations and 
may be analogous to the situation among Recent 
species of the genus Zoila Jousseaume, 1884 in 
Australia which also shows intracapsular 
development. Numerous species of Zoila are not only 
geographically restricted, but numerous intermediate 
forms are common ofif the western and Southern coasts 
of Australia (Lorenz, & Hubert, 2000; Lorenz, 2001). 
Based on mitochondrial molecular data presented 
by Meyer (2004), the two extant Barycypraea taxa B. 
teulerei (Cazenavette, 1846) and B. fultoni (Sowerby, 
1903) were placed as sister taxa to Zoila Jousseaume, 
1884, reaffirming the (validity of the subfamilial clade 
Bernayinae Schilder, 1927, whereas Muracypraea 
mus was placed in the Cypraeinae Gray, 1824. Based 
on morphological différences, Fehse (in press, a) will 
reassign the two living Barycypraea species and the 
fossil species B. zietsmani (Liltved & LeRoux, 1988) 
[Pliocène, South Africa], B. schilderi (Dey in 
Schilder, 1941) [Pliocène, India], B. gendingaensis 
(Martin, 1899) [Pliocène, Indonesia], and B. 
kendengensis (Schilder, 1941) [Pliocène, Indonesia] to 
a new genus (Dirk Fehse personal communication 
2010). Some of the shells of the lndonesian fossil 
Barycypraea group are extremely similar to those of 
Muracypraea in the Tropical American Neogene and 
could represent a convergent evolutionary process. A 
morphological feature that helps separate 
Muracypraea from Barycypraea are the flattened 
spatulate horizontal expansions on either side 
produced from the abapical tips of the inner and outer 
lips of M. henekeni and many related species. 
Occasionally these spatulate expansions are hyper 
developed in the Caribbean forms whereas they are 
somewhat obsolète in the lndonesian species of 
Barycypraea. Although the two généra may hâve a 
common Tethyan ancestor, consistent shell différences 
and the vast géographie séparation could justify 
different généra. Many of the Muracypraea shells are 
lightweight, less callused with a broad aperture and 
less developed, widely spaced dentition, whereas the 
shells of Barycapraea are heavy, strongly callused 
with a narrow aperture and a distinct, close-set 
dentition. The Muracypraea exceptions being M. mus 
donmorei (Petuch, 1979), M mus bîcornis (Sowerby, 
1870), and M. grahami (C. nigosa Ingram, 1947 
junior homonym of C. rugosa Broderip, 1827; see 
Landau & Silva 2010). The dorsal callosities of both 
généra can be variable even among the same species. 
However, in Muracypraea they tend to be restricted to 
the posterior portion of the dorsal surface whereas in 
Barycypraea there may be a second set located 
slightly anterior of the dorsum centre. Because the 
Miocene fossil record for the central Pacific islands is 
incomplète, and until a conclusive answer to whether 
two généra should be used for the two seemingly 
geographically separated species-groups, the 
classification of Schilder & Schilder (1971) for 
Muracypraea is used for the Caribbean species. 
Muracypraea hyaena (Schilder, 1939) 
Figs 1-14 
1939 Siphocypraea angustirirna hyaena Schilder, 
p. 23, fig. 24. 
1971 Siphocypraea (Muracypraea) angustirirna 
hyaena Schilder and Schilder, p. 29. 
Material. 8 specimens BL coll.; 1 specimen LACMIP 
13648 (ex BL coll.). 
Description. Shell small to medium, solid, rotund, 
dorsum very strongly humped % distance from the 
anterior end, in fully adult specimens bearing two 
small tubercles, right tubercle larger and anterior to 
left; spire depressed, covered by adapical callus; sides 
rounded, strongly callused, with the callus ascending 
progressively adapically to just below the apex 
posterior to the dorsal hump; ventrum flattened, 
slightly convex in profile; aperture almost straight. 
