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Cypraeidae of the Indo-Pacific: Cenozoic Fossil History and Biogeography

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Cypraeidae of the Indo-Pacific: Cenozoic Fossil History and Biogeography BULLETIN OF MARINE SCIENCE, 47(1): 23-34, 1990 CYPRAEIDAE OF THE INDO-PACIFIC: CENOZOIC FOSSIL HISTORY AND BIOGEOGRAPHY E. Alison Kay ABSTRACT The cowries (Cypraeidae) of the Indo-Pacific are described in terms of their origins in the Tethys and Indonesian Tertiary, and their distribution in space in the Tertiary and today. Twenty-one species groups are identified in the discussion. The earliest Indo-Pacific cowries are recorded in the Palaeocene of Pakistan and India, then part of the Tethys Sea. In Indonesia the earliest cowries are Middle Miocene. Six of 18 species groups were represented in Pakistan- India. First occurrence of four species groups occurs in Indonesia; six groups appear contem- poraneously in the Pacific and in Australia. Five groups present in the Indonesian Miocene and Pliocene are now extinct in Indonesia, although species in two groups, Barycypraea and Zoila, live today in the Indian Ocean and Australia respectively. The distribution of cowries in the Indo-Pacific is not random. The patterns of spatial distribution in the fossil record and of Recent endemism in the central Pacific and Indian Ocean indicate speciation occurred beyond the geographical boundaries of the Malay Archipelago and provide support for Ladd's hypothesis of the importance of peripheral speciation in the Indo-Pacific. The question is deceptively simple: what is the relationship of the Recent Cypraeidae (true cowries) of the Indo-Pacific to the Indonesian Tertiary? The question stems from Ekman's (1953: 18) thesis that the Indo-Malayan archipelago is "centre and focus from which the others recruited its fauna" (see also Briggs, 1974). More than 80 species of fossil cowries are reported from the Indonesian Tertiary, and it seems a reasonable assumption that knowledge of the history of these cowries can provide some insight into paleobiogeography and evolution in the Indo-Pacific biota. The following account is a first approximation, in a purely descriptive mode, towards answering the question. The Indonesian cowrie fauna is described in terms of its origins, its composition over time, and its distribution in space both in the Tertiary and today. SOURCES OF DATA AND CAVEATS This review, which involves about 250 species level taxa and 25 species complexes, has been assembled from a variety of sources. The major fossil collections studied are those of Pakistan, India and Burma described by Forbes, Sowerby, Vredenburg and others in the 19th century, and now in the British Museum (Natural History), and the Indonesian Tertiary cowries in the Martin Collection, Rijksmuseum fUr Geologie, Leiden. Collections of Recent cowries in the B. P. Bishop Museum, Honolulu, Hawaii; the British Museum (Natural History), London; C. M. Burgess collection, Honolulu, Hawaii; California Academy of Science, San Francisco, California; the U.S. National Museum of Natural History, Washington, D.C., and the Zoologisch Museum, Universitet Amsterdam, provided much of the data for present distribution records. For purposes of dating the Pakistan-India-Burma and Indonesian Tertiary faunas, Shuto's (1975; 1976; 1984) correlations of Neogene formations of southeast and south Asia have been followed. I also follow Shuto (1976) in applying a broad brush to the history of the cowries: Shuto notes that while molluscan faunas within a given subprovince (for example, Pakistan and Burma) can be correlated stage to stage, inter-subprovincial correlations (between Pakistan and Indonesia) can only be described by broad terminologies such as Middle Miocene, Upper Miocene, Pliocene, etc. Shuto's (1976) sub- province of Pakistan, India, and Burma is here referred to as Pakistan-India. The term "Indonesia" is used in referring to the Miocene to Pliocene fossils of that area, and Malay Archipelago (encompassing Indonesia and the Philippines) is used sensu Ekman (1953) for Recent distributional records. Terminology for the distribution of Recent cowries in the Indo-Pacific, Indo-West Pacific, and Western Pacific is that of Springer (1982). Indo-Pacific refers to "the Indian Ocean including contiguous 23 24 BULLIETIN OF MARINE SCIENCE, VOL. 47, NO. I, 1990 Table I. Species groups in the Cypraeidae represented in the Indo-Pacific, Pakistan-India, and In- donesian Tertiary. * = group extinct. Eras: Mio, Miocene; Olig, Oligocene; Plio, Pliocene; Rec, Recent. Locales: Afr, Africa; Austr, Australia; Carib, Caribbean; CP, Central Pacific; Ene, Marshall Islands; Indon, Indonesia; 10, Indian Ocean; IWP, Indo-west Pacific; Jap, Japan; Mal, Malay Archipelago; Med, Mediterranean; NZ, New Zealand; Okin, Okinawa; Pak-Ind, Pakistan-India; Sind, South India; WArn, Western Americas; WP, Western Pacific. Characters: ap, aperture; bc, bursa copulatrix; dent, denticle; foss, fossula; into bract, internal bract; Rad, radula Barycypraeat Foss. ribbing obsolete. Mantle brown, black. Mio: Pak-Ind., Indon, Carib. Rec: I 10, 1 Rad. tooth shaft: lateral internal bracts, later- Carib. Shell 30-60 mm; depressed; ap. den- al basal dent. Females: bc. tate or edentate. Foss. smooth. Mantle thin, Lyncina mottled tan. Rad. tooth shaft: paired basal Mio: Pak-Ind, Indon., Fiji, Guam. Rec: 4 IP, dent., into bract. Females: bc. Dev. direct. 310, 4WP, 7 CPo Shell 30-70 mm, cylindri- Bistolida calor pyriform; ap. teeth usually across base. Plio: Indon. Rec: 7 IP, 1 Mal, 3 CPo Shell 13- Foss. ribbed. Mantle dark, gray/brown. Cen- 26 mm; cylindrical, ovate; ap. teeth barely on tral rad. tooth domed; Rad. tooth shaft: no base. Foss. ribbed. Mantle gray/pale yellow. basal dent. or bracts. Females: bc. Rad. tooth shaft: no basal dent. Females: be. Mauritia Blasicrura Mio: Indon; Plio: New Hebrides, Tonga. Rec: Plio: Indon. Rec: I Mal, 4 IWP; I 10, I WP. 5 IP, 2 10, I CPo Shell 35-100 mm; mar- Shell 12-20 mm; pyriform-ovate; ap. teeth on gined; ap. teeth confined to ap. Foss. ribbed. base. Foss. ribbed. Mantle dark, gray-brown. Rad. tooth shaft: paired basal dent., internal Rad. tooth shaft: no basal dent. Females: be. and subtending bracts. Females: no bc. Chelycypraea Miolyncina* Rec: liP. Shell 100 mm; inflated; ap. teeth Eoc.-Mio. Pak-Ind, Indon. Shell 30-60 mm; confined to ap. Foss. ribbed. Mantle transpar- ovate; ap. v-shaped, labrum declivous; ap. ent. Rad. tooth shaft: no basal dent. or inter- teeth confined to ap. and in post. channel. nal bract; laterals reduced. Females: be. Foss. ribbed. Cribrarula Nesiocypraeat Mio: Indon, Fiji, Jap. Rec: liP, 1 WP, 4 10, Rec: 6 WP. Shell 30-50 mm, pyriform; ap. 4 CPo Shell 15-30 mm; ovate, cylindrical; teeth confined to ap. Foss. smooth. Rad. margined; ap. teeth on base. Foss. absent or tooth shaft: with medial dent., into bract. ribbed. Mantle crimson. Rad. tooth shaft: paired basal dent. Females: be. Notadusta L. Olig: NZ; Mio: Indon, Austr. Rec: 3 WP. Cypraea Shell 15-24 mm; ovate, rostrate; ap. teeth Mio: Indon., Guam. Rec: 2 [P, 2 10. Shell confined to ap. Foss. smooth. Col. smooth 75-100 mm; cylindrical-inflated; ap. teeth confined to ap. Foss. ribbed. Mantle gray- Ovatipsa brown. Rad. tooth shaft: basal dent., into Mio: Indon. Rec: liP, I 10. Shell 20-30 bract. Females: bc. mm; pyriform; ap. teeth coarse, across base. Foss. ribbed. Rad. tooth shaft: no basal dent. Erosaria Female: no bc. Mio: Pak-Ind, Indon, Fiji, Guam. Rec: 13 IP, I 10, 10 CPo Shell 10-50 mm; col. margin Proadusta* pitted; ap. teeth on base. Foss. ribbed. Mantle Mio: Pak-Ind, Indon. Described by Schilder dark, brown/gray. Rad. tooth shaft: single (1939) but not easily distinguished from other basal dent.; internal, subtending bracts. Fe- genera. males: no bc. Purpuradusta Erronea Plio: Indon. Rec: 4 IP, 5 IWP, I CPo Shell 6- Mio: Pak-Ind, Indon, Fiji, Ene. Rec: 7 IP, 4 20 mm, cylindrical; ap. teeth confined to ap. IWP, 7 WP, I Mal. Shell 20-40 mm; pyri- Foss. ribbed. Mantle red. Rad. tooth shaft: form, ovate; ap. teeth on base. Foss. ribbed. paired basal dent. Females: no bc. Mantle dark, brown/gray. Rad. tooth shaft: Pustularia paired basal dent., int. bract. Females: no bc. Mio: Indon, Fiji. Rec: 5 IP, I CPo Shell 15-20 Luria mm; globular to inflated-ovate; ap. finely Mio: Pak-Ind, Indon, Fiji. Rec: liP, I CP, I dentate, teeth on base. Foss. smooth. Mantle 10, I Carib., I WArn. Shell 25-40 mm; cy- yellow, gray. Rad. tooth shaft: paired basal lindrical, pyriform; ap. teeth confined to ap. dent. KAY: BIOGEOGRAPHY OF INDO-PACIFIC CYPRAEIDAE 25 Table I. Continued Ta/paria confined to ap. Foss. ribbed. Mantle red. Plio: Indon, Fiji, Okin, E. Afr. Rec: lIP, I Central rad. tooth domed; rad. tooth shaft: no 10. Shell 60-70 mm; cylindrical-ovate; ap. basal denticles. Females: bc. teeth confined to ap. Foss. ribbed. Mantle Zoi/a dark. Rad. tooth shaft: paired basal denticles, OIig-Mio: Austr. SInd; Plio: Indon. Rec: 5 into bract. Females: bc. Austr. Shell 45-100 mm; ovate, subpyriform, Ta/osto/ida humped; margined; ap. weakly dentate. Foss. Pleist: Indon. Rec.: lIP, 2 CPo Shell 10-25 smooth. Rad. tooth shaft: paired basal dent., mm; ovate, cylindrical; margined; ap. teeth internal and subtending bracts. Females: bc. t I agree with Liltved and Le Roux (1988) that Cypraea mus is a relict Barycypraea and not Siphocypraea as defined by Petuch (1979). * Luque (1980) suggests that the western Pacific deepwater cowries be included with Cypraea pyrom and other Atlantic cowries in Schilder;a. Nesiocypraea is recognized as distinct here until more anatomical evidence is available. seas, and the Pacific Ocean as far east as Easter Island but excluding the area occupied by the coast and offshore islands ... ofthe Western Hemisphere" (Springer, 1982). Indo-West Pacific refers to the Indian Ocean and the western Pacific. By western Pacific is meant the Pacific Ocean west of the western margin of the Pacific lithospheric plate which includes inland seas such as the South China Sea, Arafura Sea, and Coral Sea.
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