New species of Triviidae (Mollusca: Gastropoda) from South ... - BioOne

PROCEEDINGS

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ACADEMY

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NATURAL SCIENCES

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PHILADELPHIA 151: 23–30.

31 DECEMBER 2001

New species of Triviidae (Mollusca: Gastropoda) from South Africa, Namibia and the Philippines GARY ROSENBERG Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia PA 19103-1195, U.S.A.—[email protected]

CHARLES C. FINLEY 12117 Merricks Court, Monrovia MD 21770, U.S.A.

ABSTRACT—Three new species of triviid gastropods are described. Galeatrivia Cate, 1979 is synonymized with Triviella Jousseaume, 1884. Thirteen species named as Trivia or Galeatrivia are transferred to Triviella. SPECIES NOVAE: Trivellona bealsi (from the Philippines), Triviella immelmani (from South Africa), Trivia marlowi (from Namibia).

praeoidean morphology needs clarification, being inconsistent with that generally used for gastropod shells (see Cox, 1960 for a glossary). First, teeth on the outer lip have incorrectly been called ‘‘labial teeth’’ by some authors (e.g., Schilder, 1930a; Burgess, 1985); properly they are ‘‘labral teeth.’’ The distinction between ‘‘labrum’’ for outer lip and ‘‘labium’’ for inner lip dates back at least to Sowerby (1839). Second, teeth on the inner lip of cowries usually have been called ‘‘columellar teeth.’’ Strictly, only teeth located from the posterior end of the fossula to the anterior terminal are columellar; teeth posterior to the fossula are parietal. Because the teeth form a continuous series, however, it is convenient to have a single term to refer to them. Since the term ‘‘labial teeth’’ has been misused with cowries, we maintain for the moment use of the term ‘‘columellar teeth.’’ This requires using ‘‘columellar’’ to mean ‘‘located on the side of the aperture bearing the columella,’’ rather than ‘‘located on the columella.’’ (Such imprecise usage is noted but not condoned by Sowerby, 1839, in his definition of ‘‘columella.’’) Third, the area posterior to the fossula has been called the ‘‘columellar sulcus’’ (Schilder, 1936a: 75), the ‘‘columellar groove’’ (Burgess 1985: xiv) or the ‘‘columellar depression’’ (Cate 1973: 3; Liltved, 2000: 34); we use the term ‘‘depression’’ since the structure often is not strong enough to be called a groove or sulcus. Fourth, the ridges on either side of the fossula and columellar depression have not been named consistently. The outer ridge marks the edge of the ‘‘columellar peristome’’ illustrated by Liltved (2000: 36); the inner ridge has been called the ‘‘adaxial ridge’’ by Cate (1973: 3), but this does not differentiate it from the outer ridge: both ridges are deposited along the adaxial surface of the aperture and the inner ridge is not necessarily closer to the shell axis than the outer ridge. We refer to these structures here as the ‘‘inner columellar ridge’’ and the ‘‘outer columellar ridge.’’

INTRODUCTION Since the publication of Cate’s monograph in 1979, a number of new species of Triviidae have been described. By illustrating many type specimens, Cate facilitated species level research on triviids. Classification of triviids at the genus level, however, is still difficult because of lack of knowledge of the anatomy of most taxa. Gosliner and Liltved (1982: 126) stated of Cate’s (1979) classification of the Triviidae: ‘‘As the present generic distinctions are based solely on inconsistent conchological differences, we prefer to classify all species within the single genus Trivia, until sufficient morphological study provides additional characters for meaningful comparison of taxa.’’

In 1987 (p. 252), Gosliner and Liltved further criticized Cate’s shell-based system: ‘‘The fact that Trivia costata was placed in Triviella, T. pellucidula in Trivirostra and T. suavis in Pseudotrivia by Cate, on the basis of their conchological features, but are similar in their reproductive morphology to T. arctica (placed in Trivia s.s.) further suggests that a great deal more morphological information is required before monophyletic subdivision of the Triviidae can be obtained.’’

We agree that a monophyletic classification of the Triviidae is desirable but not yet attainable. When such a classification is realized, however, it will surely result in Trivia being split into several if not many genera, as was first proposed by Jousseaume (1884a, b). We have therefore considered what is known of taxonomy, anatomy, and shell morphology to name our new species in the genera in which they might be placed when the classification of the group stabilizes. Some of the terminology traditionally applied to cy23

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G. ROSENBERG AND C. C. FINLEY

SYSTEMATICS Trivellona bealsi, n. sp. (Figs. 1–10) Description.—Shells globose, light weight, entirely white, medium sized, ranging from 9.9 to 12.3 mm. Extremities produced, especially anterior one, so that shell has both anterior ridge and funiculum. Spire slightly produced and covered with several ribs when fully adult. Ribs total 59 to 71, counted at the periphery, often with more ribs on right that left side (32–37 vs. 29–34). Ribs somewhat triangular in cross-section, with a rounded top and concave sides. Dorsal sulcus distinct but shallow, crossed by 6–12 out of 18–22 ribs that reach it. Labrum with 16–20 denticles protruding into aperture. In some specimens some of these denticles expand slightly above base, like tabs on jigsaw puzzle pieces. Outer columellar ridge with 16–22 denticles; columellar depression with 16–22 ribs; inner columellar ridge with 18–20 denticles, including 7–9 on fossula. Outer columellar ridge continuous with anterior ridge but not with funiculum, continuing parallel to inner columellar ridge posterior to point where funiculum starts. Fossula excavated and anteriorly expanded. Denticles on both sides of aperture strong and gearlike in structure, being about as wide as their interspaces. Aperture constant in width except for slight anterior widening. Right margin callused and somewhat beveled centrally. Right marginal callus, terminals and base with microscopic granulations between ribs. Granulations also occur decreasingly up to periphery; intercostal spaces glossy smooth above periphery. Intercostal spaces, especially on right side, with slight scalloped depressions, barely visible even with magnification. Depressions evenly spaced within intercostal space and aligned between them, possibly following growth lines of juvenile shell. Type Locality.—Balut Island, south of Mindanao, Philippines, 150 m. HOLOTYPE: L 11.1 3 W 8.8 3 H 7.9 mm, live in gill net, March 1992, ANSP 406466. PARATYPES: 1) L 11.4 mm, W 8.8 mm, H 7.9 mm, live, Balut Island, south of Mindanao, Philippines, gill net, 200 m, March 1987, ANSP 408194. 2) 10.2 3 8.2 3 7.5 mm, live, Balicasag Island, off Bohol, Philippines, gill net, 100–150 m, 1990, Finley Collection. 3) 11.1 3 8.8 3 8.0 mm, dead, Balicasag Island, Bohol, Philippines, gill net, 150 m, 1990, BMNH. 4) 12.3 3 9.4 3 8.8 mm, live, Aliguay Island, off NW Mindanao, Philippines, dredged 150 m, September 2000, Beals Collection. 5) 10.2 3 8.2 3 7.5 mm, live, Balicasag Island, off Bohol, Philippines, tangle net, 150 m, December 1999, Beals Collection. 6) 9.9 3 7.7 3 6.9 mm, live, Balicasag Island, off Bohol, Philippines, tangle net, 150 m, Beals Collection. 7) The specimen illustrated as Trivellona schepmani by Dolin (2001, fig.

20a-b), from off Bohol Island, Philippines, Dolin Collection [specimen not seen, length cited as 12.0 mm]. Other Material.—One shell from the Leonard C. Hill Collection, current whereabouts unknown. Distribution.—Presently known only from the Philippines in 100–200 meters. Comparison.—Trivellona bealsi most closely resembles Trivellona sibogae (Schepman, 1909), which is the type species, by original designation, of Pseudotrivia Schilder, 1936b, a synonym of Trivellona Iredale, 1931 according to Dolin (2001). Trivellona sibogae differs in lacking a dorsal sulcus and in having much weaker apertural dentition, especially on the labrum and the fossula. Trivellona schepmani (Schilder, 1941) also lacks the dorsal sulcus, has much weaker apertural dentition, and a broader aperture. Trivellona bealsi also resembles Trivellona eos (Roberts, 1913), most notably in having weak, aligned intercostal depressions, but T. eos is a much larger shell with weaker apertural dentition and a proportionately wider aperture. Etymology.—Named for Mr. Marty Beals of Inglewood, California, who first brought this new species to our attention. Triviella immelmani, n. sp. (Figs. 11–20) Description.—Shells subglobular, medium sized, from 12.5 to 14.0 mm in length. Dorsum smooth, highly glossy. Spire whorls visible through shell. Dorsum cream to light pink; base white. One specimen (paratype 4) has five diffuse light brown spots dorsally. Outer lip strong, thickest medially, bearing 16 to 24 ribs, 11–15 of which project into aperture as prominent denticles. Outer columellar ridge with 14–15 denticles; columellar depression with 14–17 ribs, including 6–8 on fossula. Posteriormost two or three columellar teeth extend as denticles on inner columellar ridge, but teeth anterior to them do not, except near and on fossula. Columellar teeth do not extend out of aperture, except for first two or three anteriorly. Second of anterior teeth is strongest. Right margin has strong white callus, left margin not callused. Areas between labral teeth with small granulations, elongated axially. Terminal areas also bear small granulations. Mantle in alcohol preserved specimen (paratype 1) brownish yellow with numerous black spots. Type Locality.—Off East London, Cape Province, South Africa. HOLOTYPE: L 13.2 mm, W 10.8 mm, H 9.5 mm, dredged live, 100 m, July 1997, NMSA V5573/T1311. PARATYPES: Same data as holotype unless otherwise noted. 1) 14.0 3 11.2 3 10.3 mm, preserved in ethanol, ANSP A19459; 2) 12.7 3 10.8 3 9.1 mm, live, Hayes Collection; 3) 12.5 3 11.2 3 9.1 mm, live, Finley Collection; 4) 13.6 3 11.4 3 9.5 mm, dredged live, 80 m, November 1997, Beals Collection. Other Material.—Richard Kilburn (email, 8 October

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NEW SPECIES OF TRIVIIDAE

Figs. 1–10. Trivellona bealsi n. sp. 1–5, HOLOTYPE, length 11.1 mm. 6–10, base on right side in fig. 9 is a repaired break.

PARATYPE

#1, length 11.4 mm. Indentation above

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Figs. 11–20. Triviella immelmani n. sp. 11–15,


HOLOTYPE,

length 13.2 mm. 16–20,

PARATYPE

#2, length 12.7 mm.

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Table 1. Characters used by Cate (1979: 15) to distinguish Galeatrivia from Triviella. As shown in the comments, these characters are variable and do not distinguish the genera. Character

Galeatrivia

Comments

Triviella

Size

‘‘slightly smaller’’

[slightly larger]

Variable; e.g., Cate’s (1979) measurements show T. vesicularis as smaller than G.

Shell color

‘‘grey to greyish pale pink’’ ‘‘low, narrowly elongate lateral profile’’

‘‘more or less constant shade of pink to red’’ ‘‘more roundly inflated subglobular’’

Sculpture

‘‘tendency to longitudinal growth lines’’

‘‘transverse incised striations’’

Apex profile

‘‘protrudes’’

‘‘buried’’

Apex exposure

not ‘‘overlaid with nacre’’

‘‘overlaid with nacre’’

Variable within species, e.g., G. ovulata is pale or dark pink (Liltved 2000). Degree of inflation is variable; see species 67 to 87 (excluding 79) in Liltved (2000). Longitudinal growth lines occur in both groups; transverse incised striation in neither. ‘‘Spire may be barely visible or conspicuously produced’’ in G. millardi (Liltved 2000). Apex covered with shell material in both G. ovulata and G. millardi.

millardi.

Shell shape

2001) reports the following material from Transkei in the Natal Museum, South Africa (NMSA). We have not seen these specimens, having learned of them too late to include when formulating the species description. Off Sandy Point, 328 40.39 S, 288 40.49 E, 97 m, gorgonians, stylasterids, sponges, two live (one body in alcohol), NMSA C4592; off Xora River, 328 33.49 S, 288 48.09 E, 100 m, coarse sand, some sponge rubble, one dead, NMSA C4840; off Mendu Point, 328 22.19 S, 288 59.29 E, 110–112 m, stylasterids, sponge, gorgonians, one dead, NMSA C4296; off Port Grosvenor, 318 25.29 S, 298 56.89 E, 80–81 m, calcareous nodules, two dead, NMSA C921; between Mpahlana and Umyameni Rivers, 318 08.49 S, 308 16.29 E, 100 m, sponge rubble, one dead, NMSA C5383; off Mtamvuna River, 318 09.79 S, 308 15.39 E, 120–140 m, sponge rubble, 1 live (body in alcohol), NMSA C436; off Mtamvuna River, 318 08.89 S, 308 16.09 E, 115 m, sponge, rocks, 1 live (body in alcohol), NMSA C483. Distribution.—Currently known only southeastern South Africa, from East London, Cape Province to Mtamvuna River, Transkei, living in 80 to 120 meters. Comparison.—Triviella immelmani most closely resembles T. khanya (Liltved, 1986). Triviella khanya differs in having a more globular shape, being more inflated anteriorly; a wider aperture; greater dorsal extent of the labral callus medially; weaker dentition on the fossula and columellar depression; in usually having 1– 3 of the posteriormost columellar ribs extending beyond the aperture towards the left posterior margin, rather than extending within the aperture to protrude as denticles on the inner columellar ridge and in having 3–4 of the anteriormost denticles wrapping partway onto the dorsal surface of the shell. The mantle of T.

khanya lacks the black spots seen in T. immelmani, but

as the animal has been observed in only one individual of each species, it is not known if this differences is constant. Etymology.—Named for Mr. Willie Immelman of Jeffreys Bay, South Africa, who collected the type specimens. Triviella Jousseaume, 1884a, is characterized by an inflated body whorl, which results in a wide to moderately wide aperture, blunt terminals that are not much produced, and ribbing absent or reduced posterior to the fossula and sometimes reduced or absent elsewhere on the shell. As a result of the reduction of ribbing, the columellar depression is also reduced or absent. Discussion.—The type species of Triviella is Cypraea oniscus Lamarck, 1810 [non Ro¨ding, 1798] 5 Triviella aperta (Swainson, 1822), by subsequent designation of Jousseaume (1884b: 99). Triviella is sometimes regarded as a nude name in Jousseaume (1884a) (e.g., by Schilder and Schilder, 1971: 90) and Cate (1979), but six available species name were included in the new genus, thereby making it available by indication. We herein synonymize Galeatrivia Cate, 1979 with Triviella. Cate placed two species in Galeatrivia, Cypraea ovulata Lamarck, 1811 (the type species by original designation) and G. millardi Cate, 1979. None of the characters that he used to distinguish this genus from Triviella do so consistently, as shown in Table 1. One character requires clarification: exposure of the spire whorls. As far as we have been able to determine, in all species of Triviidae the spire is covered with a layer of shell material, because the mantle envelopes the adult shell. The spire is exposed only in worn shells. Cate (1979: 15) stated, however, that in Galeatrivia the

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apex is not ‘‘overlaid with nacre,’’ and Liltved (2000: 162) stated of its type species that the spire is ‘‘not obscured by enamel.’’ These statements, though similar, are not equivalent: in shells of some species, the spire whorls can be seen through the overlying shell, which presumably is what Liltved meant. In some cases, the position of the suture of the spire is physically marked in the overlying shell as a depression or angulation. Confusion of this with the suture itself may have led some workers to incorrectly regard the spire as being exposed. Species of Triviella are known only from South Africa and Namibia. Twenty-three species are referrable to the genus, as listed below. Those that had not previously been placed in Triviella are indicated as new combinations (new comb.); all of these were originally described as Trivia, except Galeatrivia millardi. Triviella Triviella Triviella Triviella Triviella Triviella Triviella

aperta (Swainson, 1822) calvariola (Kilburn, 1980), new comb. costata (Gmelin, 1791) debruini (Lorenz, 1994), new comb. eratoides (Liltved, 1986), new comb. franziskae Fehse and Massier, 2000 goslineri (Liltved and Millard, 1994), new

comb. Triviella immelmani Rosenberg and Finley, new spe-

cies Triviella Triviella Triviella Triviella

khanya (Liltved, 1986), new comb. lemaitrei (Liltved, 1986), new comb. magnidentata (Liltved, 1986), new comb. massieri (Martin and Poppe, 1991), new

comb. Triviella Triviella Triviella Triviella Triviella Triviella Triviella Triviella Triviella

millardi (Cate, 1979), new comb. multicostata (Liltved, 1986), new comb. neglecta Schilder, 1930b ovulata (Lamarck, 1810) phalacra Schilder, 1930b rubra (Shaw, 1909) sanctispiritus (Shikama, 1974), new comb. splendidissima Tomlin and Schilder, 1934 verhoefi (Gosliner and Liltved, 1982), new

comb. Triviella vesicularis (Gaskoin, 1836) Triviella virginiae (Liltved, 1986), new comb.

Liltved (2000: 204–205) notes that T. massieri and T. verhoefi may prove to be synonyms of T. millardi, and that T. goslineri and T. debruini may be conspecific but that it is unknown which name has priority as they were published almost simultaneously in 1994. Other names referrable to Triviella may be found in the synonymies of species treated by Cate (1979) and Liltved (1989), except that the status of Triviella amaryllis Schilder, 1927 is unknown. Cate (1979) regarded it as a valid species, but Liltved (2000: 42) called it dubious; it may be an older name for T. millardi.

Trivia marlowi, n. sp. (Figs. 21–30) Description.—Shells ovoid, medium sized, from 9.8 to 12.3mm in length. Overall color whitish grey, with traces of pinkish brown spots dorsally. In live taken specimens gray areas would likely be off-white. Outer lip strong, thickest medially. Spire covered by several ribs, but visible through shell. Entire surface covered with smooth, flat-topped transverse ribs, 62–74 total, counted at periphery, often with more ribs on right than left side (32–39 vs. 30–36). Dorsal sulcus absent. Outer lip with 20–23 denticles protruding into aperture; outer columellar ridge with 21–25 denticles; columellar depression with 18–23 ribs; inner columellar ridge with 18–21 denticles, including 5–7 on fossula. Fossula excavated, slightly protruding. Aperture of constant width in posterior half, widening somewhat anteriorly. Right marginal callus, terminals and base with microscopic granulations between ribs. Type Locality.—Possession Island, Namibia (27.08 S, 15.28 E), on sand reef at 135 m. HOLOTYPE: L 12.2 mm, W 9.6 mm, H 7.7 mm, dead, June 1997, NMSA V5572/T1310. PARATYPES: Same data as holotype, all dead collected. 1) 12.2 3 9.2 3 7.9 mm, ANSP 406467; 2–6) L 9.8 to 12.3 mm, W 8.5 to 10.1 mm, H 6.7 to 8.3 mm, lost in registered mail being returned to Brian Hayes, Port Elizabeth, South Africa; 7) 11.3 3 10.0 3 7.5 mm, Finley Collection. Distribution.—Known only from the type locality. Comparison.—Trivia marlowi most closely resembles Trivia suavis (Schilder, 1931), a South African species. Trivia suavis differs in having a much lighter weight shell with less callus development on the right margin. Also, its aperture is straighter medially and more curved towards the terminals than in T. marlowi. Another related species is Trivia sharonae Hayes, 1993, which is more inflated, with a lighter weight, wider aperture, and fewer ribs. Etymology.—Named for Mr. Roy Marlow of St. Francis Bay, South Africa, who collected the type specimens. Discussion.—We classify this species in Trivia, on the assumption that its reproductive anatomy will be similar to that of Trivia suavis, which we regard as the most closely related species. Gosliner and Liltved (1987) showed that T. suavis has reproductive anatomy similar to that of Trivia arctica (Pulteney, 1799), a close relative of the type species of Trivia, Trivia europaea (Montagu, 1808). Shell morphology and coloration are consistent with placement in Trivia. Trivia marlowi is the third triviid recorded from Namibia, the others being Triviella eratoides and Triviella virginiae (Liltved 1986).

ACKNOWLEDGMENTS We thank Brian Hayes of Port Elizabeth, South Africa and Marty Beals of Inglewood, California for mak-

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Figs. 21–30. Trivia marlowi n. sp. 21–25, hairline fracture around dorsum.

HOLOTYPE,

length 12.2 mm. 26–30,

PARATYPE

#1, length 12.2 mm. Specimen has a

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ing specimens available for study; Sarah Watson, Academy of Natural Sciences of Philadelphia (ANSP), who photographed the specimens and prepared the plates. Richard N. Kilburn provided valuable comments on the manuscript and data from specimens in the Natal Museum, South Africa (NMSA). LITERATURE CITED Burgess, C. M. 1985. Cowries of the World. Seacomber Publications: Cape Town, South Africa. xvi 1299 pp. Cate, C. N. 1973. A systematic revision of the Recent cypraeid [sic] family Ovulidae (Mollusca: Gastropoda. Veliger 15, Supplement. iv 1 116 pp., 51 pls. Cate, C. N. 1979. A review of the Triviidae (Mollusca: Gastropoda). Memoirs of the San Diego Society of Natural History 10:1–126. Cox, L. R. 1960. General characteristics of Gastropoda. Pp. I84-I169 in R. C. Moore, ed., Treatise on Invertebrate Paleontology, Part I, Mollusca 1. Geological Society of America and University of Kansas Press. Dolin, L. 2001. Les Triviidae (Mollusca: Caenogastropoda) de l’Indo-Pacifique: Re´vision des genres Trivia, Dolichupis et Trivellona. In: P. Bouchet and B.A. Marshall (eds), Tropical Deep-Sea Benthos, volume 22. Memoires du Museum national d’Histoire naturelle, 185:201–241. Fehse, D. and W. Massier. 2000. A new Triviella (Gastropoda: Triviidae) from South Africa. La Conchiglia 32(294–295): 123–126. Gaskoin, J. S. 1836. [Several hitherto undescribed cowries.] Proceedings of the Zoological Society of London 3:198– 204. Gmelin, J. F. 1791. Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Ed. 13. Vol.1, part 6. Lipsiae. Pp. 3021–3910. Gosliner, T. M. and W. R. Liltved. 1982. Comparative morphology of three South African Triviidae (Gastropoda: Prosobranchia) with the description of a new species. Zoological Journal of the Linnean Society 74:111–132. Gosliner, T. M. and W. R. Liltved. 1987. Further studies on the morphology of the Triviidae (Gastropoda: Prosobranchia) with emphasis on species from southern Africa. Zoological Journal of the Linnean Society 90:207–254. Hayes, B. 1993. Trivia (Pseudotrivia) sharonae n. sp. (Gastropoda: Velutinoidea: Triviidae). La Conchiglia 25(268):6–7. Iredale, T. 1931. Australian molluscan notes. No. 1. Records of the Australian Museum 18:201–235, pls. 22–25. Jousseaume, [F.] 1884a. Division des Cypraeidae. Les Naturaliste 2(52):414–415. Jousseaume, F. 1884b. E´tude sur la Famille des Cypraeidae. Bulletin de la Socie´te´ Zoologique de France 9:81–100. Kilburn, R. N. 1980. Taxonomic studies on the marine Mol-

lusca of southern Africa and Mozambique. Part 2. Annals of the Natal Museum 24:193–200. Lamarck, [J. B.] 1810. Suite du genre Porcelaine. Annales du Muse´um d’Histoire Naturelle 16:89–114. Liltved, W. R. 1986. Six new species of Trivia from southern Africa (Gastropoda: Triviidae). Veliger 29:114–122. Liltved, W. R. 2000. Cowries and their relatives of southern Africa. 2nd ed. Seacomber Publications: Cape Town, South Africa. 224 pp. Liltved, W. R. and V. G. Millard. 1994. A new Trivia (Triviidae) and Prionovolva (Ovulidae) from Southern Africa. World Shells 11:3–8. Lorenz, F., Jr. 1994. From South Africa: Trivia debruini (Gastropoda, Triviidae). La Conchiglia 26(273):8–9. Martin, P. and G. T. Poppe. 1991. A new Trivia species from South Africa. La Conchiglia 22(259):2–3. Roberts, S. R. 1913. New Cypraeidae. Nautilus 26:97–99, pl. 7. Schepman, M. M. 1909. The Prosobranchia of the Siboga Expedition. Siboga-Expeditie 49b(2):109–231, pls. 10–16. Schilder, F. A. 1927. Revision der Cypraeacea (Moll. Gastr.). Archiv fu¨r Naturgeschichte 91A(10):1–171. Schilder, F. A. 1930a. Remarks on type specimens of some Recent Cypraeidae. Proceedings of the Malacological Society of London 19:49–58. Schilder, F. A. 1930b. Beitra¨ge zur Kenntnis der Cypraeacea (Moll. Gastr.)–III. Zoologischer Anzeiger 92:67–78. Schilder, F. A. 1931. Beitra¨ge zur Kenntnis der Cypraeacea (Moll. Gastr.)–IV. Zoologischer Anzeiger 96:65–72. Schilder, F. A. 1936a. Anatomical characters of the Cypraeacea which confirm the conchological characters. Proceedings of the Malacological Society of London 22:75–112, pls. 11– 12. Schilder, F. A. 1936b. Zwei interesante fossile Cypraeacea aus Indien (Mollusca Gastropoda). De Ingenieur Ned.-Indie¨ (4) 3(12):207–209 [not seen]. Schilder, F. A. 1941. Verwandtschaft und Verbreitung der Cypraeacea. Archiv fu¨r Molluskenkunde 73:57–120. Schilder, M. and F. A. Schilder. 1971. A catalogue of living and fossil cowries: Taxonomy and bibliography of Triviacea and Cypraeacea (Gastropoda Prosobranchia). Institut royale des sciences naturelles de Belgique Memoires (2) 85:1–246. Shaw, H. O. N. 1909. Notes on the genera Cypraea and Trivia. Proceedings of the Malacological Society of London 8: 288–313, pls. 12–13. Shikama, T. 1974. On two new allied cowry and cowry shells from South African Sea. Science Reports of the Yokohoma National University (2) 21:23–26. Sowerby, G. B., Jun. 1839. A conchological manual. G. B. Sowerby: London. [ii] 1 v 1 [i] 1 130 pp., [24] pls. Swainson, W. 1822. A catalogue of the rare and valuable shells, which formed the celebrated collection of the late Mrs. Bligh. London. iv 1 58 1 20 pp., 2 pls. Tomlin, J. R. le Brockton and F. A. Schilder. 1934. Reports on the Marine Mollusca in the Collections of the South African Museum. IX. Family Triviidae. Annals of the South African Museum 30:477–479.