= : HSM | à DIS , RSA | | NASA TRS | DECO EE ES el PT : L'ÉTAT ad | | ME, re 5 : : F ; | f 2 HAN TOUS L : ; F : . ; û no fn 3 : ?. ; Ê : s ë, WE ; | D 21 Vie ads, CRT " se. “ ROUES | ÿ res e è Ke RÉAL TE TLC PE id Cl ; Lai LA “ . Dr ‘te HANREE ; AÉTE LA . b À CRTC TEE pis HARVARD UNIVERSITY t Library of the Museum of Comparative Zoology : r1 ( : (n | FYRLES LS 10 MA here Lie P2 + MAN RSS 4 AA US LE . j { | Î M . r (re se Hs MCZ LIBRARY Société Belge de Malacologie association sans but lucratif AVRIL 1994 SOMMAIRE A. Panigrahi Effect of temperature on noradrenaline and adrenaline S.K. Mahata content in the brain of a terrestrial slug, Laevicaulis alte S.K. Raut (Férussac) (Gastropoda : Soleolifera) R.G. Moolenbeek The Orbitestellidae (Gastropoda: Heterobranchia) of the Sultanate of Oman with description of a new genus and two new species R. Duchamps À note on the Museum Leskeanum B. Tursch E. Rolän The Family Triphoridae (Mollusca, Gastropoda) in R.Fernändez-Garcés Cuba. 4. The genera Monophorus, Nototriphora, Cosmotriphora and Cheirodonta, with the description of three new species Périodique trimestriel Bureau de dépôt 1180 Bruxelles 18. R. Duchamps Dr. Y. Finet L. Germain R, Houart Dr. CI. Massin Prof. B. Tursch Dr. J. Van Goethem Editeur responsable : Comité d'édition : Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s), et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. All rights of reproduction are reserved without the written permission of the board. Belgique - Belgium Etranger - Foreign Abonnement aux revues APEX & ARION Subscription to APEX & ARION [avec le service des bulletins) Membre effectif 2.02 ROUBIBEr A … 1400 BEF Membre étudiant 52 SOOIBER : Versement à effectuer par mandat postal international ou Lonsle sav ere luEie ar Se bancaire nie ad uni rh ee ge e international money order, or by bank chec Personne appartenant à la famille d'un membre effectif in Belgian Francs only. et ayant la même résidence . 400 BEF au nom de Versements à effectuer au C.C.P. n° 000097422554 de MORE la Société Belge de Malacologie c/o M. J. Buyle, Av. M. Maeterlinck, 56, 1030 Bruxelles. Av. Maurice Maeterlinck, 56, bte 8 B-1030 Bruxelles. CONSEIL D'ADMINISTRATION DE LA SOCIETE BELGE DE MALACOLOGIE e Président : MR. Duchamps, Av. Mozart, 52, 1 190 Bruxelles © : (02) 344.15.47 Dr. Y. Finet, 16 Chemin des Clochettes, CH-1206, Genève (Suisse) © : 41-22-46.77.95 M.R. Houart, St. Jobsstraat, 8, 3400 Landen ([Ezemaal) ® : (016) 78.86.16 Mme J. Masson, Rue du Merlo, 10, 1180 Bruxelles © : (02) 376.62.25 ° Trésorier : M. J. Buyle, Av. Maeterlinck, 56, bte 8, 1030 Bruxelles ® : (02] 216.68.21 ° Bibliothécaire : Mme ML Buyle, Av. Maeterlinck, 56, bte 8, 1030 Bruxelles © : (02) 216.68.21 M.G. Geeraerts, Dorpsstraat, 125, 3078 Meerbeek G :(02)757.07.47 Mme ML. Bresson, Place Guy d'Arezzo, 7, 1060 Bruxelles ® : (02) 343.62.38 M. L. Germain, Rue de Linthout, 77, 1040 Bruxelles © : (02) 734.80.11 Mme À. Langleit, Av. Cicéron, 27, bte 92, 1 140 Bruxelles ® : (02) 720.416] M. C. Van Osselaer, Chée de Waterloo, 512, 1060 Bruxelles ® : (02) 347.36.84 M. E. Waiengnier, Rue C. Wolles, 42, 1030 Bruxelles ® : (02) 241.51.80 e Vice-présidents e Secrétaire + Relations publiques + Administrateurs Panigrahi, Mahata & Raut Laevicaulis alte APEX 9(1): 1-4, April 1994 Effect of temperature on noradrenaline and adrenaline content in the brain of à terrestrial slug, Laevicaulis alte (Férussac) (Gastropoda : Soleolifera) À. PANIGRAHI, S.K. MAHATA" and S.K. RAUT Department of Zoology, University of Calcutta, 35 Ballygunge Circular Road, Calcutta 700 019, India *San Diego V.A. Medical Centre, Division of Nephrology - Hypertension (V-111-H), UCSD, 3350 La Jolla Village Drive, San Diego, CA 92161, USA KEY WORDS: Slug, brain, noradrenaline, adrenaline, temperature. ABSTRACT: The garden slugs Laevicaulis alte (Férussac) were exposed to 10°, 15°, 20°, 25°, 30° and 35°C constant temperatures to study the changes in noradrenaline and adrenaline contents in the brain. Depletions in the amount of noradrenaline at 10° and 20°C, and elevations in the amount of both noradrenaline and adrenaline have been noted at 15° and 30°C. Both noradrenaline and adrenaline contents at 25°C were found almost equal. No change in adrenaline content was noted at 10° and 20°C while the amount was appreciably high at 35°C. INTRODUCTION The garden slugs ZLaevicaulis alte (Férussac) have a wide range of temperature tolerance (GODAN, 1983; RAUT and MANDAL, 1984). They are found active in monsoon at à temperature range 14° - 35°C. In winter when the temperature falls below 12°C, they hibernate to overcome the temperature-induced environmental hazards. Though the behavioural changes are well marked, the factors influencing such changes are still not known, although the neuroendocrine system of the animal is generally presumed to be involved. The present experiments therefore examined variations in amount of noradrenaline and adrenaline in the brain of Z. alte (Férussac) under different fixed temperatures. MATERIALS AND METHODS. Gravid ZLaevicaulis alte (Férussac) were collected from their natural habitat at Sandeshkhali, 24-Parganas (North), West Bengal, India. They were reared under laboratory conditions (at room temperatures 21.6° - 30.4°C) following the method described by RAUT (1991) to obtain the slug individuals as per requirement of the study. A total of 90 laboratory-reared, adult, healthy individuals with 57-60 mm in body length, 19-21 mm in body breadth and 3.27-3.62 g in body weight were divided into 6 groups. Each group was housed separately in a terrarium measuring 30 x 20 x 15 cm, provided with loose, moist soil up to 6 cm of its height. Each of the 6 terraria was exposed to any of the constant 10°, 15°, 20°, 25°, 30° and 35°C temperature grades maintained in 6 separate Biological Oxygen Demand (B.O.D.) incubators (S.N. Mullick, Calcutta). (The atmospheric temperature of Sandeshkhali area ranges from 10° - 35°C). The moisture of the soil was kept between 37 - 40 % in all terraria throughout the experiment by spraying water at a regular interval. A favourable humidity range (80 - 90 %) was also maintained inside the chambers throughout in the same way. Also the slugs were exposed to light (400 Lux) provided by a 15 W. bulb for a period of 12 hours. The slugs were supplied with lettuce (Zactuca sativa Linnaeus) and beans (Zablab purpureus Linnaeus) regularly in excess as their food. Strict hygienic condition was maintained throughout by removing the unconsumed food matters and the faecal pellets. The experiment was continued for a period on one month. For the collection of brain (including all associated ganglionic parts), in each case, the slugs (5 individuals) were anaesthetized by applving chloroform prior to dissection. The materials were then processed for the spectrophotofluorometric determinations of noradrenaline and adrenaline following Cox and PERHACH (1973) and LAVERTY and TAYLOR (1968). The estimation of the monoamines present in the samples was made by the help of a Hitachi (Model 650-10 M) Fluorescence Spectrophotometer (for details see MAHATA and GROSH, 1989). The experiments were repeated thrice and the mean of three readings of pooled observations was considered as the actual amount of noradrenaline and adrenaline present APEX 91): 1-4, April 1994 in the brain. Statistical analysis of the data was performed following GOON et al. (1976). RESULTS The slugs survived for a period of 9-15 days at 10°C, 24-29 days at 15°C and 6-9 days at 35°C temperatures. In the remaining groups, none died during the period of 30 days. The autopsy for the collection of brain was done on the 9th, 24th and 6th day of exposure of slugs maintained at 10°, 15°and 35°C, respectively, as the death was started on these days. In these cases, five individuals were considered for autopsy. The remaining individuals (except those died on that day) were maintained as usual, at these temperatures till the death of all individuals. The average weight of the brain was 4.05 mg. Noradrenaline and adrenaline content recorded from the brain of Z. alte (Férussac) maintained under different temperature grades have been shown in Table 1. To justify the validity of the data obtained in respect to different temperatures, the two-way fixed effect homoscedastic Analysis of Variance (ANOVA) was applied (Table 2). From the results, 1t is clear that there exists a significant difference at 1 % level among noradrenaline and adrenaline in Z. alte in respect to different temperatures. Also, the degree of interaction between monoamine and temperature 1s significant throughout. To study the maximum significant effect of monoamine (noradrenaline or adrenaline), temperature and interaction due to monoamine and temperature, we adopt Student's ‘t' test through pair comparison. In general, for (ij)th pair, if absolute difference of means of (ij)th pair is greater than l Il ta /2,4f VMSE = +— ni nj; (called least significance difference (1s.d.), then, (ij)th pair is significant at «% level of significance. In our case, L.s.d. for effect of monoamine 1s : ta /2,df VMSEY2/3 =2.4940.1533V2/18 =0.3250 at 1% level of significance. Similarly, 1.s.d. for temperature is 249Y0.1533Y2/6 = 0.5629 and Is.d. for interaction between effect of monoamine and temperature 1s 249Y0.15334Y2/3 = 0.7960 It is evident that adrenaline and 35°C have maximum significant effect (Table 1). 2 Laevicaulis alte Panigrahi, Mahata & Raut DISCUSSION Ample evidences demonstrate the presence of monoamines in the central nervous system for vertebrates (TURNER and BAGNARA, 1976; LANDSBERG and YOUNG, 1985). In molluscs, catecholamine containing cell bodies have beeen demonstrated in Zimax maximus Linnaeus (OSBORNE and COTTRELL, 1971), Aplysia (GOLDSTEIN, 1984), Helisoma (TRIMBLE et al, 1984), Helicella virgata DaCosta (FRANCHINI et al., 1985) and Zymnaea stagnalis Linnaeus (AUDESIRK, 1985). The distribution of monoamines in the central nervous system of gastropods, Æermissenda crassicornis Eschscholtz and Achatina fulica Bowdich has been described by CROLL (1987a, b, 1988) VON EULER (1953) reported the presence of noradrenaline in Octopus. McCAMAN et al. (1979a, b, 1984) were successful in isolating dopamine, S- hydroxytryptamine and N-acetyldopamine from molluscan ganglia. GERSCHENFELD (1973) and LEAKE and WALKER (1980) are in opinion that dopamine 1s the major catecholamine of the snail Æelix. Though OSBORNE and COTTRELL (1970) and JUORIO and KILLICK (1972) noted the presence of significant amount of noradrenaline in the brain of Felix and some other molluscs quantitative data of the amount of noradrenaline and adrenaline in molluscs, in the true sense, was not available until the publication of OSBORNE's (1984) work on CNS of Helix aspersa Müller. Subsequently, COON and BONAR (1986) were also successful in quantifying the amount of norepinephrine and dopamine in the larval and spat stages of the Pacific Ovyster Crassostrea gigas (Thunberg). Recently, findings by PANIGRAHI et al. (1992) not only demonstrate the amount of noradrenaline and adrenaline in the brain of the slugs Z. alte (Férussac) but also provide information on fluctuations 1n the amount of these monoamines at different hours of a day. However, none of the previous workers has paid attention to study the effect of temperature on monoamine content in the brain of any mollusc. On the contrary, such à study is on record in some vertebrates - reptiles (REITER, 1981), birds (HARVEY et al., 1984) and mammals (TURNER and BAGNARA, 1976, LANDSBERG and YOUNG., 1985). The present findings clearly indicate that temperature has great influence on noradrenaline and adrenaline content in the brain of the slugs Z. alte. Variations in the amount of noradrenaline and adrenaline recorded in the brains of Z. alte in respect to temperatures seem to be related with the functional status of the enzymes involved with the process of synthesis of these amines. This Panigrahi, Mahata & Raut phenomenon has been discussed by OSBORNE (1984) in case of the gastropod snail #. aspersa. Whatever be the reasons for such changes in amine contents, it seems that the phenomenon is involved with the steps of adjustment of the concerned slugs in respect to the conditions evolved due to change of temperatures. Though further experimental studies are needed to evaluate the role of temperature in influencing the rate of release of monoamines it is sure that the said factor constitutes a potential source of interference. Acknowledgments Authors are grateful to Dr. Asok Ghosh, Sir Nilratan Sircar Professor of Zoology and to the Head of the Department of Zoology, University of Calcutta for the facilities provided. The financial assistance from the Indian Council of Agricultural Research, New Delhi is thankfully acknowledged. REFERENCES AUDESIRK, (G. 1985. Amine-containing neurons in the brain of Lymnaea stagnalis : distribution and effects of precursors. Comp. Biochem. Physiol., 81(A) : 359-365. CooN, SL. and D.B. BONAR 1986. Norepinephrine and dopamine content of larvae and spat of the Pacific Oyster, Crassostrea gigas. Biol. Bull., 171(3) : 632-639. COX RH Jr and: IE. Jr. PERHACH 1973. A sensitive, rapid and simple method for the simultaneous spectrophotofluorometric determinations of norepinephrine, dopamine, 5- hydroxytryptamine and 5-hydroxyindoleacetic acid in discrete areas of brain. J. Neurochem., 20: 1777-1780. CROP, RP. 1987a: Distribution of monoamines in the central nervous system of the nudibranch gastropod, Æermissenda crassicornis. Brain Res., 405 : 337-347. CEROEL RP. _1937b. Distribution of monoamines in the nervous system of the hatching snail, Achatina fulica. Bull. Can. Soc. 20011823. CROLL, RP. 1988. Distribution of monoamines within the central nervous system of the juvenile pulmonate snail, Achatina fulica. Brain Res., 460: 29-40. FRANCHINI, A, E. OTTAVIANI and E. CASELGRANDI 1985. Biogenic amines in the snail brain of Hellicella virgata (Gastropoda, Pulmonata). Brain Res., 347 : 132-134. GERSCHENFELD, HM. 1973. Chemical transmission in invertebrate central nervous system and neuromuscular junctions. Physiol. Rev., 53 : 1-119. Laevicaulis alte APEX 9(1): 1-4, April 1994 GODAN, D. 1983. Pest slugs and snails, Springer-Verlag, Berlin, Heidelberg, New York. vi + 445 pp. GOLDSTEIN, R.S., 1984. Immunocytochemical, histofluorescent and ultrastructural studies of monoaminergic neurons and their processes in Aplysia. Ph. D. thesis, Columbia University, New York. GoOoN, A.M., MK. GUPTA and B. DASGUPTA 1976. Fundamentals of Statistics. Vol. 2, World Press, Calcutta. xin + 431 pp. HARVEY, S., J.G. PHILLIPS, A. REES and TR. HALL 1984. Stress and adrenal function. J. Exp. Zool., 232 : 633-645. JUORIO AN an TS WRI LICRLO 72: Monoamines and their metabolism in some molluscs. Comp. Gen. Pharmacol., 3 : 283- 295. LANDSBERG, L. and J.B. YOUNG 1985. Catecholamines and the adrenal medulla. In "Williams Textbook of Endocrinology" (JD. Wilson and D.W. Foster, Eds.). Saunders and Igaku-Shoin/Saunders. Philadelphia/Tokyo. pp. 891-965. LAVERTY, R. and K.M. TAYLOR 1968. The fluorometric assay of catecholamines and related compounds Improvements and extensions to the hydroxyindole technique. Anal. Biochem., 23 : 269-279. LEAKE, LD. and RJ. WALKER 1980. Invertebrate neuropharmacology, Blackie, Glasgow. MAHATA, S.K. and A. GHOSH 1989. Influence of splanchnic nerve on reserpine action in avian adrenal medulla. Gen. Comp. Endocrinol., 73 : 165-172. McCAMAN, MW. JK. ONO and RE. McCAMAN 1979a. Dopamine measurements in mollluscan ganglia and neurones using a new sensitive technique. J. Neurochem., 32 : 1111- 111S McCAMAN, M. W.; RE. McCAMAN and J. STETZLER 1979b. A rapid radioenzymatic assay for dopamine and N-acetyldopamine. Anal. Biochem., 96 : 175-180. McCAMAN, MW. JK. ONO and RE. McCAMAN 1984. 5-hydroxytryptamine measurements in molluscan ganglia and neurones using a modified radioenzymatic assay. J. Neurochem.., 43 : 91-99. OSBORNE, NN. 1984. Phenylethanolamine-N- methyltransferase and dopamine-B-hydroxylase immunoreactivity and the occurrence of noradrenaline and adrenaline in the nervous system of the snail Felix aspersa. Cell Tissue Res., 237 : 605-608. APEX 9(1): 1-4, April 1994 OSBORNE, NN. and GA. COTTRELL 1970. Occurrence of noradrenaline and metabolites of primary catecholamines in the brain and heart of Helix. Comp. Gen. Pharmacol., 1 : 1-10. OSBORNE, N.N. and G.A. COTTRELL 1971. Distribution of biogenic amines in the slug, Limax maximus. Z. Zellforsch., 112 : 15-30. PANIGRAHI, À., S.K. MAHATA and S.K. RAUT 1992. Circadian rhythm in norepinephrine and epinephrine contents in the brain of the garden slug, Laevicaulis alte (Férussac). APEX., 7(2) : 59-65. RAUT S.K. 1991. Laboratory rearing of medically important molluscs. In: "Snails, flukes and man" (Director, Zoological Survey of India, Ed.), pp. 79-83. Calcutta. Table 1. Laevicaulis alte Panigrahi, Mahata & Raut RAUT, S.K. and R.N. MANDAL 1984. Natural history of the garden slug Zaevicaulis alte. J. Beng. Nat. Hist. Soc., 3 : 104-105. REITER, R.J. 1981. The Pineal. Vol. 6, Eden Press, 265 pp. TRIMBLE, D.L., D.L. BAKER and B.J. BULLARD 1984. Dopamine in a molluscan nervous system: synthesis and fluorescence histochemistry. J. Neurobiol., 15 : 27-36. TURNER, C.D. and J.T. BAGNARA 1976. The adrenal medulla : Chromaffin tissue. In "General Endocrinology", 6th edn., pp. 291- 323. W.B. Saunders Company, Philadelphia, London, Toronto. VON EULER /"U:S’, 1953; Presence of catecholamines in visceral organs of fish and invertebrates. Acta Physiol. Scand., 28 : 297- 305. Noradrenaline and adrenaline content (average + S.E.) in the brain of the garden slugs Laevicaulis alte maintained under different temperature grades for a different length of time (not more than 30 days in any case) depending upon their withstanding capacity under such conditions [Remarks refer to the concentrations of noradrenaline and adrenaline at the same temperature]. Temperature (°C) Monoamine (ug/£ tissue + S.E) t-value Remarks Noradrenaline Adrenaline 646021 9.62 + 0.24 10.514 (p=0.01) 14.58 +0.29 11.34 + 0.25 8.510 (p=0.01) 232-204 9.45 + 0.32 21.221 (p=0.01) 9PYÆUMTS 941+0.11 0.623 10.16 + 0.26 10.28 + 0.22 0.208 10125075 1931007 35.868 (p=0.01) * The slugs survived for a period of 9-15 days. se The slugs survived for a period of 24-29 days. 1 The slugs survived for a period of 6-9 days. S = Significant, NS = Not significant. Table 2. Analysis of Variance (ANOVA) to justify the effect of monoamines (noradrenaline and adrenaline in respect to different temperature grades in the brain of the garden slug Laevicaulis alte. Source of variation df SS MS FCal Ftap (@ = 1%) Effect of monoamines (E) Il 64.7220 Temperature (T) 5 305.9037 Interaction (E x T) 5 1633672 Error 24 3.6797 64.722 422.131 787 61.1807 399.034 3.9 3216734 20213"105 29 0.1533 > = 25 537.6726 Moolenbeek Orbitestellidae of Oman The Orbitestellidae (Gastropoda: Heterobranchia) of the Sultanate of Oman with description of a new genus and two new species* R.G. MOOLENBEEXK. Zoôlogisch Museum Amsterdam, P.O. Box 4766, NL-1009 GT Amsterdam, The Netherlands ABSTRACT. Three species of the family Orbitestellidae are recognized from Oman . The first, Orbitestella bermudezi (Aguayo & Borro, 1946) was previously known from the tropical Atlantic Ocean. It is also recorded from the Red Sea. The two remaining species are new to science and require a new genus. Boschitestella nov. gen. with B. donaldi nov. sp., known from Oman, Red Sea, Thailand and Indonesia and B. eloiseae nov. sp. only known from Oman are assigned to this new genus. KEY WORDS: Gastropoda, Orbitestellidae, Orbitestella , Boschitestella nov. gen, Indian Ocean, Oman. INTRODUCTION Because of their minute size, species of the family Orbitestellidae have been overlooked in most marine faunas. However, during recent years much progress has been made in the understanding of these very small gastropods. PONDER (1990) showed that the orbitestellids are primitive heterobranchs. Up to-now most species were recorded from the southern hemisphere (especially [sub]Antarctica, Australia, and New Zealand) but recently some recordings have been made from the northern hemisphere. KAY (1979) described a new species from Hawaï. FABER (1991) and ROLAN & RUBIO (1992) recognized their existence in the western and mid Atlantic Ocean. During fieldwork along the coastline of the Sultanate of Oman in November 1991, a limited number of these microgastropods were found in the intertidal zone, beneath rocks and among coralline algae. In our samples two types with different characters are distinguished. One with the typical outline of the genus Orbitestella Iredale, 1917 and shells of the other group are distinguished by only one single, sharp carina on the periphery of the teleoconch. These are considered here to belong to a new genus. *Studies on the marine molluscan fauna of Oman, n°. 9. TAXONOMY Orbitestella bermudezi (Aguayo & Borro, 1946) Figs 1-8 Cyclostremiscus bermudezi Aguayo & Borro, 1946: 9-12 Orbitestella similis Rolân & Rubio, 1992: 17-18 Orbitestella cubana Rolän & Rubio, 1992: 18- 19 Distribution. West Indies, Cape Verde Islands; Red Sea (Gulf of Aqaba), Oman (Al Hallaniyah, Sta. 91/60, 5 specs: Masirah Island, BERS camp, Sta. 91/95, 1 spec.; Ra's al Ya, Sta, 91/105, 1 spec., all November 1991, leg. R.G. Moolenbeek & H. Dekker). Remarks. FABER (1991) recorded ©. bermudezi , originally described as a Cuban fossil of Miocene or Pleistocene age, being an extant species known from several West Indian islands. ROLAN & RUBIO (1992) described two new species respectively ©. similis from the Cape Verde Islands and ©. cubana from Cuba. If the figure numbers in ROLAN & RUBIO (1992) are correct (and not reversed) it 1s obvious that the figured specimen of ©. similis 1s nearly identical to Faber's figured specimen from the West Indies. APEX 9(1): 5-10, April 1994 APEX 9(1): 5-10, April 1994 After comparing specimens from several West Indian islands it is my opinion that both taxa from ROLAN & RUBIO (1992) fall within the variability of ©. bermudezi. . Furthermore, their identical protoconchs support this view. More remarkable is the occurrence of ©. bermudezi in the northern Indian Ocean. Specimens from Oman (Figs 1-3) have the same type of protoconch with the minute granulations and a mid dorsal ridge (Figs 4-5). Also the sculpture of the teleoconch falls within the variation known in ©. bermudezi. The only difference noticed is the height of the two peripheral spiral ridges. It appears slightly smaller on the figured specimen but varies in the Oman population. In a small grunge sample from the Red Sea, Gulf of Aqaba (Nuweiba). collected by Mr. C. Steenman in October 1992 this species was rather common (Figs 6-8). Again, the specimens show variability in shell morphology but are still considered to be identical to ©. bermudezi. This exceptional large distribution pattern may be explained by its likely Tethyan origin. TURTON (1932) described Æomalogyra gemmulata from South Africa. This species 1s very similar to Orbitestella bermudezi, however his figures and description are too vague to give a definitive opinion. À study of the type material (Oxford University Museum, U.K.) and additional material from the Natal Museum (Pietermaritzburg, South Africa) has been initiated. Also Cyclostrema bastowi Gatliff, 1906, the type species of the genus Orbitestella might be identical and would be the oldest name for this taxon. Boschitestella nov. gen. Type Species: Boschitestella donaldi nov. Sp. Diagnosis. Shell minute, discoidal, width 0.5-0.9 mm, height 02-03 mm, widely umbilicate, translucent white , with numerous fine spiral threads, which consist of small granules (at high magnification), weak axial ribs or knobs and one spiral ridge on the periphery. Etymology. The genus and the type species were named in honour of Dr. Donald Bosch, who stimulated my interest in the marine molluscs of Oman. Distribution. Red Sea, Thailand and Indonesia. In Oman, from the Muscat area and from Al Hallaniyah (=Kuria Muria Islands). Orbitestellidae of Oman Remarks. The genus 1s based on differences in shell morphology between species of this genus and species of the genus Orbitestella Iredale, 1917. The type of this genus is Cyclostrema bastowi Gatliff, 1906, described from Victoria, Australia. Boschitestella differs by its single sharp carina on the periphery, being larger and a different structure of its protoconch. Radula and soft parts are unknown. The genus Vitrinorbis Pilsbry & Olsson, 1952 from the Panamic province has superficial resemblance to Boschitestella . However, it differs in being twice as large and having a different sculpture of fine spiral threads, cut into fine beads by close, retractively radial grooves. Boschitestella donaldi nov. sp. Figs 9-13 Description of the holotype (ZMA Moll. 3.93.002; width 0.86 mm, height 0.24 mm). Shell minute, translucent white, discoidal with a flat spire, widely umbilicate, strong axial plicae on the periphery and numerous fine spiral threads. Protoconch I of 0.6 whorl (diameter 0.09mm), initial part with a hexagonal, crateriform structure gradually becoming smooth and terminated by a varix. Apex slightly inrolled. Protoconch II of 0.5 whorl, smooth, terminated by a distinct varix (diameter Pc I + Pc II: 0.17 mm). Colour light tan. Teleoconch of approximately 1.8 whorls with numerous fine spiral threads on upper side and rather strong knobs on the mid part. Well developed plicae on the periphery and on the base the spiral threads diminish. Base with about 13 spiral rows gradually disappearing towards the protoconch and approximately 38 axial ribs on the body whorl. Type Locality. Sultanate of Oman, Haramal near Muscat. Sta. 91/83. low tide. in tidal pools with rocks, 28.11.1991, leg. R.G. Moolenbeek & H. Dekker. Variability. Two paratypes (ZMA Moll 3.93.003) from the type locality, conform favorably in all sculpture details with the holotype. Paratype 1 (fig. 11) width 0.76 mm: paratype 2, a subadult specimen (fig. 10) width 0.58 mm. Other Material Studied OMAN: Al Bustan near Muscat, Sta. 91/51, XI.1991, leg. R.G. Moolenbeek & H. Dekker ( 1 spec.). RED SEA: EGYPT, Gulf of Aqaba, Nuweiba, X.1992, leg. (C. Steenman (4 specs). INDONESIA: NE. Sumbawa, Bima Island, 10 m, IX.1987, leg. J. Veth (10 specs);, Bunaken Moolenbeek Moolenbeek Island, 20 m, VIL.1988, leg. J. Veth (1 spec.). THAILAND: Phuket Island, Patong Beach. XIL.1983, leg. J. de Visser, coll. T. Keukelaar- Van den Berge (2 specs). Remarks. Boschitestella donaldi nov. sp. differs from Orbitestella bermudezi by having (probably) planktotrophic larval development and only one ridge on the periphery. It differs from Boschitestella eloiseae nov. sp. by having planktotrophic larval development, and fewer spiral threads on the bodywhorl. Boschitestella eloiseae nov. sp. Figs 14-16 Description of the holotype (ZMA Moll. 3.93.003; width 0.55 mm, height 02 mm, subadult specimen). Shell minute, translucent white, discoidal with flat spire. widely umbilicate, small axial plicae on periphery and numerous fine spiral threads. Protoconch of 1.3 whorls (diameter 0.18 mm), initial part with a hexagonal, crateriform structure gradually becoming smooth and terminated by a varix. Colour white. Teleoconch of about 1.3 whorls with many fine spiral threads on upper side (about 25) and rather strong axial ribs (about 21). Under side with 25 fine spiral threads of which only about 5 are visible near the protoconch and about 38 axial ribs. Type Locality. Sultanate of Oman. Al Hallaniyah (=Kuria Muria Islands) Sta. 91/60. tidal pools with rocks, 12.11.1991, leg. RG. Moolenbeek & H. Dekker. Variability. One paratype ( 0.5 mm width: 0.2 mm height) from Masirah Island, Ra's Abu Rasas, tidal pools, Sta. 91/90, 19 November 1991, leg. R.G. Moolenbeek & H. Dekker, closely resembling the holotype. Remarks. A single species comparable to this taxon is Boschitestella donaldi nov. sp. However, it can be distinghuished by its different type of protoconch, indicating a non- planktotrophic larval development. ACKNOWLEDGEMENTS. I express my gratitude to Dr. Donald Bosch, who invited us to participate in the Oman expedition and to Mrs Eloise Bosch for hospitality during our stay in Muscat. Thanks are due to the staff of the BERS station on Masirah Island for accomodation and support. Peter and Una Dance, Christine and Valter Hägstrom, Donald and Eloise Bosch and Henk Dekker were companions during our collecting activities and always helpful. Without the enthusiastic support of RH. de Bruyne, S. Dekker. J. Hoenselaar. Orbitestellidae of Oman APEX 9(1): 5-10, April 1994 and T. Keukelaar-Van den Berge in sorting out most sediment samples, this study could not be achieved. M. J. Faber, J. de Visser, T. Keukelaar-Van den Berge and C. Steenman kindly collected and donated sediment samples to our Museum or presented their private collection for study purposes. Dr. G. Rosenberg (Academy of Natural Sciences of Philadelphia) kindly send in loan the type species of the genus Vitrinorbis. KLM Oman (Mr J.W. Creutzberg and J. Simpson) kindly arranged a courtesy air ticket from Amsterdam to Seeb. Dr. HE. Coomans, M.J. Faber and two unknown referees gave valuable suggestions and Mrs K. Kaiser corrected the English text. SEM photos were made at the Laboratorium voor Elcktronenmikroskopie (University of Amsterdam) by the author. APPENDIX: In the final stage of this paper, a congress lecture was published in Venus 51(1- 2), 1992: 133-135 by S. Kaneko. He figured two unidentified orbitestellids from Japan which definitely belong to the genus Boschitestella. Whether these are conspecific with the Oman species needs further research. APEX 9(1): 5-10, April 1994 REFERENCES AGUAYO, C.G. & P. BORRO. 1946. Nuevos moluscos del Terciario Superior de Cuba. Rev. Soc. Malac . 4: 9-12. FABER, M.J. 1991. Cyclostremiscus bermudezi : first record of a recent member of the Orbitestellidae (Gastropoda, Prosobranchia) in the Caribbean. APEX 6(3-4): 77-79. KAY, E.A. 1979. Hawaïian marine shells, reef and shore fauna of Hawaun. Bernice P. Bishop Museum, Honolulu: 1-653. Orbitestellidae of Oman PONDER, W.F. 1990. The anatomy and relationship of the Orbitestellidae (Gastropoda: Heterobranchia). J. Moll. Stud . 56: 515-532. ROLAN, E. & F. RUBIO. 1992. Two new species of the genus Orbitestella Iredale, 1917 from the Atlantic Ocean. La Conchiglia 23(262): 17-20. TURTON, W.H. 1932. The marine shells of Port Alfred, South Africa. Oxford University Press, London: 1-331. Figs 1-8. (opposite page) Orbitestella bermudezi (Aguayo & Borro, 1946). 1-5. Oman, AI Hallaniyah, Sta. 91/60. 1. Dorsal view, width 0.50 mm. 2. Lateral view, height 0.2 mm. 3. Ventral view, width 0.60 mm. 4-5. Detail of protoconch, dorsal and ventral view. 6-8. Red Sea, Nuweiba, X.1992. 6. Dorsal view, width 0.55 mm. 7. Lateral view, height 0.2 mm. 8. Ventral view, width 0.55 mm. Moolenbeek Moolenbeek Orbitestellidae of Oman APEX 9(1): 5-10, April 1994 APEX 9(1): 5-10, April 1994 Orbitestellidae of Oman Moolenbeek — 23.0 jun : D 23.0 ym on Figs 9-13. Boschitestella donaldi n.sp., Oman, Haramal, Sta. 91/83. 9. Dorsal view of paratype 1, Width 0.76 mm.10. Ventral view of paratype 2, Width 0.58 mm. 11. Lateral view of holotype, height 0.26 mm. 12-13. Details of protoconch, dorsal and ventral views of paratype 1. Figs. 14-16. Boschitestella eloiseae n.sp., Oman, Al Hallaniyah, Sta. 91/60, holotype. 14. Dorsal view, width 0.55 mm. 15. Details of protoconch. 16. Lateral view, height 0.20 mm. 10 Duchamps & Tursch Note on the Museum Leskeanum APEX 9(1): 11-16, April 1994 A note on the Museum Leskeanum R. DUCHAMPS (°) and B. TURSCH Laboratoire de Bio-Ecologie, Faculté des Sciences, Université Libre de Bruxelles, 50 av. F.D. Roosevelt, B-1050 Brussels, Belgium. (°) Chercheur associé ABSTRACT. The Museum Leskeanum of D.L.G. Karsten (1789) should be considered as an available work, in the meaning of the International Code of Zoological Nomenclature. RESUME. Le Museum Leskeanum de D.L.G. Karsten (1789) doit être considéré comme un travail disponible, dans le sens du Code International de Nomenclature Zoologique. KEY WORDS: Zoological nomenclature, Karsten, available work, Museum Leskeanum. The names in Karsten's Museum he was called by the minister Heynitz. Karsten Leskeanum are generally dismissed in the malacological literature as being "non- binominal". Careful re-examination of this work shows that this opinion should be reversed. Dietrich Ludwig Gustav KARSTEN was born in Bützow (Mecklenburg) on April 5, 1768 in a family of great scientific achievers. His father, Wenceslas J.G. KARSTEN (1732-1787) was one of the foremost German mathematicians of the 18th century, the author of many books and a professor at several universities. His uncle Franz C.L. KARSTEN (1751-1829) was a prominent agronomist, professor at Iena. His cousin Karl JB. KARSTEN (1782-1853) was a mineralogist. author of many works. and ended as chief adviser for the mines in Berlin. Albeit rarely cited in malacology, Dietrich Ludwig Gustav KARSTEN is very well known in mineralogy, a field of which he is considered to be one of the most important founders. He first studied mathematics and physics with his father, then enroled in 1782 in the Freyberg school of mines, where he studied for vears under A.G. Werner. He was designated in 1788 to classify the mineral collection of N.G. LESKE, that was also very rich in animals. This resulted the following year in the publication of the Museum Leskeanum, consisting in two in 8° volumes. The second volume proposed a new method of classifying minerals on the basis of their natural characters and was epoch-making in the history of mineralogy. The first volume (Regnum Animale) is the subject of this note. After this, Karsten published numerous works on mineralogy and related fields. In 1789 he was lecturing at the University of Berlin, where progressed rapidly in the scientific hierarchy and in 1810 he was nominated to head of the administration of sciences. He died shortly later, in Berlin on May 5, 1810. Nathanel Gottfried LESKE, whose collection is described in the AMfuseum Leskeanum, was a German naturalist born in Muskau on October 22, 1751. He was a professor at Leipzig and Marburg and published books on botany, ichtyology, physiology, etc. He died in Marburg on November 25, 1786. The Museum Leskeanum (of which we will only consider the first volume. the second being devoted to minerals) consists of 320 pages and 9 plates. An exemplary 1s present at the Dautzenberg Library (Institut Royal des Sciences Naturelles de Belgique, Brussels). The front page 1s reproduced in Fig. 1. The Volume 1 of the Museum Leskeanum consists of 6 parts: Mammalia (86 specimens), Aves (161 specimens), Amphibia (120 specimens), Pisces (72 specimens), /nsecta (2576 specimens) and Vermes (1430 specimens). The numbering in the book concerns specimens, not species. The voluminous part on Insects 1s not of the hand of Karsten., but is from one J.J. Zschach as clearly stated on the first page of the chapter (see fig.2). This interesting but separate work will not be discussed here and should be referred at as "Zschach in Karsten". Availability of the Museum Leskeanum. To be available à work must: a/ be published within the meaning of the International Code of Zoological Nomenclature after 1757. 11 APEX 9(1): 11-16, April 1994 b/ not be suppressed by the International Commission of Zoological Nomenclature for nomenclatural purposes. c/ consistently apply the Principle of Binominal Nomenclature. We shall consider these three points in succession. a/ The book was printed in Leipzig in 1789 ("LIPSIAE, SUMPTIBUS HAEREDUM 1IG. MULLERI"). It obviousiy meets the requirements of Chapter III (Criteria of Publication) of the Code. b/ The Museum Leskeanum was not found in the Official Index of Rejected and Invalid Works in Zoological Nomenclature. c/ AIl the descriptions of Karsten (numbering 1869) have been checked. By large, Karsten's text is consistently binominal, never referring to an animal by a vernacular name only. One entry is clearly non-binominal: "Pinna haud ignobilis" (p. 186, 403-404). This should present no nomenclatural problem, as it simply means: "a Pinna that 1s not of the species ignobilis". It is obviously not a name but, on the contrary, a disclaimer approximating our modern "Pinna aff. ignobilis". The remainder of the text still contains a number of trinomens as well as a few names that are questionable for the modern taxonomist. As examples of the most disputable cases we found, let us cite: Arca Rhomboidalis I. Orient. (p.173), Cypraea Caput Serpentis (p. 204), Turbo Cidaris rufescens (p. 275), Turbo Tectum Persicum (p. 275). Note on the Museum Leskeanum Duchamps & Tursch On the one hand, our first reaction was that Karsten's work 1s not consistently binominal and should be rejected. On the other hand, we are well aware that such departures from strict binominal nomenclature are commonplace in ancient works that are officially available in the meaning of the International Code of Zoological Nomenclature. It is obvious that the rigid requirements of the Code are de facto more flexible in the case of very old authors. But by how much should the rules be stretched ? We strongly feel that it is not for us to rule on this point. In the present case, we do not even need to air Our own opinions on the subject. The acceptable level of departure from strict binominality can be very simply determined by comparing (see Table 1) the major deviations to binominal nomenclature found in the Museum Leskeanum with some present in the Museum Boltenianum (Rôding, 1798), a work that is approved by the International Commission of Zoological Nomenclature (Official List of Works Approved as Available for Zoological Nomenclature. Direction 48. Title 26. Publ. 21 nov. 1956). Such cases are actually very common in Rôding's Museum Boltenianum. Amongst many others (not utilised in Table I) let us cite: Cassis Caput Bovis (p. 28, n° 342). Cassis Mitella Polonica (p. 29, n° 357), Conus Mille punctatus (p. 47, n° 605), Conus cutis anguina (p. 619, n° 48), Zrochus Tectum chinense (p. 81, n° 1057), Neptunea Corona Mexicana (p. 116, n° 1492). Museum Leskeanum (Karsten, 1789) Museum Boltenianum (Rôding, 1798) Arca Rhomboidalis I. Orient. (p.173) Conus Cauda Erminea (p. 191) Cypraea Caput Serpentis (p. 204) Turbo Cidaris rufescens (p. 275) Turbo Tectum Persicum (p. 275) Conus Archithalassus Indiae (p. 43, n° 545) Bulla Ovum Vanelli (p. 15. n° 181) Cypraea Caput Serpentis (p. 23, n° 23) Bulla Ampulla Striata (p. 15, n° 182) Cidaris Tectum persicum (p. 84, n° 1089) Table 1. Examples of some departures from strict binominality in Karsten (1789) and in Rôding (1798). The short comparison table given here could be considerably extended, with the same result. Karsten's occasional deviations to strict binominalism are all exactly matched in Rôding's Museum Boltenianum, published a 12 decade later. If only for the sake of coherence, what is accepted for Rôding should also be accepted for Karsten and we see no reason why the Museum Leskeanum names should not be available. Duchamps & Tursch Comments. The work of Karsten is of incomparably higher scientific standing than the sterile, uninteresting enumeration of Rôding. It is perfectly clear from the Praefatio that the author is conversant with the Linnean system and well aware of the distinction between generic and specific characters. He 1s also familiar with natural variation, as evidenced all along the text. His descriptions (see examples in fig. 3) are objective and informative. For molluscs, Karsten gives measurements of length and width and is thus à precursor of shell morphometry (the ratios of his measurements on Oliva were checked by us and found most accurate). In contrast to most of his contemporaries, Karsten gives a detailed list of references and carefully analyses his sources. Reading the Museum Leskeanum always gave us an impression of modernity. The author was obviously quite in advance on his time, as attested by his fame in another field. mineralogy. ACKNOWLEDGEMENTS. We thank Dr. J. Van Goethem (I.R.Sc.N.B.) for access to the books of the Dautzenberg Library. We are specially grateful to Mr Antoine Lievrouw (LR.Sc.N.B.) for his kind and constant help. We thank Dr. Henry Coomans (Zoëlogisch Museum, Amsterdam) for his friendly and valuable advice. Note on the Museum Leskeanum APEX 9(1): 11-16, April 1994 References DIDOT, Firmin Frères, 1858. Nouvelle bibliographie générale, depuis les temps les plus reculés jusqu'à nos jours. HOEFER, Ed. Vol 27: 467-469. Paris. DUNKER & HUMBOLDT, 1977. Neue Deutsche Biblographie. Herausgegeben von der Historischen Kommission bei der Bayerischen Akademie der Wissenschaften. Band 11: 304. Berlin. ibid. Band 14: 328. KARSTEN, G.; 1789. Museum Leskeanum. Regnum Animale. OQuod ordine systematico …. Müller, Lipsiae. RODING, P.F., 1798. Museum Boltenianum sive Catalogus Cimeliorum ….. Pars Secunda. Hamburg. 13 APEX 9(1): 11-16, April 1994 Note on the Museum Leskeanum Duchamps & Tursch a —————_—_—_—_—" "77 Duchamps& Turs LESKEANVM _ REGNVM ANIMALE QYOoD ORDINE SYSTEMATICO DISPOSVIT ATQVE DESCRIPSIT D. L. GVSTAVVS KARSTEN, SOCILT. NAT. CVRIOS. HALENS. SODALIS. l KGLCHIR.ACAN VaO | Cum IX. iconibus piléis. / EVE SEE. SVMPTIBVS HAEREDVM.I G. MVLLERS D EL : BALE, Fig. 1. The cover page of the Museum Leskeanum. Duchamps & Tursch Note on the Museum Leskeanum APEX 9(1): 11-16, April 1994 CLASSIS V. INSECTA CORP EP EU ZSCHACHIL Haec claffis a Clariff. J. J. Zfchachio Med. Bacc. elaborata, jam ante annum et quod excurrit fub titulo: Mufeum N. G, Leskeanum, Pars entomnologica, ad fyitema entomologiae CI. Fabricii ordinata 8 maj. in bibliopolio Mulleriano typis exprefla eft, Quae entomologica colleétio, cum fingula- rem colleétoris curam expofcat, fortailis a reliquo mufeo fejunéta, naturac ferutatori, cuius impriwis interft, iftam polidere, feparatim divendetur. Fig.2. The cover page of the part on insects un APEX 9(1): 11-16, April 1994 Note on the Museum Leskeanum 252 MUS. LESK. REC. ANIM. Murex Olearium. Linn.S. N. Gen, 325. Sp. 530. a Born, teft, muf, Vind. pag. 257. 919 M. OI. tefla pallida transverfim ftriata, occllis ferrugineif feriatim cinéta, apcrtura dentitulata. Chemin, Konch. Kab. °F, 4, tab. 127. fig. 1223. Long, 6 poll. 6 lin, lat. 3 poll. 8 lin. 920 M. Ol. telta /ubferruginea albido maculata, varicibus ‘. alternis tuberculatis; apertura la@tea ad labruim incats nata fufco maculata, labioque fufco, Long. 6 poll. lar, ; poll. 921 M. OI. tefta albida unicolor; labrum dentibus /o/itariis obfitum; paullulum deftru&um. Long. $ poll. 8 lin. lat. 2 poll, € lin. Not. Hacin fpecice plane fecuti fumusPerill. a Borniums' nulla enim habita ratione fententiae Chemnitzii fecun, ‘‘dum quai fig. fupra citata M, Olear. :Linn. propteres non refponderet, quod apertura effet denticulata; fed Archiater Linn. etiam im Lampade, Feimorali et in pluribus tefkis, aperturam edentulan: docet, ubi Con- chiologiftae recentiores veritatem ct conflantiam cha« ra@eris huius non confirmatam viderunt, Cypraea amethyftea. Linn, S. N. Gen. 3120, Sp. 334. | 540 C. aim. tefta fubfufca, antice ac poflice violaceo undata, lateribus gibbis, fufco maculatis. Martini Konch: Kab. T. 1. tab. 25. fig. 248. ‘Long. 2 poll, $ lin. lat. r poll. 4 lin. Conus Ruflicus. | Linn, $. N. Gen, 319. Sp. 306, 463 C.R. tefta ex livido flavefrens albida, in medio fa/ciata, area poflica punétis albidis clevatisin fericbus cinéta. Ind. Occ. Martini Konch, Kab. T. 2. tab, 63. fig. 694. Long. 1 poll, 8 lin. lat. « poll, 464 C.R. tefla Jublivida, spertura intus gibbofa. ; Long. 1 poli, 7 lin, lat, 14 Jin. 465 C. R. tefta livida albo fafciata, area poñica #ndique filis grañulofis, antica duobus tantum cinéla. Long. 1 poil. 2 lin. lat, 9 lin 466 C.R. tefla flavefcens fafcia albida in medio nu/{a, Long. 1 poll, s lin. lat, us lin, Fig. 3. Examples of descriptions of mollusc species 16 Duchamps & Tursch Rolän & Fernandez-Garcés Triphoridae of Cuba The Family Triphoridae (Mollusca, Gastropoda) in Cuba. 4. The genera Monophorus, Nototriphora, Cosmotriphora and Cheirodonta, with the description of three new species Emilio ROLÂN Cânovas del Castillo 22, 36202 Vigo, España Raul FERNÂNDEZ-GARCÉS Poder Popular, Cienfuegos, Cuba KEY WORDS: Triphoridae, Caribbean Sea, Cuba. PALABRAS CLAVE: Triphoridae, Mar Caribe, Cuba. ABSTRACT. New information on the species known of the genera studied are reported. One new species of the genus Monophorus and two of Cheirodonta are described. RESUMEN. Se realizan nuevas aportaciones a las especies ya conocidas de los géneros estudiados y se describen tres especies nuevas, una del género Monophorus y dos de Cheirodonta. INTRODUCTION Following the publication of the first works on the family Triphoridae in Cuba. in which the species of the genera Metaxia Monterosato, 1884 (ROLAN & FERNANDEZ-GARCES. 1993a). Iniforis Jousseaume. 1884 (ROLAN & FERNANDEZ-GARCES, 1993b) and /sotriphora (ROLAN & ESPINOSA, in press) were studied, we continue the revisory present work in which four genera are studied. New information on some of the previously known species 1s reported and three species new to science are described. Additional material was recently examined from Bahamas Islands loaned by Colin Redfern, of Boca Raton, Fla. USA. Abbreviations: MNCN: Museo Nacional de Ciencias Naturales, Madrid IES: Instituto de Ecologia y Sistemäatica, La Habana AMNH: American Museum of Natural History, New York BMNH: The Natural History Museum, London MNAN: Museum National d'Histoire Naturelle, Paris ZMA: Zoologisch Museum, Amsterdam RESULTS SUBFAMILY TRIPHORINAE Gray. 1847 Genus Monophorus Grillo, 1877 Monophorus olivaceus (Dall, 1889) (Figs. 1-3, 6, 8, 30 MO) = ornatus auct. non Deshayes, 1832 Material examined. NORTH OF CUBA: 3 specimens and 2 shells at 3 m, off the Hotel Comodoro, La Habana,; 3 shells at 4 m, Jibacoa: 2 shells at 3 m, Baracoa. SOUTH OF CUBA: 6 shells at 17 m, Punta Pedernales: 4 shells at 15 m, Cayo Matias, 3 fragments at 15 m, Cayo Avalos, Archipiélago de Los Canarreos;, 2 specimens and 16 shells at 3 m, Rancho Luna: and 20 shells and some fragments at 45 m, Cienfuegos Bay. Description Shell (Fig. 1-3) sinistral. oval-elongated with pointed apex and two or three nodulous cord on each whorl, the nodules being white or brown. Protoconch (Fig. 6) of dark brown colour with three and a half whorls. On the first whorl there are tubercles with arrow-head shape. The other whorls have two spiral cords which are crossed by uninterrupted axial striae. 17 APEX 9(1): 17-27, April 1994 APEX 9(1): 17-27, April 1994 Teleoconch with about 10-12 whorls in larger specimens. The first whorls have two spiral cords from the beginning and, around the fifth or sixth whorl, a third cord begins between the previous two. This last cord is narrower, but it increases gradually to be similar to the lower one around the tenth whorl. Large and round nodules appear in the intersections of the axial ribs with the spiral cords. The colour is brown and white, the subsutural cord being brown with one white nodule between each two or three brown ones. The lower cord is always white. The intermediate cord has some white nodules and some brown. Under magnification, a very fine axial striation in the spaces between the cords and the ribs may be seen. Dimensions: the biggest shells can reach as much as 10 mm. Animal à little translucent with variously sized spots formed by very small points of white-milk colour. The propodium has a yellowish colour marked in the anterior border of the foot. Radula (Figs. 8 and 30 MO) with formula 15-1-1-1-15. The rachidian tooth has five cusps of similar size. The lateral tooth 1s very similar. The marginal teeth also have five cusps but the most peripheral have the external cusps shorter than the internal. Remarks. FABER & MOOLENBEEK (1991) consider that the correct name for this species should be Cosmotriphora olivacea (Dall, 1889) instead of "Zriphora" ornata Deshayes, 1832. Its position in the genus Cosmotriphora seems not adequate according to the characteristics of protoconch and radula: hemispherical tubercles in the protoconch and three cusps in the marginal teeth, in Cosmotriphora. In contrary to the other known species of Monophorus, the present one has an animal without red colour However, for this reason we do not think that it should be placed in another genus and we agree with the opinion of BOUCHET (1984) on the generic value of the radula and protoconch in Monophorus. | Monophorus ateralbus n. sp. (Figs. 4. 5, 7. 9, 30 MA) Material examined. NORTH OF CUBA: 2 specimens at 2 m, Marianao Beach. and 7 shells at 4 m, off the Hotel Comodoro Beach, La Habana. SOUTH OF CUBA: 2 shells and 4 fragments with protoconch at 15 m, Cienfuegos Bay. BAHAMAS: 1 shell in beach drift, Abaco Island. Description Shell (Fig. 4-5) sinistral, with an oval- elongated form, a little wider near the base and with pointed apex. 18 Triphonidae of Cuba Protoconch (Fig. 7) with four whorls and of uniform dark brown colour. The first whorl has T-form tubercles. The others have two spiral cords crossed without interruption by axial threads, which are a little 1rregular and shightly oblique in some parts. Teleoconch with 7-9 whorls, which begins with two nodulous, spiral cords. These nodules are a little bigger in the lower cord. Around the sixth to seventh whorls a new spiral cord appears, situated near the upper one. In the body whorl, at the begining, there are five nodulous cords, and near the anterior end, new spiral cords appear, there being eight by the end of the shell. Among these eight, the lower one 1s smaller, not nodulous and very close to the siphon. The aperture 1s rounded and the anal sinus is only slightly deeper but open. The siphonal canal is short, curved and closed by a fold from the external lip. The distribution of the dark brown and white colour in the teleoconch 1s in bands. The lower nodulous cord is white and the others are brown. This white nodulous cord ends in the anal sinus. Dimensions between 3 and 6 mm, but the exact size of most of the collected specimens 1s difficult to determine because the shells with completed development of the body whorl are frequently decollated. The animal has à whitish colour with numerous red-brown spots on the head and on the dorsum of the foot. The tentacles have very small white dots. There is a bigger white spot behind the head and additional spots on the posterior part of the foot. Laterally on the base of the tentacles, at the same level of the eyes, there 1s a small lateral prominence. Radula (Figs. 9 and 30 MA) with formula 6-1-1-1-6. Rachidian tooth with five cusps from which two are longer. The lateral tooth has five cusps the smaller being most external. Marginal teeth have four cusps which are a little longer in the outermost ones. Type material. Holotype (of 3.9 mm), MNCN, n° 15.05/11141; 1 paratype in JIES, AMNH n° 226469, ZMA and 4 in the collections of R. Fernaändez-Garcés and E. Rolan. Type locality: Marianao Beach, La Habana (Cuba). Etymology. The specific name 1s due to the dark brown (almost black) and white banded coloration of the shell. Remarks. Monophorus ateralbus n. sp. has a shell with brown and white spiral cords. Because of this kind of coloration, the shell must be compared with the following species: Monophorus olivaceus (Dall, 1889) has the spiral cords with the same colours but in each cord the nodules may be brown or white: also Rolän & Fernändez-Garcés Rolän & Fernandez-Garcés different are the animal coloration and the radula. "7riphora" intermedia (C. B. Adams, 1850) has smaller and more numerous nodules and three spiral cords from the third whorl of teleoconch. “7riphora" ellyae De Jong & Coomans, 1988 has the position of the spiral cords inverted, the upper one being white and the same occurs with "7riphora" elvirae Jong & Coomans, 1988. /niforis turristhomae (Holten, 1802) has smaller nodules and a tubular anal hole far from the aperture. The differences of the shell with those of the Cheirodonta verbernei (Moolenbeek & Faber, 1989) and C. decollata n. sp. are based on the different position of the brown and white colour in the spiral cords in most parts of the teleoconch. Genus Nototriphora Marshall, 1983 Nototriphora decorata (C. B. Adams, 1850) (Figs. 10, 14, 16, 30 ND) Material examined. NORTH OF CUBA: 4 shells at 6 m, Jibacoa, 2 shells at 8 m, Herradura. SOUTH OF CUBA: 2 specimens and 4 shells between 4 and 20 m. Punta Francés, and 1 shell and 3 fragments between 20 and 50 m, Punta Pedernales, Isla de la Juventud; 10 shells at 15 m, Cayo Matias and 1 shell at 2 m, Cayo Diego Perez. Archipiélago de Los Canarreos: 6 specimens and 15 shells between 15 and 50 m, Cienfuegos Bay Description Shell (Fig. 10): see ADAMS (1850) and CLENCH & TURNER (1950). This description should be complemented by the following information: Protoconch (Fig. 14) of brown colour. It has between four and half to five spiral whorls. The first one with hemispheric tubercles very dense, the rest of the whorls with uninterrupted axial ribs crossed by one spiral cord in the first whorls and two in the lower. Near the anterior end both cords are fused into one. The teleoconch presents a very fine, spiral striation in the spaces between the axial ribs, as mentioned by BOUCHET (1984). Animal of hyaline white colour with very small white-milk spots irregularly distributed in the head and the dorsal part of the foot. Tentacles translucid. GARCIA & LUQUE (1986) mention the presence of some red spots on the flanks near the operculum; we could not find these red spots in several animals examined. We think that it is not a constant character. Tniphoridae of Cuba APEX 9(1): 17-27, April 1994 Operculum rounded, with a central nucleus and a translucent vellowish-white colour. Radula (Figs. 16 and 30 ND) with formula 18-1-1-1-18. The rachidian tooth has three equal cusps. The lateral tooth has five cusps, of which the second one is less prominent. The first marginal tooth has four cusps. the two central ones being filiform. The rest of the marginal teeth have three cusps the central one being longer and narrower. Remarks. Some shells collected in Cienfuegos present a violet coloration instead of brown. alternating with white. As we can not find any other differences, this coloration must be considered as an ecological variation. Genus Cosmotriphora Olsson & Harbison, 1953 Cosmotriphora melanura (C. B. Adams, 1850) (Figs. 11, 25, 26, 30 CM) Material examined. NORTH OF CUBA: 5 shells at 10 m., Herradura: 5 shells at 4 m., Jibacoa; 3 shells at 4 m. off the Hotel Comodoro Beach, La Habana; 2 shells at 6 m, Baracoa. SOUTH OF CUBA: 6 shells between 8 and 17 m, Cayo Matias, Archipiélago de Los Canarreos; 4 shells at 50 m. Punta Pedernales, Isla de la Juventud: 40 shells between 10 at 20 m, Cienfuegos Bay. Description Shell (Fig. 11), see BOUCHET (1984). It has been figured by BOUCHET (1984, p. 36, fig. 27) and by FERNANDES & ROLAN (1986, pl. 1, fig. 1, pl. 2, fig. 1), for specimens from the eastern Atlantic. WARMKE & ABBOTT (1961, pl. 13, fig. 1) and ABBOTT (1974, fig. 1132) showed Caribbean specimens. The radula is drawn in BOUCHET (1984, fig. 16). The protoconch of a shell from Cuba is represented in the Fig. 25. Animal of opalescent whitish colour with numerous white spots which are slightly vellowish in the propodium. Behind the eyes there are subcutaneous yellow areas. Radula (Figs. 26 and 30 CM) with formula 10-1-1-1-10. It has a rachidian tooth and very similar lateral teeth, each one with four cusps. Marginal teeth have three cusps: the inner ones have their three cusps of almost equal size, while the external ones have their lateral cusps shorter and the central one longer. In the most external, the central cusp becomes filiform. 19 APEX 9(1): 17-27, April 1994 Remarks. This species 1s variable in size: the smallest shell is only 4 mm while others can be as much as 10 mm. The normal white coloration can become cream in some specimens. The shells from the Caribbean have been compared with specimens from Ghana and Cape Verde Islands, showing small differences: in the African shells, the third spiral cord of the teleoconch begins between the 6th and &8th whorl and always has smaller nodules than those of the other cords, except in the body whorl. On the other hand, in the shells from the Caribbean, the third spiral cord begins around the 3rd whorl and, between the 6th and the 8th, it 1s of similar size to the other two. Nevertheless, these differences seem not enough to consider both populations in different specific position. The protoconchs are equal and the radulas, after the examination of several specimens from Cuba and Ghana, have no significant differences. So, the observation of BOUCHET (1984) in relation to the marginal external teeth 1s not confirmed. It is considered an amphiatlantic species. Cosmotriphora arnoldoi Faber & Moolenbeek, 1991 (Figs 1215"15) Material examined. 5 shells and 3 fragments at 20 m, Cienfuegos Bay. Description Shell, see FABER & MOOLENBEEK (1991). In the Figs. 12 and 13 shells are shown with normal colour distribution which was not evident in original figures because a SEM- photograph was used. Dimensions: although the holotype 1s of a size smaller than 3 mm, some shells from our material reach 6 mm and have 10 whorls (Fig. 12). Remarks. In FABER & MOOLENBEEK (1991) the assignation of this species to the genus Cosmotriphora is not explained. Perhaps it could be on similarity of its protoconch with that of Cosmotriphora melanura. The lack of knowledge about the radula and operculum makes this assignation only a provisional effort. Genus Cheirodonta Marshall, 1983 Cheirodonta verbernei (Moolenbeek & Faber, 1989) Figs. 17, 18, 22, 30 CV) Material examined. 1 specimen, 5 shells and 2 fragments with protoconch, in sédiments at 25 m, Cienfuegos Bay. 20 Triphoridae of Cuba Description Shell (Figs. 17 and 18), see MOOLENBEEK & FABER (1989). Some shells of our material are similar to the description of the holotype, in which a brown colour with knobs of a lighter shade is mentioned. Other specimens have a lighter lower cord in the penultimate whorl and, in the body whorl, the upper one white. The protoconch (Fig. 22), has the apex covered with hemispheric tubercles (it can be observed in the picture in spite of a fracture) and it is not smooth, as is mentioned in the description of the holotype, which has this part polished by erosion. Animal translucent white with milk-white spots irregularly distributed on the dorsum. The radula (Fig. 30 CV), studied from one live collected specimen and partially destroyed during the protographic process, showed a rachidian tooth with two cusps at each side and a lateral tooth with shortish cusps. The operculum is rounded, light yellow, translucent, with a central nucleus, the external border obliquely elevated outwards and with a small depression in the centre of the internal part. Remarks. The inclusion of this species in the genus Cheirodonta is based on the radular characteristics, similar to the Cheirodonta labiata (see MARSHALL, 1983, fig. 8 C) and C. pallescens (see BOUCHET, 1984, fig. 10-11). Cheirodonta decollata n. sp. (Figs: 19/20M102372730CD) Material examined. NORTH OF CUBA: 8 specimens at 2 m, Marianao Beach, and 1 specimen at 3 m, Marina Hemingway, La Habana; 1 specimen and 6 shells at 3 m, Baracoa. SOUTH OF CUBA: 8 specimens and 10 shells at 3 m, Rancho Luna, 1 specimen, Cable Inglés and 8 shells at 10 m, Cienfuegos Bay. Description Shell (Figs. 19, 20 and 21) sinistral, ovoid- elongated, slightly pyriform, usually with the protoconch lost (only present in 1/6 of the shells studied). Protoconch (Fig. 23) with about four whorls. The first whorl is covered by hemispheric tubercles; the rest have two spiral cords crossed by axial ribs. Dark brown colour. Teleoconch with seven or eight whorls which present two spiral cords with rounded and rather big nodules, being slightly larger in the upper cord. On the last whorls these cords are separated, especially in the penultimate one. Rolän & Fernändez-Garcés Rolän & Fermändez-Garcés In the beginning of the body whorl there are five cords, a new one appearing below the upper one. At the end of the spire there are seven or eight cords by the presence of several others. The axial ribs between the nodules are a little oblique, being more evident in the lower whorls. Towards the end of the body whorl the axial ribs are slighter and more closed; at the same time, the spiral cords are finer, bifurcated and attenuated, almost disappearing near the free border. The aperture has the form of an inverted arc; the anal sinus is deep but open. The siphon is short, curved and closed by a fold of the external lip. There is a microsculpture of microscopic tubercles spiraliy aligned. The coloration is very characteristic and constant: the two first whorls of the teleoconch are of a cream colour but with the subsutural cord brown. From the second whorl, both nodulous cords change to uniform brown, but from the 5th whorl can be observed that the nodules of the upper cord are slightly bigger and whitish. This white colour is more evident in the penultimate whorl. In the body whorl the upper cord is bifurcated, a finer cord appearing below. The colour of the upper cord continues white until the end where its nodules are smaller and brown, finishing in the anal sinus. The base 1s brown. Animal translucid white with opaque spots formed by very small white-milk dots. Tentacles translucid. Operculum white, translucid, multispiral and with a central nucleus. Radula (Figs. 24 and 30 CD) with formula 7-1-1-1-7. Rachidian tooth with 9 cusps of which the central one is shorter. Lateral tooth with 8 shortish cusps. Marginal teeth of comb- like form with elongated cusps. Dimensions. The holotype is 3.95 mm of length. Other specimens with protoconch are slightly smaller. In most shells 1t 1s not possible to know the real size due to their decollation. Type material. Holotype of 3.95 mm and one paratype, in MNCN n° 15.05/11142. Two paratypes each in IES, AMNH n° 226470, BMNH n° 1993062, MNHN, ZMA and 11 paratypes in the collections of R. Fernändez- Garcés and E. Rolän. Type locality. Habana. Habitat. On rocky bottoms, under rocks or outside of the coral barrier, under dead corals. Etymology. The species is named after the fact that it looses its apex during maturity. Remarks. The shell of Cheirodonta decollata n. sp. at a superficial look may remind one of Monophorus ateralbus n. sp. but this last species has the lower cord white instead of brown. Also there are differences in the Marianao Beach, La Triphoridae of Cuba APEX 9(1): 17-27, April 1994 microsculpture of the protoconch and the radula when these characters can be studied. From Cheirodonta verbernei (Moolenbeek & Faber, 1989) it must be differentiated because this latter species has the first whorls of the teleoconch of uniform brown colour instead of cream with a brown cord. Also, C. verbernei has the lower cord white in the penultimate whorl, and the white colour of the nodules is less evident. This species was also collected in Bahamas (Redfern, pers. com.). Cheirodonta apexcrassum n. Sp. (Figs:27728:29) Material examined. NORTH OF CUBA: 4 shells and 9 fragments with protoconch, in sediments at 7 m, Jibacoa. BAHAMAS: 3 shells at 10 m, Chub Rocks, Abaco Island. Description Shell (Figs. 27 and 28) sinistral, ovoid- elongated, a little pyriform. Protoconch (Fig. 29) relatively large, with uniform brown colour. It begins with a well- differentiated nucleus in a vertical position, and consists of between 2 and 2 1/2 spiral whorls. These whorls present two prominent spiral cords which are irregular at the beginning and nodulous after. Another small cord is on the suture. At the end of the protoconch both cords are fused in one. The beginning of the teleoconch is 1ll-defined. Teleoconch with 5-6 whorls. It begins with the lower cord a continuation of the only cord of the protoconch. Later, the upper cord appears smaller, but increasing quickly and achieving the same size as the lower one. Both have evident nodules which are connected by axial ribs. Towards the fourth whorl a new cord appears between the last ones, nearer the upper one and with smaller knobs. On the body whorl there are six cords from which the three upper ones are nodulous, the three lower being smooth. Aperture slightly ovoid with a prominent cutting external lip and a superior open anal sinus. In the base there is a fold which closes the siphonal canal. The siphon is short and curved. The columellar lip has a basal prominence towards the beginning of the siphonal canal and another up near the sinus. Coloration is almost uniform brown, the upper cord a little lighter in the last whorls and also lighter in the external lip of the aperture. Type material. Holotype (Fig. 27) of 2.78 mm, in MNCN n° 15.05/11143. One paratype each in IES, AMNH n° 226471, BMNH n° 1993061, ZMA, MNEN and the collection of R. 21 APEX 9(1): 17-27, April 1994 Fernändez-Garcés; three (from Abaco) in that of Redfern and six in that of E. Rolän. Type locality. Jibacoa, in North of Cuba. Etymology. The specific name makes reference to the thickness of the protoconch. Remarks. The position of the present species in the genus Cheirodonta is only tentative, based on the great similarity (shell and protoconch) with the species shown by MARSHALL (1983), Cheirodonta labiata (A. Adams. 1851) from Australia. Cheirodonta apexcrassum n. sp. can be differentiated from "Zriphora" calva Faber & Moolenbeek, 1991 because this last species has a smaller protoconch and smaller nucleus: also lacks the two constant cords of the protoconch. It differences from the other species of the genus described in the present work by having a paucispiral protoconch. ACKNOWLEDGEMENTS. We are indebted to Francisco Guitiän Ribera, of the Câtedra de Edafologia of the Facultad de Farmacia and to Maria de los Angeles Rodriguez Cobos, of Anatomia of the Facultad de Medicina, both of the University of Santiago de Compostela, and also to Enrique Porto of the AIMEN (Universidad de Vigo) for the scanning electron micrography: to the collegues of the collecting expeditions in Cuba, especially to Angel A. Luque, José Templado, Diego Moreno, Jesüs Ortea and Enrique Vidal, for their help in the obtaining the micromollusc material: to José Espinosa for his contribution to the collecting of material, on part of which this work 1s based; to Colin Redfern for his loan of material from Bahamas; to Kevin M. Plumley for his help in the english text. REFERENCES ABBOTT, R. T. 1974. American seashells. (2nd. Ed.). Van Nostrand Reinhold Co. New York. 663 pp., 24 pls. ADAMS, C. B. 1850. Description of supposed new species of marine shells which inhabit Jamaica. Contributions to Conchology (4): 56- 68 22 Triphoridae of Cuba BOUCHET, P. 1984 Les Triphoridae de Méditerranée et du proche Atlantique (Mollusca, Gastropoda). Lavori S.IM., 21: 5- 58. CLENCH, W. J. & R. D. TURNER, 1950. The Western Atlantic marine mollusks described by C.B. Adams. Occasional Papers on Mollusks, 1 (15): 233-403. : FABER, M. J. & R. G. MOOLENBEEK, 1991. Two new shallow water triphorids and a new name in Metaxia from Florida and the West Indies. Apex. 6 (3-4): 81-85. FERNANDES, F. & E. ROLAN, 1986. A Familia Triphoridae (Mollusca: Gastropoda) no Archipélago de Cabo Verde. Publicaciones Ocasionais Sociedad Portuguesa de Malacologia. (11): 17-32. GARCIA, M. T. & A. A. LUQUE, 1986. Contribuciôn al conocimiento de los gasterépodos prosobranquios de la Isla de la Juventud y del Archipiélago de los Canarreos (Cuba). Revista de Investigaciones Marinas, 7 (29: 31-52. MARSHALL, B. A. 1983. A revision of the recent Triphoridae of Southern Australia. Records of the Australian Museum, supp. 2: 1-119. MOOLENBEEK, R. G. & M. J. FABER, 1989. Two new Triphora species from the West Indies (Gastropoda, Triphoridae). Basteria, 53 (4-6): 77-80. ROLAN, E. & J. ESPINOSA, (in press). The family Triphoridae (Mollusca, Gastropoda) in Cuba 3. The genus /sotriphora. Basteria. ROLAN, E. & R. FERNANDEZ-GARCES., 1993a. La familia Triphoridae en la isla de Cuba 1. El Genero Metaxia. Bolletino Malacologico, 28 (5-12): 169-176. ROLAN, E. & R. FERNANDEZ-GARCES, 1993b. The family Triphoridae (Mollusca, Gastropoda) in Cuba 2. The genus /niforis Jousseaume, 1884. Apex, 8 (3): 95-106. WARMKE, G. L. & R. T. ABBOTT, 1961. Caribbean Seashells. Livingston Publishing Co. Wynnewood, Pennsylvania. 348 pp... 43 pls. Rolän & Fernändez-Garcés Rolän & Fernandez-Garcés Triphoridae of Cuba APEX 9(1): 17-27, April 1994 Wu d d 44 LU ER MiTICITETP) DANS TA "ES d'A: : S'LLEEEST Pas E send ( Dress NI", \ ch Le 1-3. Monophorus olivaceus; 4. Monophorus ateralbus n. sp. Holotype (MNCN); 5. Monophorus ateralbus n. sp. Paratype (coll. E. Rolän); 6. Monophorus olivaceus. Protoconch; 7. Monophorus ateralbus. Protoconch; 8. Monophorus olivaceus. Radula. 9. Monophorus ateralbus n. sp. Radula. (Scale bar: shells: 1 mm; protoconchs: 0.1 mm; radulas 0.01 mm) APEX 9(1): 17-27, April 1994 Triphoridae of Cuba Rolän & Fernändez-Garcés 1 & LL Figs. 10-16. 10. Nototriphora decorata; 11. Cosmotriphora melanura; 12. Cosmotriphora arnoldoi: 13. Cosmotriphora arnoldoi, 14. Nototriphora decorata. Protoconch; 15. Cosmotriphora arnoldoi. Protoconch; 16. Nototriphora decorata. Radula. (scale bar: shells: 1 mm; protoconchs: 0.1 mm; radulas 0.01 mm) 24 Rolän & Fernändez-Garcés Triphoridae of Cuba APEX 9(1): 17-27, April 1994 CSL ES ï | Figs. 17-24. 17-18. Cheirodonta verbernei, 19. Cheirodonta decollata n. sp. Holotype (MNCN). 20-21. Cheirodonta decollata n. sp. Paratypes (col. E. Rolän); 22. Cheirodonta verbernei. Protoconch; 23. Cheirodonta decollata n. sp. Protoconch; 24. Cheirodonta decollata n. sp. Radula. (scale bar: shells: 1 mm; protoconchs: 0.1 mm; radulas 0.01 mm) 25 APEX 9(1): 17-27, April 1994 lriphoridae of Cuba Rolän & Fernändez-Garcés Figs. 25-29. 25. Costrotriphora melanura. Protoconch; 26. Costrotriphora melanura. Radula. C central tooth; 27. Cheirodonta apexcrassum n. sp. Holotype (MNCN); 28. Cheirodonta apexcrassum n. Sp. Paratype (Col. E. Rolän); 29. Cheirodonta apexcrassum n. sp. Protoconch. (scale bar: shells: 1 mm; protoconchs: 0.1 mm; radulas 0.01 mm) 26 Rolän & Fernändez-Garcés Triphoridae of Cuba APEX 9(1): 17-27, April 1994 IGN CS OMR RE RS LUE NP LAAER ETES Fig. 30.- Radular teeth: MO: Monophorus olivaceus; MA: Monophorus ateralbus; ND: Nototriphora decorata CM: Cosmotriphora melanura; CV: Cheirodonta verbernei; CD: Cheirodonta decollata C- rachidian tooth; L- lateral tooth; M1-2-etc.- marginal teeth. 27 LARGE CHOIX D'OUVRAGES ET DE | Werner PERIODIQUES DE FMALACULOGIE EN FRANCAIS, NEERLANDAIS, Massier ANGLAIS ET ALLEMAND, Liste sur demande. Specimen Shells Verte par correspondance. Exposition permanente de coraux et Librairie de coquillages de collection, UNIVERS SOUS - MARIN KONINKLIKE BAAN 90 B 8460 KOKSIJDE Æ& 058/512821 Largest specimen shell dealer in Africa a —— | Specialized in shells of the Southern African region HIGH QUALITY OF SPECIMEN SHELLS Good selection ofworid-wide BRAZILIAN SEASHELLS Eur eu, AND LANDSHELLS _. one MAIL ORDER RETAIL. d ÿ à FREE LISTS. Donax Seashols Werner Massier PO. Box 1671 MAURICIO ANDRADE LIMA : Bus ne 0 ne 2 | 4275 Margate/Natal-SOUTH AFRICA Tel (081) 241-9862 Tamarineira CEP 52051 | Tel.+ Fax 3931/76153 RECIFE - PE - BRASIL Collection F. Lorenz, jun. Rosenstr. 4, 35418 Buseck; Germany Telephone/FAX. 06408-1098 Write for free price list, wantlists welcome. Many impossible-to-set rarities of Cowries, Maroinellas, Cones and many other families, from the collection of Felix Lorenz, jun. WANTED RARE OVULIDAE - PLEUROTOMARIIDAE HARPIDAE - LAND SHELLS SHELLS FROM THE COLLECTION Guido T. 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XVIII The distribution of Oliva species and the variation of their colour patterns in Hansa Bay (Papua New Guinea). Studies on Olividae. XIX. Where is the suture of Oliva shells ? Studies on Olividae. XX. The pre-Lamarckian names for Oliva species. Contribution to the knowledge of the family Caecidae. 1. À new Caecum from Canary Islands (Caenogastropoda: Rissooidea) Révision des Muricidae de l'Eocëne de la falaise de la Côte des Basques à Biarritz (Pyrénées-Atlantiques, France) R. Duchamps Dr. Y. Finet L. Germain R, Houart Dr. CI. Massin Prof. B. Tursch Dr. J. Van Goethem Editeur responsable : Comité d'édition : Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s), et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. 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XVII APEX 9(2/3): 29-46, juil. 1994 EN CR ES PR NE À ee ES Studies on Olividae. XVIII. The distribution of Oliva species and the variation of their colour patterns in Hansa Bay (Papua New Guinea). C. VAN OSSELAER (‘), J. BOUILLON (**), J.M. OUIN (**) and B. TURSCH (*) * Laboratoire de Bio-Ecologie. ** Laboratoire de Zoologie, Faculté des Sciences. Université Libre de Bruxelles, 50 av. F.D. Roosevelt. B-1050 Brussels, Belgium. ABSTRACT. Sediments of 64 stations where Oliva species have been collected in Hansa Bay (Papua New Guinea) have been analysed for carbonates, organic carbon and granulometry. Comparison between the classification of sediments and the distribution of 30 species shows that the latter fall into stenotopic and eurytopic species. Crypsis 1s generalised in both groups of species. The taxonomic consequences of crypsis are discussed. RESUME. Les sédiments de 64 stations auxquelles des espèces d' Oliva ont été récoltées dans Hansa Bay (Papouasie Nouvelle-Guinée) ont été analysés pour leur teneur en carbonates. en carbone organique et leur granulométrie. La comparaison entre la classification des sédiments et la distribution de 30 espèces montre que ces dernières se divisent en espèces sténotopiques et eurvtopiques. La crypsis est généralisée dans les deux groupes. Les conséquences taxonomiques de la crypsis sont discutées. KEY WORDS. Mollusca. Gastropoda. Oliva. habitat. sediments. stenotopy. eurytopy. crvpsis. polymorphism. taxonomwy. 1. INTRODUCTION. 1.1. The problem. One of the main biological causes of the old "Oliva problem" 1s the great variability of their colours and colour patterns. Colour variations said to be habitat-related have been observed by field collectors (such as HAMLYN-HARRIS. 1970) and the possibility that some described "species" could be ecovariants (GREIFENEDER. 1981) or colour morphs (PETUCH & SARGENT. 1986) has been raised. So the topic of habitat does not only concern ecology and ethology: it also has taxonomic implications. 1.2. Colour variation in molluscs. À great number of studies (see ETTER. 1988) have been prompted by the obvious variations in the shell pigmentation of some molluscs. Much caution is necessary in the interpretation of such data. Many factors -such as the influence of environmental factors (COLTON, 1916: CREESE & UNDERWOOD, 1976: ETTER. 1988: MITTON, 1977). food (COLE. 1975; INO, 1949; LEIGHTON, 1961: MOORE, 1936). genetic determinism (ADAMKIEWICZ & CASTAGNA. 1988, INNES & HALEY. 1977: PALMER. 1985). selective predation (ELEK & ADAMKIEWICZ , 1990: REIMCHEN. 1979). behavioural polymorphism (GIESEL. 1970) and combinations of these can be (and have been) invoked to account for the observed facts. The classical example of colour polymorphism in terrestrial molluscs populations is that of Cepaea nemoralis (CAIN & SHEPPARD, 1954: LAMOTTE, 1959; MURRAY, 1962; JONES et al. 1977) where the relative proportion of genetic morphs varies with habitat, even in localities separated by short distances and this has been correlated with the ability of a predator (thrushes, in that case) to detect their prev. Another visual predator, man, was shown to be responsible for the maintenance of colour polymorphism in an unidentified African Achatinid snaïl (OWEN & REID, 1986). Contribution n° 285, Station Biologique Léopold Il, Laing Island, Papua New Guinea. 29 APEX 9(2/3): 29-46, juil. 1994 Conflicting interpretations have been proposed for some species. The degree of pale striping in the shell of the blue mussel Mvtilus edulis was interpreted as having an adaptive significance, the morphs varving in the proportion of incident sunlight transformed into heath (MITTON, 1977). On the contrary. INNES & HALEY (1977) concluded that the coloration polymorphism of Mvtilus edulis was determined by genetic rather than bv environmental differences. Colour changes associated with a change of food were first reported for Vucella lapillus by MOORE (1936), after COLTON (1916) noted that the shell colour of this mollusc was influenced by the environment. It was later shown that the interpopulation variation in shell colour of Vucella lapillus is in part a response to a selective gradient in the physiological stress due to temperature and desiccation and that selection for crypsis by visually hunting predators does not appear to play a prominent role (ETTER. 1988). In summary. the better studied cases in marine molluscs point at an ‘“/ntimate relationship between genotvpic and environmental factors which influence prosobranch shell colour" (COLE. 1975). The enormous number and varietv of colour forms in some natural populations could be an adaptation in itself, providing protection against visual predators. This hvpothesis. known as reflective selection was proposed by MOMENT (1962) for extremelv variable Donax species. 1.3. Purpose. Any attempt at defining a potential correlation between the colour pattern of Oliva specimens and their habitat had to start with the accumulation of detailed habitat data. Very little detailed information on the habitat of Oliva species 1s available, excepted for a few qualitative reports (HEMMEN. 1981: WIDMER, 1981; WITTIG-SKINNER. 1981). It was felt that useful information could be gained by a methodical study of the distribution of a number of Oliva species between different. well-characterised sediments within a restricted geographic range, followed by the analvsis of a possible habitat-pattern relationship. 1.4. Location. À most suitable location for such a study is provided by Hansa Bay, on the North coast of Papua New Guinea. It is a small semicircular bay (approximately 10 km in diameter) located in Madang Province, near the mouth of the Ramu River. about 110 nautical miles West of Madang. Laing Island (4°10'30"S - 30 Studies on Olividae. XVIII 144°52'47"E). lving roughlv at the middle of the bay is a raised coral reef, covered with vegetation and separated from the mainland by depths of 45-50 m, muddy bottom. Climatic and hydrological data are described by BOUILLON et al. (1986) and CLAEREBOUDT et al. (1989). Detailed environmental data can be found in CLAEREBOUDT (1989). All the coast of Hansa Bay is lined with a long black sand beach. except for small Boro Beach that is formed of white, coarse coral sand. 2. MATERIAL AND METHODS. 2.1. The collection of Oliva. The Oliva species of Hansa Bay have been under survey for nearly 20 vears, since the establishment of King Leopold IIT Biological Station on Laing Island in 1974. Oliva specimens have been obtained over the vears by a variety of methods including dredging (using a small rectangular steel mesh dredge with an opening of 60 x 22 cm). trawling (with a small mesh 3 m rigid frame trawl). SCUBA diving (day and night dives). snorkelling in shallow waters. or beach collecting at the turn of the low tide. When diving, small rigid steel mesh hand "dredges" {in the shape of a dustpan. about 20 x 30 cm) have been especially productive. Baiting and trapping have been often used. The 30 following species have been collected in Hansa Bay: ©. amethvstina (Rôding. 1798), ©. athenia Duclos. 1835. ©. buelowi Sowerby, 1888, ©. bulbiformis Duclos. 1835, O. caerulea (Rôding., 1798), ©. carneola (Gmelin. 1791), ©. ceramensis Schepman. 1911, ©. concavospira Sowerby. 1914 ©. concinna Marrat, 1871, ©. sp. DHB (abbreviation for species "D" of Hansa Bay). O. dubia Schepman, 1911, ©. elegans Lamarck. 1811. ©. funebralis Lamarck. 1811, ©. longispira Bridgman, 1906, ©. mantichora Duclos, 1835, ©. miniacea (Rôding. 1798). O. mucronata Marrat, 1871. ©. panniculata Duclos. 1835, ©. parkinsoni Prior. 1975, ©. paxillus Reeve, 1850. ©. reticulata (Rôding, 1798), ©. rufula Duclos, 1835, ©. semmelinki Schepman, 1911, ©. sericea (Rôding. 1798). ©. smithi Bridgman, 1906, ©. solomonensis Petuch & Sargent, 1986, O. tesselata Lamarck, 1811. O. vidua (Rôding, 1798), ©. cfr. volvaroides Duclos, 1835, ©. sp. ZHB (abbreviation for species "Z" of Hansa Bay). As far as we know, this is by far the largest number of Oliva species ever recorded for a single locality, but this might only reflect an unusual collecting effort. Nearly all species have VAN OSSELAER, BOUILLON, OUIN, TURSCH VAN OSSELAER, BOUILLON, OUIN, TURSCH been obtained in adequate numbers and most specimens have very accurate locality data. All species have been kept and observed in aquaria. some for several months. The taxonomy of the genus Oliva being what it is (see ABBOTT. 1991), some of the above names will undoubtedly have to be corrected in the future. For the time being, the names ©. longispira and ©. smithi were selected only because of the existence of adequate type material. These two taxa are respectively the "species L+X" and "species G" of the "Oliva oliva complex" (see TURSCH & al, 1992). O. amethystina and ©. mantichora were formerlv part of ©. annulata Gmelin, 1791, a nomen dubium encompassing a mixture of species (TURSCH, GERMAIN & GREIFENEDER. 1986). O. sp. ZHB (the specific rank of which is still open to question) and ©. sp. DHB have been collected in numbers but could not be positively identified. Decision on their taxonomic status depends upon future biometric comparison with all possibly related type material. 2.2. The analysis of sediments. During 20 years of study. Oliva specimens have of course been collected at many more locations than we could analyse for sediment and representative stations had to be selected. Two maps (Figs. 1 and 2) show the points where samples were collected in the early months of 1992. Individual sampling stations can be identified in Table 1. AIl sediments were collected by diving. about 800 g being taken in the first 6 cm of substrate. corresponding to the maximum observed burrowing depth observed for Oliva species (VAN OSSELAER & al. 1993). The samples were homogenised. dried at 70-80°C in an oven at Laing Island. individually sealed and sent to Brussels for analysis. The colour of the dried sediments was determined with the Rock Colour Chart of the Geological Society of America. Each sample was separated (homogeneous fractionation by inquartor) into a fraction of about 10 g for carbon analysis and a fraction of about 100 g for granulometric analysis. All weightings were effected with a 0.01 g precision. Granulometric analysis was effected by sieving, using the Udden scale modified by Wenthworth. Sieves were selected for the sand's range. as commonly used in studies of benthic fauna (see for instance JONES et al. 1990). The samples were passed through a series of sieves (2000, 1000, 500, 250, 125 and 63 pm mesh) in a vibrating shaker (HAVER and BOEKER) for 20 Studies on Olividae. XVIII APEX 9(2/3): 29-46, juil. 1994 minutes (15 minutes for calcareous sands that are subject to rapid mechanical wear). For total carbon analysis, an aliquot of about 10 g of sample was finelv ground in a FRITSCH apparatus (type WRR 731 1/4) for 5 min at 98 rpm. Analysis was effected by pyrogenation followed by measurement of the released carbon dioxide with a STRÔHLEIN carbon doser calibrated with Standard B.CS. Steel 163/2. Temperature and atmospheric pressure corrections were accounted for (using the Strôühlein's "Umrechnungstabelle zum Kohlenstoffbestim-mungsapparat"). The same technique was used for organic carbon analysis, but after prior elimination of the carbonates by treating with 50% HCI until no more effervescence was observed. then drying on a hot plate. Carbonate content is calculated from the difference between the total carbon and the organic carbon. Every sample was thus characterised (see VAN OSSELAER. 1992) by a reference number. a date and the 13 following parameters: locality. depth. colour. % total carbon, % organic carbon. % carbonates and the percentages of the seven textural classes (>2000 um. 2000-1000 p m. 1000-500 pm, 500-250 um, 250-125 um, 125-63 pm and < 63 pm). 3. RESULTS. 3.1. The classification of sediments. The data obtained on the sediment samples are given in Table 2. Multivariate analysis soon revealed that some order was hidden in that apparent chaos. Application of the classical U.P.G.M.A. (Unweighted, Pair-group. Method using arithmetic Averages) clustering method (see SNEATH & SOKAL, 1973) using squared Eucliduan distances to the matrix containing six textural classes. the percentage of organic carbon. the percentage of carbonates and depth vields the dendrogram shown in Fig. 3. Taking all seven classes into account would introduce redundancy, as the sum of the classes is necessarily 100%. Sediments are characterised mainly by either fine or coarse particles, so we elected to eliminate one of the 3 intermediate textural classes. Amongst these, the class 250- 125 um has the smallest contribution to the total variation (in the analysis of principal components on raw data) and was therefore discarded. It can be seen that the sediments fall into two clusters that are very clearly separated (over 50% of the maximal distance between groups). 31 APEX 9(2/3): 29-46, juil. 1994 One of these clusters corresponds to light- coloured coral sands. with coarse particles and a high carbonate content. Colours of dry samples of this group were: bluish white 5B 9/1. dusky vellow 5Y 6/4, gravish yellow 5Y 8/4, light olive gray 5Y 6/1. very pale orange I0YR 8/2, vellowish gray and yellowish gray 5T 7/2. The other cluster corresponds to dark terrigenous sediments with fine particles and low carbonate content. Colours of dry samples of this group were: dark greenish gray, dark greenish gray 5GY 4/1, grayish olive 10Y 4/2. greenish gray 5GY 6/1, light olive gray 5Y 5/2. olive gray. olive gray 5Y 3/2 and olive gray 5Y 4/1. For the sake of simplicity we shall not use the rich vocabulary available for sediments (see for instance COLLINSON & THOMPSON. 1989) and these groups will be here under referred to as "black substrate" and "white substrate". UP GMA. clustering works on distances in the attribute hyperspace and gives no information on the relative importance of the different factors under consideration. This was obtained by the equally classical FAC. (Factorial Analysis of Correspondences) method. first effected on all the factors considered in the analysis here above. The same analysis. performed with all textural classes (this hardly brings any modification in this case) 1s 1llustrated in figure +. It fully confirms not only the existence of the two groups "black" and "white" obtained by UP. GMA. but also the correctness of their interpretation. The principal axis (52.7 % of the total inertia) corresponds to the textural classes. ordered bv size. The least important factors are depth and organic carbon. The same analysis. effected without considering depth hardly shows anv modification. This might appear surprising at first sight but is quite logical because organic matter is related to depth. the role of which depends upon the nature of the sediment. This is confirmed by the observation that when F.A.C. are performed separately on each group of sediments (not illustrated here). the contributions of depth and organic carbon become apparent, especially in the case of black sediments. Very similar results were obtained by A.P.C. (Analysis of Principal Factors, not illustrated here). The clustering of sediments into two groups can even be evidenced in a much simplified representation (Fig. 5) in which all stations are reported on a scatter diagram of the carbonate content vs. depth. 32 Studies on Olividae. XVIII It must be stressed that this classification into two clear-cut, compact groups does not encompass all the sediments of Hansa Bay but onlv those in which Oliva specimens have been found. The two groups of sediments are most probably bridged by intermediate. deep water points, where O/iva have not been met with. 3.2. The distribution of species. The occurrence of the various Oliva species at the different stations. together with a brief description of their common habitat, is given in Table 3. 3.2.1. Correlation with depth. The distribution of the Oliva species in Hansa Bay is obviously related to depth, as can be seen on the graph of figure 6. Some species are restricted to deep water and some others to shallows. One will notice that the observed depth range of some species is rather extensive. Specimens of ©. amethvstina, ©. panniculata and ©. paxillus have exceptionally been collected at much greater depths than normal. These rare findings occurred only on very steep reef slopes and have not been reported in the graph. as they are most probably accidental (on such slopes. an unsuccessful attack by a predator could result in a considerable fall). With the exception of the widespread ©. carneola, wide depth ranges seem to be a feature of deep water species. The graph of figure 7 gives the number of species as a function of depth. The curve sharply culminates at 5m (where two thirds of the Hansa Bay species can be found) then shows a rapid. regular decrease. The distribution of Oliva species 1s certainly not a function of depth alone. If this would be the case, all shallow water species would be expected to coexist, which is not at all the observed situation. One should be aware that the correlation between depth and distribution 1s most probably indirect: Olives are not known to possess any pressure-sensitive anatomical structure and specimens can be rapidly brought to the surface from -70 m. then kept for extended periods in aquaria without any apparent disability. 3.2.2. Correlation with nature of the sediment. The distribution of each species was established by drawing contour lines around its points of presence both in the F.A.C. diagram and in the depth vs. carbonate scatter diagram. For the sake of space economy. only two species. Oliva parkinsoni and ©. reticulata will VAN OSSELAER, BOUILLON, OUIN, TURSCH VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olividae. XVII APEX 9(2/3): 29-46, juil. 1994 be given as examples. It can be seen (Figs. + and 5) that these two species have different distributions (this is also apparent on the UPGMA. Fig. 3). ©. parkinsoni is found only in a very restricted zone of “white” sediments. while ©. reticulata occupies disjunct portions of both "black" and “white” sediments. The following groups of species are observed: Species found on "white" substrate only: ©. amethystina (°). O. buelowi. O. caerulea (©), ©. concinna, ©. mantichora, ©. miniacea, ©. panniculata, ©. parkinsoni. ©. paxillus, ©. semmelinki, ©. sericea (°). ©. tesselata. The species marked (°) are found also in small patches of somewhat darker "white" coral sand (coloured with terrigenous sediments) formed around World War 2 wrecks lving in "black" sediment. Species found on "black" substrate only: ©. athenia. O. ceramensis. O. concavospira. O. sp. DHB. ©. dubia. ©. funebralis. O. mucronata. ©. rufula. O. sp. ZHB. Species found on both "black" and "white" substrates: ©. bulbiformis. ©. carneola. ©. elegans. ©. longispira. ©. reticulata, O. smithr, O. solomonensis. O. vidua. O. cfr. volvaroides. The species restricted to onlv one type of sediment will be here under referred to as stenotopic and the species occupving different types of substrate as eurvtopic. Experiments in aquaria (VAN OSSELAER & al. 1993) have shown that it is highly unlikely that restriction of habitat is caused by a choice of substrate by adult specimens. 3.3. Observations on colour patterns and crypsis. Oliva species are famous for the variability of the colour pattern of their shells. but this variability is not entirely haphazard. Our observations fully confirm the previous impressions that the colour pattern can vary with the substrate. Of the 30 Ofiva species collected in Hansa Bay 28 are cryptic (coloration and markings of the shell and the soft parts resemble the surroundings and aid in concealment). For instance ©. amethvstina and O. mantichora that live exclusively in coral sand in proximity to live reefs. are easily mistaken for fragments of dead Acropora coral. The crypsis of more colourful species like ©. semmelincki, ©. parkinsoni and the brightlv coloured ©. buelowi is not evident when specimens are seen in a drawer but quite convincing in the field: these species live in much deeper water, where the colours red and orange are seen dark brown and brown. One must also remember that the sediment is generally not uniform but contains debris and rubble of various sizes. On such a substrate. the reticulated or variegated pattern of many olive shells constitutes a most effective camouflage. One should note that in eurytopic species both the mantle and the shell are cryptic. The general aspect can vary strikingly within the same species (see PI. 1, Figs. 6 and 7). The two non cryptic species are ©. carneola and ©. rufula, which will be discussed later. There is an obvious (and probably continuous) variation in the "colour strategy" of the Oliva species in Hansa Bay. On the one hand. for most species every local micropopulation is cryptic and quite homogeneous in colour pattern. An experienced local collector can often guess the exact origin of a given specimen because the colour pattern is often characteristic of a given locality. On the other hand. local populations of a species can be extremely heterogeneous. Of 35 specimens of ©. concinna (found exclusively on white sand off Boro Beach) 25 (71%) were white. 9 were all black and 1 was orange. Of nearly one thousand ©. longispira specimens observed on the black beach at Sisimangum 76% were "black", 15 % were "white" and 9 % did not fit into these categories. In these populations. it is only the large mayority of the specimens that 1s cryptic. These are clear cases of population polymorphism. It is interesting to note that the "colour strategy" of a given species can differ from one locality to another. We have just seen that ©. longispira is polymorphic on the black sand of Sisimangum beach but 1t 1s not so on the white sand of Boro Beach where all of the +2 collected specimens of this species were "white". This can also be observed for ©. carneola. some populations of which are verv homogeneous while others are highly heterogeneous. Some of the homogeneous populations of this species are cryptic. others not. Intermediate strategies do occur. For instance. the colour pattern of the ©. vidua populations is always cryptic but very variable within à given micropopulation. 4. DISCUSSION. The Oliva species habitats reported here are based only upon observations in Hansa Bay. It is conceivable that the same species might occupy different habitats in other localities. The present data so far agree with the habitats we have observed in other localities in Papua New Guinea (Boesa I. Legoarant Is: Bogia. 33 APEX 9(2/3): 29-46, juil. 1994 Madang. New Ireland) and in other regions of the Indo-Pacific (Indonesia. Sevchelles. Sri Lanka, Thailand. Vietnam). 4.1. Crypsis. One might wonder what olives that are burrowing and nocturnal could gain from crypsis, a strategy obviously directed at diurnal predators endowed with good vision. But olives do come in full light when they detect prey at the sediment surface (they burrow back immediately after the capture of prev). As they do not bury deep (only a few centimetres. see VAN OSSELAER & al. 1993) they could also easily be exposed by any of the many digging or rummaging predators present in their biotope. Crypsis is then very efficient. especially when compounded with very fast burrowing (many a diving collector will recall a coveted Oliva specimen literally vanishing under his eyes). A strong argument for crypsis being due to predator pressure stems from the observation (see Plate 1) that crypsis is more convincing on the dorsal face and the mantle (the parts a predator is more likelv to see) than 1t is on the ventral face. In Hansa Bay. there are two exceptions to the generalised crypsis of Oliva species. The first is arguably ©. rufula (Fig. 8). restricted to deep. dark. very soft sediments. Although its background colour blends with its surroundings. its strikingly disruptive colour pattern could be interpreted as an aposematic (warning. protective) signal. indeed the very contrary of crvpsis. The pattern is indeed seen as very contrasting nearlv black-and-white 1f on a colour picture of ©. rufula one filters off the colours red and vellow (that do not reach the depth where the animal lives), This interpretation 1s tenable because we have often observed captive. stressed specimens of this species to produce a deep-green. highly toxic exudate. The second. quite obvious exception 1s the abundant ©. carneola (neariy ubiquitous from 0 to -30m). The populations of this very variable species are not homogeneous in coloration: those living on "black" sediment consist mostly of brownish (cryptic) specimens while populations from "white", shallow substrates consist very predominantiy in bright orange individuals. sharply contrasting with their surroundings. We still lack data on the possible defences of this form (which produces a bright yellow exudate of unknown toxicity) and have no interpretation to propound for this puzzling case. Studies on Olividae. XVII The Hansa Bay Oliva species display a large spectrum of "colour strategies" and strict analogy with any of the previous studies on shell colour variation (see Introduction) is not obvious. In our case the problem is even more complex because the colour pattern of Oliva specimens can vary during the lifetime of an individual, as attested by the abrupt colour pattern changes often observed on the shells of many species. Svnchronism of such colour pattern transitions in a population of the East African ©. bulbosa, has led GREIFENEDER (1984) to suggest that colour pattern changes could be used as a "chronicle of the habitat". The possibility that the shell colour of Oliva specimens depends upon food cannot be rejected at this point. The colour of the small bivalves that seem to constitute an important part of the diet of Oliva species 1s also often matched to the colour of the sediment. Oliva species could constitute an ideal experimental material for the study of shell colour variation. if one could solve the problem of raising their veliger larvae. These are verv easy to obtain but our first. crude attempts at raising them have so far been unsuccessful. In summary. crypsis in Oliva species is still far from being understood. but whatever its interpretation. the phenomenon is the rule rather than the exception in Hansa Bay. 4.2. Taxonomic consequences. The colour pattern of Oliva shells is a character that has been of paramount taxonomic importance and still constitutes a large part of contemporary species descriptions. Our observations call for some comments on the use (and possible abuse) of this character. as manv authors working on the genus Oliva seem completely unaware of the vast literature available on shell colour variation. In Hansa Bay. most species of Oliva are cryptic and many are eurvtopic. On the one hand. within the same eurytopic species. spécimens imitating very dissimilar habitats will acquire greatly different aspects. On the other hand. syntopic populations of different species will mimic the same substrate and will thus tend to resemble each other (see Plate 1). forming local assemblages at first sight reminiscent of Müllerian groups of mimics (but this concept should be restricted to aposematic and not to cryptic colorations). So crypsis has two consequences: divergence within eurytopic species and convergence between syntopic species. This can obviousiy cause much taxonomic confusion and crypsis is thus likely VAN OSSELAER, BOUILLON, OUIN, TURSCH | to have been a major contributor to the old "Oliva problem. The taxonomic value of gross colour features should be considered with great caution. In Hansa Bay. the overall colour pattern of an Oliva specimen (especially with depth correction for colours) often gives more information on the underwater aspect of its habitat than it does on its taxonomic status. This remark does not apply to features of the ventral face (not seen by the predator) or to small details (the patterns of the spire. the fasciolar band. the subsutural zone. etc.) that hardly affect concealment. Such features. being less adaptive, are more likely to constitute reliable identification characters. We see no reason why this situation should be restricted to the genus O/iva and one can expect crypsis to occur in other controversial groups of molluscs. with similar taxonomic consequences. Acknowledgements. This work was supported by the FRF.C. (grant n° 2.9008.90), the F.N RS. the Fonds Leopold III. the Fonds Lefranc and BIOTEC. S.A. The authors are grateful to Sid Johnson. Müillar Magap. and especially Jean Pierret for invaluable help in the field. We thank W Gruber. Prof. J. Herbauts. Prof. A. Herbosh, M. Bertrand and A. Preat for allowing the use of their facilities and giving advice for the analysis of sediments. We thank Dr. D. Greifeneder for discussing the manuscript and P. Willecomme for his kind help REFERENCES. ABBOTT, R.T. 1991. Those puzzling Olive shells. 4. Conch. 19(3): 23. ADAMKIEWICZ. L & M. CaASTAGNA. 1988. Genetics of shell color and patterns in the Bay * Scallop Argopecten irradians. J. Hered. 79: 14- IgÉ BOUILLON. J.. M. CLAEREBOUDT & G. SEGHERS. 1986. Hydroméduses de la baie de Hansa (Papouasie Nouvelle-Guinée): répartition. conditions climatiques et hydrologiques. /ndo- Malavan Zoology 3: 105-152. CAIN. A. J. & P.M. SHEPPARD. 1954. Natural selection in Cepaea. Genetics, 39: 89-116. CLAEREBOUDT, M. 1989. Répartition spatiale et diversité des Scléractiniaires sur un récif corallien de Papouasie Nouvelle Guinée. Thèse de doctorat. Université Libre de Bruxelles. VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olividae. XVIII APEX 9(2/3): 29-46, juil. 1994 CLAEREBOUDT. M. CL. MASSIN & J. BOUILLON, 1989. A general survey of Laing Island environment (Papua New Guinea). /ndo- Malavan Zoology, 6: 1-23. COLE, T.J.. 1975. Inheritance of juvenile shell colour of the oyster drill Urosalpinx cinerea. Nature 257: 794-795. COLLINSON, J.D. & D.B. THOMPSON, 1989. Sedimentary structures. Unwin Hyman Ltd. London. 207 pp. COLTON, H.S.. 1916. On some varieties of Thais lapillus in the Mount Desert region, a study of individual ecology. Proc. Acad. nat. Sci. Philad. 68: 410-154. CREESE, RG. & A.J. UNDERWOOD, 1976. Observations on the biology of the Trochid Gastropod Austrocochlea constricta (Lamarck) (Prosobranchia). I. Factors affecting shell- banding pattern. /__ exp. mar. Biol. Ecol. 23: 211-228. ELEK. J.A & S.L. ADAMKIEWICZ. 1990. Polymorphism for shell colour in the Atlantic Bay Scallop Argopecten irradians irradians (Lamarck) (Mollusca: Bivalvia) on Martha's Vinevard Island. 4. Nalac. Bull. 7(2): 117- 126. ETTER. R.J. 1988. Physiological stress and colour polymophism in the intertidal snail Nucella lapillus. Evolution 42(4): 660-680. GIESEL. JT. 1970. On the maintenance of a shell pattern and behavior polymorphism in Acmaea digitalis. a limpet. Evolution 24: 98- 119. GREIFENEDER. D. 1981. What do we know about Olividae. Contributions to the study of Olividae. Acta Conchvliorum 1: 1-90. GREIFENEDER. D. 1984 Die Farbmuster von Oliva-Gehäusen. Club Conchvlia 5/6: 53-65. HAMLYN-HARRIS. A.G.. 1970. Oliva rubrolabiata. Hawaii. Shell News 18(7): 5. HEMMEN, JD. 1981. Olividae of Jaco (Costa Rica) and Aruba (Ned Antilles) Acta Conchvliorum 1: 128-130. INNES, D.J. & L.E. HALEY: 1977. Inheritance of a shell-color polymorphism in the mussel. Z Hered. 68: 203-204. INO, T.. 1949. The effect of food on growth and coloration of the Topshell (7urbo cornutus Solander).J. mar. Res. 8(1): 1-5. JONES. GP. D.J. FERREL & P.F. SALE. 1990. Spatial pattern in the abundance and structure APEX 9(2/3): 29-46, juil. 1994 of mollusc populations in the soft sediments of a coral reef lagoon. \/ar. Ecol. Progr.: Series 62: 109-120. JONES.J.S.. B. H. LEITH & P. RAWLINGS. 1977 Polymorphism in Cepaea: a problem with too many solutions ? .1nn. Rev. Ecol. Svst. 8: 109- 143. LAMOTTE., M.. 1959. Polymorphism of natural populations of Cepaea nemoralis. Cold Spring Harbor Svmp. Quant. Biol. 24: 65-86. LEIGHTON, DL. 1961. Observations on the effect of diet on shell coloration in the Red Abalone. Haliotis rufescens Swainson. The Veliger 4(1): 29-32. MITTON. J.B.. 1977. Shell color and pattern variation in A/vtilus edulis and its adaptative significance. Chesapeake Sci. 18(4): 387-390. MOMENT, G.B. 1962. Reflexive selection: a possible answer to an old puzzle. Science 136: 262-263. MOORE. H.B.. 1936. The biology of Purpura lapillus. I. Shell variations in relation to environment. J. \{ar. biol. Assoc. UK. 21: 61- 89. MURRAY. J.D. 1962. A pre-pattern formation mechanism for animal coat markings. Z_Theor Biol. 88: 161-199. OWEN, DF. & JC. REID. 1986. The white snails of Africa: the significance of Man in the maintenance of a striking polymorphism. Oikos 46(2): 267-269. PALMER. A.R. (1985). Genetic basis of shell variation in 7hais emarginata (Prosobranchia. Muricacea). I. Banding in populations from Vancouver Island. Biol. Bull. 169: 638-651. PETUCH, E.J. & DM. SARGENT, 1986. Atlas of the Living Olive Shells of the World., CERF editions, Charlottesville, VA. 253 pp. REIMCHEN, PE” 1979. Substratum heterogeneitv, CTYPSIS, and colour polymorphism in an intertidal snail (Littorina mariae). Can. J. Zool. 57: 1070-1085. SNEATH, P.H. & R.R. SOKAL. 1973. Numerical taxonomy. WH. Freeman and Co, San Francisco, 573 pp. TURSCH, B.. L. GERMAIN & D. GREIFENEDER. 1986. Studies on Olividae. IV. Oliva annulata Gmelin. 1791 (of authors) : a confusion of species. /ndo-Malavan Zoology 3 : 189-216. 36 Studies on Olividae. XVII TURSCH. B. ©. Missa & J. BOUILLON. 1992. Studies on Olvidae XIV. The taxonomic structure of Oliva oliva (auct.). Apex 7(1): 3-22. VAN OSSELAER. C.. 1992. Contribution à l'étude écologique du genre Oliva (Mollusca. Gastropoda) à Hansa Bay (Papouasie Nouvelle- Guinée). Travail de fin d' Etudes. Université Libre de Bruxelles. VAN OSSELAER. C., J. BOUILLON & B. TURSCH. 1993. Studies on Olividae XVII. Data on depth of burrowing, motion and substrate choice of some Oliva species. Apex 8(4): 151-158. WIDMER., M., 1981. Olividae of Dar es Salaam. Acta Conchvyliorum 1: 115-127. WITTIG-SKINNER, R., 1981. Olividae of Indonesia. Acta Conchvliorum 1: 91-114. VAN OSSELAER, BOUILLON, OUIN, TURSCH VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olividae. XVII APEX 9(2/3): 29-46, juil. 1994 Map coord. Station Map (diving) | coord. (m) Station (diving) H5/b Station (diving) Al A2 J4/b 3 A3 J4/b 2 At J4/b il AS J4/b 6.5 A6 J4/b 65 A7 J4/b 4.5 A8 H/b 2 A9 U1/b 0 A10 12/7 10 All K3/a 16 A12 Z7/b 9 A13 Z7/b A14 Z7/b 5 A15 F3/b 20 A16 S2/d 5 A17 N3/d 25 A138 N4/a 12 A19 H5/d 0.5 A20 T2/a-c + A21 T1/d 4.5 A22 V1/b 7 A23 T1/b 0.5 A2+4 Z5/b (Dredging) | coord. (Dredocing) | coord. (m) D1 FL) VI 5 D2 Z+ D3 UI 3 D{ jiil 3 DS T1 3 D6 UI 3 D7 UI 5 D8 UI1 10 D9 U2 15 D10 2 10 D11 d2 VI Table 1. Identification of sampling stations. Coordinates refer to maps figures 1 and 2, where each surface unit was subdivided into a (upper left), b (upper right), c (lower left) and d (lower right). 37 APEX 9(2/3): 29-46, juil. 1994 Studies on Olividae. XVII Granulomet m DEEE SEE 2 >1000 >500 >250 >63 [1069 | 3128 | 3847 | 1050 | 616 | 205 | oss | | 12601-24250") 23842) 03020 NL 60) | Re ne BC RTS RON RE CN EE ET CE D _o19 | ose | 576 | 874 | 2882 | 3404 | 2159 | | =020 «| "561. 5:55 20199410 /ns67 1er [ 005: ©], or [076 | © 635, 1[E 5300 1je55.96 | Eloice13] | 020’ [som | ORNE PSS TE Ne | TON BR SOC NON EEE EC C'o47 6097 so 1937 6310) Moss |Bu0 00] DOTE NET OS CON ES ES [0031 080 7 sp ns 20 IR 200 | | = PEN RENTE TS RCE EN TN ET [_030 | 067 | 233 | 186 | 1017 | 7616 | 851 | D 52 [_009 | 008 | 034 |! su |essss | 3883 ru | [_o18 | o26 | 187 | 315 | 2338 | 5510 | 1607 | [_000 | 012 |" 135 | 335 | 3508 15270 |740, [_000 | 004 | 028 | 491 | 1742 | 4810 | 2926 | [_000 | 002 | 043 | 161 | 2323 | 4826 | 2615 | [_2807 | 1968 | 4763 | 268 | 116 | o68 | o10 | PRET CNT CET TE CN ETC EE ON TC [_oow | oi | #o36 | 038 | 376 | 6891 | 2647 | [_o00 | oo | o19 | os | 5031 | 402 | 00 | [_4987 | 2094 | 2011 | 376 | 35 [nc 005%) [_3707 | 710 | 798 | 582 | 2594 | 1608 | 000 | [_2960 | 1384 | 2211 | 1748 | 1332 | 3.4 |" oo! BNC TON CET NE A LR PTE RE Table 2. Characteristics of sediment samples. 38 Carbon analysis 93.18 | 010 | 001 | [ o11 | 000 | BOT | 016 | %.04 | [ 010 | 1428 | [ 016 | 9140 | [ 015 | os | [_ 014 | 109 | [ 011 | 094 | [ 018 | 8300 | [ 006 | 087 | [| 006 | 066 | [ o19 | 9186 | ETS EUX VAN OSSELAER, BOUILLON, OUIN, TURSCH Substrate group White While White White White White White Black Black White White White White White Black White White Black Black Black Black Black White White Wlute White Black White Black White Black Black White Black Black White White Black White White White Black Black Black Black Black Black Black White Black Black Black White Wlhute Black Black Black White White White White Black Black Black VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olividae. XVIII APEX 9(2/3): 29-46, juil. 1994 HABITAT EXAMPLES coral sand near living coral, mostlv 1-10m. A2, A3, A4, A8, A28, A3, A60 5. SPECIES amethystina Also around wrecks athenia "black" sediment, 5-10m. D13, DI4 buelowi coral sand, bottom of reets, 20-50m. A32 bulbiformis "white" or "black" sediment, 1-8m (shallow only in quiet | Al14, A24, A35, A42, D3. waters). "black" or "white" sediment, 0.5-30m (not in very agitated shallow waters). A2 to A8, AIO, AIS to A19, A21, A22, À 26 to A28, A36, A40, A42 to A45, A47, AS1 to AS3, ASS to A58, A60, A63, A64, A70, D7 to D9, DI4, DI16, D20. A10, A29, A31, A40, ASI to AS3, ASS, A56, AS8, A61, D8 to DI0, D19, D20, D24. Al, A19, A63, A64, A76, A77, A78 carneola ceramensis "black" sediment, 10-31 m (generally 18-25m). coral sand, 0.5-3 m in Laing I. lagoon, 6m on Duranagit Reef. "black" sediment, 8-22 m (generallv 12-18m) caerulea concavospira A56, AS8, A62 concinna "white" sediment, 5-10 m (off Boro Beach onlv). Al4, A35. "black" sediment, 3-12 m A13, A16, A20, A33, A45, A56, A58, A7O, D12 to D15, D21, D23 very fine "black" sediment, 40-60 m. "black" sediment, 0.5-7 m near Sakula and Aivar Rivers; dubia elegans "white" sediment (0.5-1 m) at Mandi Beach unebralis "black" sediment; 3-7 m A70 “black” or "white" sediment, surf-exposed beaches onl\ longispira mantichora coral sand near living coral, mostly 20-40 m miniacea coral sand, 10-18 m. Rare in Hansa Bay mucronata "black" sediment, 5-8 m, mostly near Sakula River panniculata coral sand, 6-20 m, agitated water (top of Durangit Reef). parkinsoni coral sand near reef, 12-42 m. Al1, A26, A27, A30, A32, A38, | © A39, AS4, A7S paxillus coral sand, 6 m, agitated water. top of Durangit Reef. A13, A19, A37, A4], A44, A45, A57, A63, A64, A76, A77, A78, DI. rufula "black" sediment, 18-35 m (mostly 25-35m). A29, A31. sericea coral sand, mostlv 1-10 m. Also around wrecks A35, A63, A64 semmelincki coral sand, bottom of reefs, 35-70 m. smithi "black" or "white" sediment, 0.5-12 m. Al, A12,A13, A14, A35, A45, A46, A57, A70, DI to D6, DII, D12, D15, D21. solomonensis "black" or "white" sediment, 5-10 m. A14, A35, A43, D13, DI4. tesselata coral sand near reef, 3-7 m, only in lagoon. vidua "black" or "white" sediment, 0.5-12 m. Al, A12, A13, A34, A45, A7O, D3, D13, DI4, DI16, D17, D18, D20. Cfr. volvaroides "black" or "white" sediment, 6 m. Very rare in Hansa A22, A35. Bay. ZHE on Fond ar one Loc En Table 3. Brief notes on habitat of Ofiva species, with collecting stations in February-March 1992. "black" or "white" sediment, 0.5-12 m. reticulata 39 APEX 9(2/3): 29-46, juil. 1994 Studies on Olividae. XVII VAN OSSELAER, BOUILLON, OUIN, TURSCH Plate 1. (opposite) . Oliva bulbiformis. Laing Island lagoon. 1 m, "white" substrate. . Oliva. solomonensis. Off Boro Beach. 5 m, “white” substrate. . Oliva caerulea. Laing Island lagoon. 0.5 m, "white" substrate. . Oliva concinna. Off Boro Beach. 6 m, “white” substrate. . Oliva elegans. Mandi Beach. 0.5 m, "white" substrate. . Oliva reticulata. Laing Island lagoon. 1 m, "white" substrate. . Oliva. reticulata. Off Sisimangum. 5 m, "black" substrate. . Oliva vidua. NE of mouth of Sakula River. 5-7 m, "black" substrate. . Oliva sp. DHB (see text, section 2.1). NE of mouth of Sakula River. 5-7 m, "black" substrate. 10. Oliva athenia. NE of mouth of Sakula River. 5-7 m. "black" substrate. 11. Oliva elegans. NE of mouth of Sakula River. 5-7 m, "black" substrate. 12. Oliva reticulata. NE of mouth of Sakula River. 5-7 m, "black" substrate. © © —J O O1 BR CO) MN D — 40 VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olhvidae. XVIII APEX 9(2/3): 29-46, juil. 1994 41 APEX 9(2/3): 29-46, juil. 1994 Studies on Olividae. XVIII VAN OSSELAER, BOUILLON, OUIN, TURSCH D A 2 + eDURANGIT Ar £ ee Reef 3 À Fig. 1. Hansa Bay. Black circles represent locations of sediment samplings at sites where Oliva species have been collected during February-March 1992. Dredgings are represented by thick gray lines. Stars represent coral reefs. VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olhividae. XVII APEX 9(2/3): 29-46, juil. 1994 e D j 22: hé “> ten UNE T CRE ETS RAT TP a x Fig. 2. Laing Island. Black circles represent locations of sediment samplings at sites where Oliva species have been collected during February-March 1992. Stars represent the limit of coral reefs emerging at low tide. APEX 9(2/3): 29-46, juil. 1994 Studies on Olividae. XVII VAN OSSELAER, BOUILLON, OUIN, TURSCH 12000 10000 2 = = E 7 3 = 8000 £ = == — 2 An v eo” = o Z = C Z = $ = = D 6000 == a = = LU] Le] q B4 C 3 4000 2000 ô ———— RS ES ER D EE en RE OS 29221755 21712414222144277747142< 172222227227 <<72771<22< << © parkinsom P Pp£rPpoP P © reticuiata r r r r (A < Co TAC raRrar Fig. 3. U.P.G.M.A clustering (squared euclidian distances) of sediment samples. Variables: six textural classes (>2000 um, 2000-1000 um, 1000-500 um, 500-250 um, 125-63 um and < 63 um), percentage of organic carbon, percentage of carbonates and depth. The disjunct distributions of Oliva reticulata (r) and ©. parkinsoni (p) are shown below the dendrogram, as an example. : AXI ÆZ. DIT... Are. ROIDTS A AN BLACK WHITE De LR. D Era © == Vs A D c SL = NES 72 ns" ——_. LR En — SES SN A L > Fé ' \ \ ÿ À À se TS 5 1 CE PET F «€ ., ” e EXC LI PQ A Sie» NS Ces «. r ' UC" TRES Car 30 res : RS ax: RER PARL) TT mn 288 : ES _ 7 32 QSON.. = PARA pars [ - à O Pi OEE FE ! D = #2: ss 34, 3 “ t LP 2 A 0, rt), - CA mn "x : 24 st — . 416 — — © Fig. 4. F.A.C. (Factorial Analysis of Correspondences) analysis of sediment samples. Variables: seven textural classes [>2000 um (g), 2000-1000 um (f), 1000-500 um (e), 500- 250 um (d), 250-125 um (c) 125-63 um (b) and < 63 um (a)], percentage of organic carbon (OC), percentage of carbonates (CA) and depth (D). The disjunct distributions of Oliva reticulata and ©. parkinsoni are shown as an example. point point seen hidden CA 4 5 6 12 13 20 23 ++ point point point point seen hidden seen hidden g 28 3 63, 64 22 36 24 70 32 39 46 71 20 47 f 76 depth (m) 70 60 50 40 30 20 10 VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Olividae. XVII APEX 9(2/3): 29-46, juil. 1994 depth (m) EI] 40 + P Il S'S à 30 + £ Black substrate PF White substrate Per 25 " n P RO | a Tiré P: 15 + P lR2 n 10 : 2 RR ee ë 5 SEE or ce) R K ES e [e) GS > + ; +—£ FRPK 0 10 20 30 40 50 60 70 80 90 100 % CaCO; Fig. 5. Scatter diagram of % carbonates vs. depth. The two groups of sediments are again separated on this simplified representation. The disjunct distributions of Oliva reticulata (R) and ©. parkinsoni (P) are shown as an example. [e) 1 [er] (a) [e [æ) ] Lai :) dubia mm 2HB@ DHB Sith 8 © | 4 Vidua DB 0 | elegans BO | D semmelinki © buelowi mantichora rufula w parkinson: ceramensis minacea paxillus athenia & concinna mucronala BB funebralis & tesselata sericea renculatam © |+————— y caerulea —— + (l Al ! ns | 24 C—— | x] Conan | Le | C bulbiformis @ 0 | + camelaB Q |——————————————_—_—————————4 concavospira panniculata “volvaroides"@ © solomonensis @ amethystina Fig. 6. Observed bathymetric range of Oliva species in Hansa Bay. Data from 1974 to 1993 (see text section 3.2.1). Black and white squares indicate occurence in "black" and "white" sediments, respectively. longispira @ © APEX 9(2/3): 29-46, juil. 1994 Studies on Olividae. XVII VAN OSSELAER, BOUILLON, OUIN, TURSCH Fig. 7. Number of Oliva species in Hansa Bay as a function of depth. Fig. 8. Aposematic pattern of the toxic species Oliva rufula (trapped NW of Laing I., 35 m, "black" substrate). Scale bar: 10 mm. 46 VAN OSSELAER & TURSCH The suture of Oliva APEX 9(2/3): 47-S0, juillet 1994 Studies on Olividae. XIX. Where is the suture of Oliva shells? C. VAN OSSELAER and B. TURSCH Laboratoire de Bio-Ecologie, Faculté des Sciences, Université Libre de Bruxelles, 50 av. FD. Roosevelt, B-1050 Brussels, Belgium. ABSTRACT. The suture of Oliva shells is hardly perceptible and has no functional link with the so-called "sutural channel", for which the more descriptive name "filament channel" 1s proposed. RESUME. La suture des coquilles d' Oliva est à peine perceptible et n' a pas d'association fonctionnelle avec le soi-disant "canal sutural", pour lequel le nom plus descriptif de "canal du filament" est proposé. KEY WORDS: Oliva, shell, suture, channel, posterior filament, filament channel. 1. INTRODUCTION. AIl authors agree broadly on the definition of the suture of coiled shells. This is for instance: "fhe line of contact where two whorls meel" (FRETTER & GRAHAM, 1962). or ‘a continuous line marking the junction of whorls in a gastropod shell" (ARNOLD, 1965), or "the junction of each whorl against the other" (ABBOTT, 1974). The spire of all Oliva shells displays a wide, conspicuous spiral channel that LAMARCK (1811) called "/e canal de la spire". In their descriptions of the general characters of the genus Oliva, FISCHER von WALDHEIM (1807), MONFORT (1808), LAMARCK (1811) and later DUCLOS (1844) all repeated verbatim the same expression: "fours de spire séparés par un canal". If the whorls were "separated by a channel", then it was quite logical to associate the channel with the suture and DUCLOS (1844) indifferently used "canal", "canal spiral" and "canal sutural". From there on, the habit was established and until today nobody questioned the appropriateness of the channel-suture association. For instance, one finds the terms: "groove on the suture" in GRAY (1842), "sutural canal" in MARRAT (1871) and PETUCH & SARGENT (1986); "sutural channel" in TURSCH & GERMAIN (1985); "suture canaliculated" in TRYON (1883): "channeled suture" in ZEIGLER & PORRECA (1969), ABBOTT (1974) and KANTOR (1991); "suture" in KANTOR (1991), TURSCH & VAN OSSELAER (1987), VAN OSSELAER & TURSCH (1988), "open suture" 1n GREIFENEDER (1981). Amongst gastropods, the "channeled suture" is found only in the family Olividae where it is a hallmark of the genera Oliva, Olivella, Olivancillaria and Agaronia. It has been shown to constitute an operational taxonomic character in the genus Oiva by TURSCH & GERMAIN (1985), TURSCH & VAN OSSELAER (1987) and VAN OSSELAER & TURSCH (1988). The "channeled suture" seems to be a very important feature, as it 1s always maintained open (at least on nearly one full volution) even in the many species where the spire is covered with a thick callous layer. The channel 1s also present in freak specimens. It 1s a very old feature, clearly displayed in the oldest Oliva shells (such as the Miocene fossil ©. dufresnei Basterot, 1925). The "channel" appears right after the protoconch transition and is already present in very juvenile specimens. One would predict that a structure so carefully preserved both in phylogeny and in ontogeny has to be functional, but what could that function be? From early days, the channel was related to a peculiar organ, the posterior Jilament. The first observation we could trace was in QUOY & GAIMARD (1834): "/e manteau … se termine en arrière par un filament plus ou moins long qui se loge dans le canal tout particulier que forment les sutures de la spire". GRAY (1842) vwrites of ‘"fhe thread-like elongation at the hinder angle, which forms the groove on the suture". TRYON (1883). in the general characters of the subfamily Olivinae. 47 APEX 9(2/3): 47-50, juillet 1994 writes: "an appendage behind which reposes in the channeled suture". In OLSSON (1956). one finds: "/n Oliva, the channel in the suture is maintained open and deep by a slender, tail- like appendage attached to the back of the mantle. This appendage lies along the channel when the animal is expanded but is lifted out as the soft parts are pulled back in the shell. Its real purpose is unknown". KEEN (1971) writes: "The mantle edge also has an unusual threadlike extension that lies along the suture, called a filament, which probably has a sensory function". 2. OBSERVATIONS 2.1. Observations of shell sections. Examination of polished longitudinal sections of the shell of several Oliva species brought unexpected results. Figure 1 shows the part of the shell that will be examined here. The two external crystal layers of each whorl are very easily recognised and allow easy localizing of the external boundary of each whorl. In all the sections (figures 2. 3 and 4) one can see in each whorl that the channel (c) lies in the third. cross-lamellar layer, counting from outside. On the sections of Oliva reticularis Lamarck, 1811 (Fig. 2) and of ©. reticulata (Rôding. 1798) (Fig. 3) the suture (s) is clearly separated and well above the channel (c). For the latter, one should note that the suture now lies close to a channel, but it 1s the channel of the preceding whorl | This is the most usual case amongst Oliva. Albeit easily detectable on polished cuts, the real sutural line of these Oliva is nearly invisible on the intact shell, even under magnification. The location of the suture cannot be guessed at by changes of coloration on the whorls of the spire: these are generally due to variations in the thickness of the outer layer. An interesting and common case is that of some heavily calloused shells. such as the specimen of Oliva carneola (Gmelin. 1791) illustrated in Figure 4. The channel is still distinct from the external boundary of the preceding whorl, as in the previous examples. But in this shell, every volution entirely covers the whole spire and there is no external line marking the separation between consecutive whorls. In such cases the common concept of suture (as a continuous line marking the junction of whorls) does not make sense. 48 The suture of Oliva 2.2. Observations on live specimens The posterior filament has been routinely observed for about thirty species of Oliva of which we have studied the live animal. For every species, the filament (when extended) could be seen lying inside and along the spiral channel, as schematically depicted in Figure 5. This fully confirms the relationship described by earlier authors. A clear sketch of the positioning of the posterior filament of Olivella biplicata has been published by BURCH (1988). One should note that the posterior filament is not always obvious because in many species it 1s nearly translucent. 3. DISCUSSION Our observations show clearly that the spiral channel of Oliva shells is completely distinct from the suture. The real suture is never "canaliculated" or "channeled" and is hardly visible on the shell. The classical names "sutural canal", "sutural channel" or "suture" that have been applied to the channel are misleading and have to be replaced. Because of its obvious association with the posterior filament, we suggest that it be named the "filament channel". The shape of the transversal section of the filament channel differs from species to species and several examples have been 1llustrated in TURSCH & VAN OSSELAER (1987). These features can be utilised as operational taxonomic discriminants (see TURSCH & VAN OSSELAER, 1987 and VAN OSSELAER & TURSCH. 1988). The function of the channel is most probably that of a protecting sheath for the posterior filament. The function of the posterior filament itself still remains a mystery, in spite of the anatomical study of MARCUS & MARCUS (1959). The possibility of the filament being a sensory organ was raised by KEEN (1971). BURCH (1988) reports that the posterior filament senses water currents. Several alternative hypotheses (among which the production of chemical messengers) could be considered. Work on this subject 1s being pursued in this laboratory for some vears but no firm conclusion has been reached so far. The exact function of the filament 1s a fascinating problem but has no bearing on the conclusion reached here above, 1.e. that the channel 1s not related to the suture. This case is à fine demonstration of the necessity of checking old postulates. The assumption that the channel was related to the VAN OSSELAER & TURSCH VAN OSSELAER & TURSCH suture was so logical that for nearly two centuries it was never questioned by any of the many students of Oliva. Ironically, this was also the case for the authors of this paper, who performed numerous, detailed measurements on a feature (the filament channel) that was erroneously called "the suture". The name of the character does of course not affect the taxonomic applications (TURSCH & GERMAIN, 1985; TURSCH & VAN OSSELAER, 1987; VAN OSSELAER & TURSCH, 1988) for which these measurements were proposed. Acknowledgements. We thank Mr. G. Bernardinis (Department of Geology) for his kind help in preparing the shell sections and Mrs. N. Van Mol (Department of Animal Biology) for the line drawings. REFERENCES ABBOTT, RT.,1974. American Seashells. Van Nostrand, N.Y. 663 pp. ARNOLD, WH. 1965. A Glossary of a thousand-and-one terms used in conchology. Veliger 7 (Supplement): 1-50 BURCH, BL. 1988. Olividae and its posterior tentacle. Hawaiian Shell News 36(8): 12. DUCLOS, P.L., 1844. Oliva in J.C. CHENU, Illustrations Conchyliologiques. Paris. FISCHER von WALDHEIM, G., 1807. Museum Demidoff ou Catalogue systématique et raisoné des Curiosités de la Nature et de l'Art. Données à l'Université Impériale de Moscou etc. Moscow. FRETTER, V. & A. GRAHAM, 1962. British Prosobranch Mollusces. Ray Soc. Lond. 755 pp. GRAY, M.-E. 1842. Figures of molluscous animals, selected from various authors. Etched for the use of students. London. GREIFENEDER. D. 1981. What do we know about Olividae. Contributions to the study of Olividae. Acta Conchyliorum., 1: 1-90. KANTOR, Y., 1991. On the morphology and relationships of some oliviform gastropods. Rufhenica 1(1-2): 17-52. The suture of Oliva APEX 9(2/3): 47-50, juillet 1994 KEEN, M. America. 2nd edition. Press. 1064 pp. 1971. Seashells of tropical West Stanford University LAMARCK. JBPA. 1810-11. Détermination des espèces de Mollusques testacés continuation du genre Ovule, Tarrière, Ancillaire et Olive. Ann. Mus Hist. Nat. 16 (for 1810): 300-328. Paris (Jan. -Mar. 1811). MARCUS, E. & E. MARCUS. 1959. Studies on Olividae. Bolm. Fac. Filos., Ciénc., Univ. S Paulo, 232 (Zool.22): 96-188. MARRAT, FP., 1870-71. Monograph on the genus Oliva Bruguière. In G.B. SOWERBY, Thesaurus Conchyliorum, 46 pp. MONTFORT, D. de, 1808. Conchyliologie systématique, et classification méthodique des coquilles. Vol. 1. Paris. OLSSON, A.A., 1956. Studies on the genus Olivella. Proc. Acad. Nat. Sci. Philad. 108: 156-225. PETUCH, E.J. and DM. SARGENT, 1986. Atlas of the Living Olive Shells of the World. CERF editions, Charlottesville, VA. 253 pp. QUOY, JRC. & JP. GAIMARD, 1834. Voyage de découverte de l'Astrolabe executé par ordre du Roi … sous le commandement de M. J. Dumont d'Urville. Zoologie. Tome troisième. Tastu. Paris. TRYON, GW. 1883. Manual of Conchology. Vol. 5. Marginellidae, Olividae, Columbellidae. Philadelphia. 267 pp. TURSCH, B. & L. GERMAIN. 1985. Studies on Olividae. L A morphometric approach to the Oliva problem. Indo-Malayan Zoology 2: 331- 352. TURSCH, B.& C. VAN OSSELAER, 1987. Studies on Olividae. VI. Suture measurements as taxonomic characters in the genus Oliva. Apex 2 : 69-84. VAN OSSELAER. C. & B. TURSCH, 1988. Studies on Olvidae. IX. Ten additional suture characters for Oliva taxonomy. Apex 3(4) : 81- 87. ZEIGLER, R.F. & H.C. PORRECA, 1969. Olive Shells of the World, Rochester Polychrome Press, Rochester, N.Y., 96 pp. 49 APEX 9(2/3): 47-50, juillet 1994 The suture of Oliva VAN OSSELAER & TURSCH ins K Fig. 1. Plane of the observed cuts in Ofiva shells. Fig. 2. Section of Ofiva reticularis Lamarck, 1811. s: suture. c: filament channel. LWh: last whori. Scale bar: 1 mm. Fig. 3. Section of Oliva reticulata (Rôding, 1798). s: suture. c: filament channel. LWh: last whorl. Scale bar: 1 mm. Fig. 4. Section of Ofiva carneola (Gmelin, 1791). s: suture. c: filament channel. LWh: last whorl. Scale bar: 1 mm. Fig. 5. Schematic view of a portion of the posterior filament lying in the filament channel. In reality, the filament (represented in black for the sake of clarity) is nearly translucent in many species. Îts relative size has also been exagerated. 50 TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 Studies on Olividae. XX. The pre-Lamarckian names for Oliva species Bernard TURSCH, Ralph DUCHAMPS (°) and Dietmar GREIFENEDER (°) Laboratoire de Bio-Ecologie, Fac. Sci. Université Libre de Bruxelles, 50 av. F.D. Roosevelt, 1050 Brussels, Belgium. (©) research associate ABSTRACT. The nomenclatural status of each of the Oliva names introduced by Linnaeus (1758), Born, (1778), Schrôter (1782), Lightfoot (1786), Abel (1787), Karsten (1789), Gmelin (1791), Rôding (1798), Link (1807), Fischer (1807), Montfort (1808) and Perry (1811) has been re-examined. Of the 88 names proposed by these authors 14 have been retained as valid Oliva names. We have been compelled to replace two well known names: Oliva vidua (Rôding, 1789) by ©. nigrita (Karsten, 1789) and ©. tessellata Lamarck, 1811 by ©. olivacea (Karsten, 1789). RESUME. Le statut nomenclatural des noms d' Oliva introduits par Linné (1758), Born, (1778), Schrôter (1782), Lightfoot (1786), Abel (1787), Karsten (1789), Gmelin (1791), Rôding (1798), Link (1807), Fischer (1807), Montfort (1808) and Perry (1811) a été réexaminé. Des 88 noms proposés par ces auteurs, 14 ont été retenus comme noms valides pour des Oliva. Nous avons été forcés de remplacer deux noms bien connus: Oliva vidua (Rôding, 1798) par ©. nigrita (Karsten, 1789) et ©. tessellata Lamarck, 1811 par O. olivacea (Karsten, 1789). KEY WORDS : Mollusca, Gastropoda, Olividae, Oliva, nomenclature. INTRODUCTION sources was thus deemed necessary. A chronological approach 1s clearly the simplest Is a review of the old Oliva names still way of automatically detecting junior necessary”? It is true that nearly all the names analysed here are well known to recent authors and can be found in the indexes of popular publications. The problem is that many of the attributions are uncritical citations, mainly from the pioneering works of BURCH & BURCH (1960, 1967). This can be demonstrated for instance by the ubiquitous use of aurata (Rôding., 1798), a very obvious romen nudum, to designate à variety of the species known as Oiva vidua (Rôding, 1798). There is simply no way of guessing what aurata might be and it is obvious that the original description has not been checked. frequent, as will be seen hereunder. Both factors Divergent attributions are also causes of the well-known nomenclatural confusion in the genus Oliva. A are major critical, de novo analysis of all the original homonyms and synonyms. Some of the points discussed in this paper might appear to be insignificant details. For instance, 1s it really important to know if the name given to a "bad species" is a zomen nudum or a nomen dubium”? The matter is actually of importance and does shape the subsequent nomenclature, because a nomen nudum remains available for future re- use. The identification of the species described by the old masters 1s straightforward only when the type material has been preserved. For Oliva, this is the case only for LINNAEUS (studied by OLSSON & DANCE, 1966) and FISCHER von WALDHEIM (studied by IVANOV & KANTOR, 1991). For all the problems are the rule rather than the exception. others, identification 51 APEX 9(2/3): 51-78, juillet 1994 The descriptions of the pre-Lamarckian authors are frequently insufficient for positive identification. They are nearly always extremely short and often based upon characters (such as the colour pattern) that are known today to be highly variable in the genus Oiva. We doubt that even a single species could be unambiguously recognised by its description alone. Fortunately, the authors do mostly (but not constantly) refer to previously published illustrations, which then constitute "indications" in the sense of art. 12 b (7) of the Code. The principal sources of these illustrations (amounting to over 90% of the total) are ADANSON (1757), d'ARGENVILLE (1742), BONNANI (1681. 1709), GUALTIERI (1742), KLEIN (1753). KNORR (1760-73), LISTER (1682- 1695), MARTINI (1769-1795$), PETIVER (1767). RUMPHIUS (1705), SEBA (1734-65) and SCHRÔTER (1782, 1783). Some of these figures are of very high quality and can be interpreted without ambiguity. Many others (quite rightly qualified as "medieval cartoons" by irreverent young students) are entirely unidentifiable. In some cases one may even doubt that the figure depicts an actual specimen. Problems of image identification are compounded by the fact that the concept of species of ancient authors was quite different from ours. This is evidenced by the very common use of conflicting illustrations in support of a given name and the use of the same illustration for different names by the same author. Let us cite BURCH & BURCH (1967): "1 is tragic that we are compelled to abandon such solid material (note: this refers to names supported by type material) and accept references to a series of poorly drawn old wood cuts. Typical of these are some of Rôding in the notorious Museum Boltenianum in which for some, Rôding lists as many as four references all to entirely different species, some of them unrecognizable, and the actual shell has been sold as a curio and lost. What the species may have been is known only to God". The frequency of such contradictions would make sense only if the descriptive conventions of the 18th century were different from ours. It is our 52 The pre-Lamarckian Oliva feeling that many of the ancient authors cited previous illustrations to report resemblance rather than conspecificity in its present meaning. In such conditions, réconciling the ancient species concepts with the rigid rules of the modern Code. of Zoological Nomenclature Code") necessarily entails some interpretation of the (hereunder referred to as "the message the old masters might have wanted to convey. In doing so, our main concern has been stability. (Changes to the presently accepted names have been made only nomenclatural when this was inevitable. This paper deals only with recent species. For each author, only new names have been considered, previously utilised names being nomenclatural without interest (they are necessarily junior homonyms), the only exception being a previous nomen nudum (which remains available). Names that are obviously incompatible with Oliva species have not been considered. For each name, the original description has been reproduced verbatim in order to allow a verification of our conclusions. The only modification brought to the original texts 1s that for each author, species have been presented in alphabetical order (in their original spelling), for the facility of the reader. Divergent attributions in recent works frequently consulted by Oliva students are given in notes. 1. The Oliva of Linnaeus, 1758. The species of OUliva described by Linnaeus in the tenth edition of the Systema Naturae were originally placed in FOLUTA, Cylindroidae f. subcylindricae. In citations of these species, the name of Linnaeus should thus be enclosed in parentheses (Code, art. 51 c). The Oliva of Linnaeus pose no more problem, having been studied by HANLEY (1855) and adequately revised by OLSSON & DANCE (1966), who carefully examined the type material at the Linnean Society, London. No specimen of Oliva has been reported in the Linnean collection at Upsala (HOLM, 1957). TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER ispidula (p.730, sp.351) ORIGINAL DESCRIPTION: Ispidula. 351. Vtesta cylindroide laevi, spira prominente margine unico. Rumph.mus.t.39.f.6,7. Pet.gaz.t.59.f.8. Habitat .. DISCUSSION. This name, cited here only because it has brought so much confusion in the Oliva literature, has already been discussed in great detail by OLSSON & DANCE (1966). The selected lectotype is not an Oliva but à fossil Agaronia (Agaronia plicaria Lam., 1811). STATUS: not an Oliva. Note: the name ispidula has not been rejected as a secondary homonym and remains available for an Oliva species not described under Voluta. The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 Habitat in M.Indico. Varietates coloribus infinite ludentibus; Litterata praefertur. DISCUSSION. This name has already been discussed by Olsson & Dance (1966), who selected and illustrated a lectotype, preserved at the Linnean Society, London. Linnaeus very fittingly called attention to the extreme variability of this species and the name should be used with caution. The lectotype seems to correspond to one of the species of the "Oiva oliva complex", the taxonomic structure of which has been discussed by TURSCH, MIssA & BOUILLON (1992). STATUS: valid name. Fig. 1. "Voluta" ispidula L., 1758. Type specimen, Linnean Society of London. Scale bar: 1 cm. oliva (p.729, sp.350) ORIGINAL DESCRIPTION: Oliva. 350. V.testa cylindroide laevi, spirae basi reflexa. List.Conch.4.f.10.c.l.t.2. Rumph.mus.t.39.f.2-5. Gualt.test.t.23.f.B. Argenv.conch.t.16.f.R. Litterata. Kratzenst. Regentf. 2.t.1.f.2. Fig. 2. Oliva oliva (L., 1758). Lectotype, Linnean Society of London. Scale bar: 1 cm. porphyria (p.729, sp.349) ORIGINAL DESCRIPTION: porphyria 349. V.testa cylindroide laevi, spirae basi oblitterata, labro medio retuso. List.Conch.t.727. Rumph.mus.t.39.f.1. Gualt.test.t.24.f.0.P. Argenv.conch.t.16.f.K. Kratzenst.Regenf.8.t.2.f.15. Habitat... 53 APEX 9(2/3): 51-78, juillet 1994 Affinitas tanta cum sequenti, ut potius varietas, quam dlistincta species, quamvis pretium eam nobilitaverit. DISCUSSION: This name has already been discussed by OLSSON & DANCE (1966), who selected and 1llustrated a lectotype. preserved at the Linnean Society, London. The lectotype corresponds to the unanimous. traditional concept of this species. STATUS: valid name. Fig. 3. Oliva porphyria (L., 1758). Lectotype, Linnean Society of London. Scale bar: 1 cm. 2. The Oliva of Born, 1778, 1780. No new olive name was given by Born. but one should consider his ispidula, which name being still available for an Oliva since ispidula (L. 1758) has been shown not to belong to this genus) ORIGINAL DESCRIPTION: In the /ndex (dating from 1778 according to RUTSCH, 1956), Born started his description: F.I1.4. Voluta ispidula. Die Spizdattel. Linn. S.N. Sp. 400 Testa subcylindrica laeui, spira conica, suturis acutis, columella incrassata oblique plicata. Die einigermassen walzenfôrmige glatte Schale hat Un ES The pre-Lamarckian Oliva einen dglatten Kkegelformigen scharfrandingen Schnirkel, und eine dichte schief gespaltene Spindel. Born then listed a number of colour varieties (from & up to y) over the next two pages, starting with: a albida, brunno maculata. Weisslich, mit braunen Flecken. In the Zestacea (1780). one finds the same varieties, with the same reference figures and a description: Testa emarginata subcylindrica, laevis; spira conica, longior, Anfractuum suturae acutae; Columella incrassata, oblique plicata. Dignoscitur a. V. Oliva, cui multum est affinis, basi spirali tumida, neque reflexa; Colorum differentia varietates, quas adduximus distinguit. Long 2 poll 9 lin. lat. 1. poll Patriam ubi constitit, varietatum descriptioni addidimus. DISCUSSION. This case presents obvious contradictions and it deserves careful consideration, as identical situations will occur for the same species treated by subsequent authors. On the one hand, there is an explicit reference to "Linn. S.N. Sp. 400" (ispidula in the 12th edition of the Systema Naturae). On the other hand, the description of colour varieties and the large size "2 poll 9 lin".(about 74.6 mm) are quite incompatible with the fossil French Agaronia of Linnaeus. The many illustrations supporting the descriptions of the numerous varieties are very conflicting. Bearing in mind that the concept of species at the time was quite different from ours, it is safer by far to conclude that the reference to "Linn. S.N. Sp. 400" indicates that Born referred to the species already described by Linnaeus. Note: this species 1s /lammulata Lamarck, 1811 according to BURCH & BURCH (1960). TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER 3. The Oliva of Schrôter 1782, 1783. No new olive name was given by Schrôter. but (as in the case of Born) one should consider his ispidula, this name being still available for an Oliva since ispidula (L. 1758) has been shown not to belong to this genus). The argument used in the case of Born also applies here and explicit references (both in the Musei Gottwaldiana and in the Einleitung) are taken to indicate that Schrôter referred to the species already described by Linnaeus. 4. The Oliva of Lightfoot-Solander, 1786. Lightfoot the author of the Portland Catalogue, as shown by DANCE (1962) and REHDER (1967) (see also IREDALE, 1916. DALL, 1921, Kay, 1965). Lightfoot utilised manuscript names given by Solander (his notes, Was real formerly in the Banks collection, are now in the library of the British Museum, according to Rehder) and added a few names of his own invention Only one species of Oliva was published in the Portland Catalogue. It was described as Voluta and in citations of this species, the name of Lightfoot should parentheses (Code, art. 51 c). One should note that Lightfoot did not describe species as such, but described lots of an thus be enclosed in auction. The same species can thus appear in two different lots. This is the case of: incrassata (p. 13, # 264) This shell appears in lots 264, 2315 and 3696. ORIGINAL DESCRIPTION: 264. Voluta incrassata, S. Martyn, 499, 500 very rare. 2315. "A fine specimen of Voluta incrassa, S. Martyn, Vol. || - very rare, f. 499. 500." 3696. "A very fine pair of Voluta Incrassata, S. extremely scarce ….Martyn, vol. II. 499. 500." DISCUSSION. Figures 499 and 500 of Martini (written "Martyn" by the author) clearly The pre-Lamarckian Oliva correspond to the unanimous present concept of this species. STATUS: valid name. Rehder draws attention to "incrassa", which he considers to be a misspelling of the trivial DILLWYN (1817) manuscript Oliva names of Solander. These are: aurora MSS. baltheata MSS. cruenta MSS. maculata MSS. ventricosa MSS. name. refers to other These names of Solander, being manuscript, have no nomenclatural standing. They are best referred to as (Solander) Dillwyn., 1817. Fig. 4. Oliva incrassata (Lightfoot, 1786). 4a: Martini, fig. 499. 4b: Martini, fig. 500. Scale 1:1. Un Un APEX 9(2/3): 51-78, juillet 1994 APEX 9(2/3): 51-78, juillet 1994 Notes: - aurora (Solander) Dillwyn, 1817 is carneola (Gmelin, 1791) for BURCH & BURCH (1960), a color form of the same for PETUCH & SARGENT (1986). - baltheata (Solander) Dullwyn, 1817 :1s annulata (Gmelin, 1791) for BURCH & BURCH (1960), WAGNER & ABBOTT (1978). - cruenta (Solander) Dillwyn, 1817 1s annulata (Gmelin, 1791) for BURCH & BURCH (1960). WAGNER & ABBOTT (1978).and ZEIGLER & PORRECA (1969) and emicator Meuschen (non binominal) for DAUTZENBERG (1927). - maculata (Solander) Dillwyn, 1817 is figrina Lamarck, 1811 for BURCH & BURCH (1960) and WAGNER & ABBOTT (1978). - ventricosa (Solander) Dillwyn, 1817 is bulbosa (Rôding, 1798) for BURCH & BURCH (1960), WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). 5. The Oliva of Abel, 1787. Abel (1787) introduced new names such as Voluta reticulata and Voluta porphyrea. But there are good reasons to consider Abel as being not consistently binominal. In the Voluta section alone, one species has two names (p.66, n°1: Voluta oliva - Vidua mauritana). Others like sepultura principis (p; 67, n°9) and vellus aureum (p. 68, n° 13) bear no generic name. Many species were referred to only by a vernacular name, for instance “Die grosse Panamarolle” (p. 69, n° 21). In our opinion, these facts constitute sufficient grounds for rejecting all the names of Abel. 6. The Oliva of Karsten, 1789. In the Museum Leskeanum - a work that appears to be consistently binominal (DUCHAMPS & TURSCH, 1994)- introduced two new names (o/ivacea and Karsten nigrita) on p. 216. Both names bear explicit reference to Martini but the work of that author has been officially rejected as non-binominal. Under the provisions of the Code, his names 56 The pre-Lamarckian Oliva remain available and have to be considered here. The Oliva of Karsten were described as Voluta and in citations of these species, the name of the author should thus be enclosed in parentheses (Code, art. 51 c). olivacea (p.216, # 638) ORIGINAL DESCRIPTION: Voluta olivacea Mart. 638 V. oliv. testa albida, punctis ex violaceo lutescentibus maculata, apertura atque columella omnino amethystinis. Martini Konch. Kab. T.2.tab. 46. fig. 498. 94. Long. 10 lin. lat 5 lin. DISCUSSION: The original description and the colour figures of Martini undoubtedly represent the very characteristic shell now known as tessellata Lamarck. The figures were indeed cited by Lamarck himself for fessellata. The measurements given by Karsten (a precursor of biometry) indicate the shell 1s twice as high as wide. We have verified that this is nearly exactly the case for tesselata. With much reluctance we are compelled to conclude that olivacea Karsten is the earliest name for the well-known fessellata Lamarck (the type material of which has disappeared), which becomes an objective Junior synonym as Lamarck referred to the same figures as Karsten. STATUS: valid name. S # : 4 ss 5a 5b Fig. 5. Oliva olivacea (Karsten, 1789). 5a: Martini, fig. 493. 5b: Martini, fig. 494. Scale Tate nigrita (p. 216, # 639, 640, 641) Under the name nigrita Karsten grouped his sections 639, 640 and 641. The note under 641 clearly indicates that the author considerred these as varieties or forms of the same species. TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 ORIGINAL DESCRIPTION: Voluta Nigrita Mart. 639 V.N. testa ferruginea, columella ex albescenti - sanguinea. 2. Martini Konch. Kab. T.2.tab. 47.fig.501. Long. 2 poll.1 lin. lat. 10 lin. 640 V.N. testa ex fusco nigra immaculata, apertura cum columella albida. Amboina, 2. Martini loc. cit. tab. 45, fig. 472.73. Long. 1 poll. 8 lin. lat. 10 lin. 641 V.N. testa ex fusco nigra, ad labri marginem externum luteo striata et characteribus obscurioribus scripta; apertura albida, columella rufescens. 2. Long. 2 poll. 2 lin. lat.9 lin. Note. Specimina 639-641 ad singularem speciem cum b. Martinio retuli propterea, quod columella in ipsis, postice duntaxat inprimis est striata, et magnitudine, hujus speciei omnes fere V. Olivae varietates Linn. superat. DISCUSSION. Figures 501, 472, 473 of Martini undoubtedly refer to the shell known today as vidua Rôding. With much reluctance we are compelled to conclude that nigrita Karsten 1s the earliest name for the well-known vidua Rôding (of which there is no type material), which becomes an objective junior synonym as the same figs. 472 and 473 are the only illustrations cited by Rôding, 1798 for vidua STATUS": valid name. Note: This species 1s oliva (L.. 1758) for DAUTZENBERG (1927). BURCH & BURCH (1960) and WAGNER & ABBOTT (1978) (who consider the work non binominal). One should also consider Karsten's ispidula, as this name 1s still available for an 6c Oliva, since ispidula (L. 1758) has been shown not to belong to this genus). The argument used in the case of Born also applies here and the Fig. 6. Oliva nigrita (Karsten, 1789). 6a: explicit reference is taken to indicate that Martini, fig. 472. 6b: Martini, fig. 473. 6c: Karsten referred to the species already described Martini, fig. 501. Scale 1:1. by Linnaeus. APEX 9(2/3): 51-78, juillet 1994 7. The Oliva of Gmelin, 1791. Two new names (annulata and carneola) were introduced by Gmelin in the 13th edition of the Systema Naturae. The Oliva of Gmelin were described as Foluta and in citations of these species, the name of the author should thus be enclosed in parentheses (Code, art. 51 C). annulata (p. 3440, # 18) ORIGINAL DESCRIPTION: annulata. 18. Vtesta laevi alba; dorsi annulo carinato. List.Conch.t.717. f.1. Martin. Conch. 2.t.51. f. 564. B) Martin. neuest. Mannigfalt., l.p.446.t.2.f.21 ? Habitat ... B) rufescente undulata. DISCUSSION. This case has been discussed in detail by TURSCH, GERMAIN & GREIFENEDER (1986) who concluded that it was a nomen dubium, used by subsequent authors to encompass both ©. amethystina Rôding and ©. mantichora Duclos (demonstrated to be two distinct species). The last reference of Martini will not be considered. as it was given with a question mark in the original description. The figure of Lister 1s very ambiguous. The "ring" alluded to by subsequent authors does not clearly show on the profile of the shell and the difference in shadowing of the body whorl could also be interpreted as indicating a difference in coloration. The figure could very well depict some other species. Fig. 564 of Martini 1s even more ambiguous as the details of the body whorl are suspiciousiy similar to those of Lister's figure. Here the shell is clearly ringed. WEINKAUFF (1878) correctly remarked "Die Martini'sche Figur kann aber ebensogut auf die gekielte und farblose Varietät der ©.peruviana gedeutet werden". Although the shells of ©. mantichora are frequently ringed and very occasionally occur in a white form, it would take quite a stretch of imagination to identify either ©. amethystina Rôding or ©. mantichora Duclos with any of the above illustrations. STATUS: nomen dubium. The pre-Lamarckian Oliva Note: This is. - valid for ZEIGLER & PORRECA (1969), PETUCH & SARGENT (1986). - a form of emicator (Meuschen) (rejected work) for DAUTZENBERG (1927). carneola (p 3443, # 24) ORIGINAL DESCRIPTION: Carneolus. 24. Vtesta aurantia fasciis caeruleis, spira complanata et apertura albis. Martin Conch.2.t.46.f.495. Habitat ... DISCUSSION. The small coloured illustration of Martini depicts a shell with white spire. white fasciole. orange body whorl decorated with several brown horizontal stripes and a blue square blot. Although the spire and the general shape are not correctly depicted, this figure 1s quite compatible with the present concept of ©. carneola (a justified subsequent correction of the original spelling carneolus by LAMARCK. 1811. p. 321) a name which should be preserved in the Interests of stability. Subsequent descriptions of very similar species (such as ©. kwajalainensis da Motta, 1985).and of numerous varieties (by Dautzenberg, 1927) might eventually require the designation of a neotype. STATUS: valid name. Fig. 7. Oliva carneola (Gmelin, 1791). Martini, fig. 495. Scale 1:1. crassa (p. 3421 # 108) ORIGINAL DESCRIPTION: C. testa crassa subflava : fasciis tribus albidis, ore caerulescente. List. Conch. t. 664. f. 8. Habitat . .. pollices longa. carneolae affinis, testa ultra 4 TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER DISCUSSION. This species was described in the section Cypraea. It is considered here only because Gmelin deemed it close to carneola. The figure of Lister unmistakably depicts a Cypraea. STATUS: not an Oliva. Note. This is: - Pseudoliva crassa for BURCH & BURCH (1960). - an indeterminate species for WAGNER & ABBOTT (1978). One should also consider Gmelin's ispidula, this name being still available for an Oliva since ispidula (L, 1758) has been shown not to belong to this genus. The argument used in the case of Born also applies here and an explicit reference "Mus.Lud. Ulr." (the Ludovicae Ulricae of Linnaeus, 1764) indicates that Karsten refers to the species already described by Linnaeus. "O. leucophaea Gmelin" is given by Môrch, Museum Reginae 1850 and 1s sometimes cited in the subsequent literature. We have not found this species in Gmelin. 8. The Oliva of Rüding, 1798. The Museum Boltenianum, written by Peter Friedrich Rôding, is an inventory of the rich collection of J.F. Bolten, a leading physician in Hamburg. The rediscovery of this work at the beginning of this century and its subsequent recognition as an available work (in the sense of the Code) caused a major upset in the nomenclature of molluscs Bolten 1s reported to have been a lifelong student of conchology, dissatisfied with the "crude method of Linnaeus". In contrast, we believe that his friend Rôding simply intended to produce an inventory with no scientific pretensions. It can indeed be seen in the "descriptions" reported here below that he gives very accurate indications on the number (e.g. "15 St.") and the disposition of the specimens in the display cabinets (Oliva were in "Lade" 18 to 21) while presenting only very minimal information on the characteristics of the shells The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 themselves. It is not surprising that the interpretation of such an inventory as a scientific publication would lead to poor results. Only 6 (less than 20%) of the 46 Rôding's Oliva names can be identified, possibly a sad world record. For information on the Museum Boltenianum see DALL (1915) and REHDER (1945). The Oliva of Rôding are all grouped in the genus Porphyria. AI descriptions start with two numbers, the first -in a separate column indicates the number for the species and the second -in the text- indicates the number in the section Porphyria). In citations of Rôding's species, the name of the author should thus be enclosed in parentheses (Code, art. 51 c). amethystina (p.35, # 440) ORIGINAL DESCRIPTION: 440134 P. Amethystina. Die amethystfarbene Dattel. Gmel.sp.23.V.ispidula. Martini 2 .t.46. f.491.492. Knorr Vergn. 2.t.10.f 6 7. 14St. DISCUSSION. This name has already been treated in detail by TURSCH, GERMAIN and GREIFENEDER, 1986, who amethystina Rôding, 1791 from ©. mantichora separated ©. Duclos, 1835 (so far mixed under the nomen dubium ©. annulata Gmelin, 1791, q.v.). The adjective “amethystfarbene" correctly applies to shells of the species amethystina many specimens of which have a deep violet colour pattern. This 1s very rarely (1f ever) the case for shells of mantichora. Figures 6 and 7 of Knorr can be interpreted as depicting a specimen of amethystina With no markings on the suprafasciolar band. In the figures of Martini the body whorl 1s a rather dark vellow- beige and the interior of the aperture is deep orange. Figure 492 shows a suprafasciolar pattern compatible with the pattern characteristic of amefhystina. STATUS: valid name. Note. This is: - annulata (Gmelin, 1798) for BURCH & BURCH (1960). 59 APEX 9(2/3): 51-78, juillet 1994 The pre-Lamarckian Oliva TURSCH, DUCHAMPS & GREIFENEDER a form of annulata (Gmelin, 1798) for spicata 1s à name in very common use 1t should WAGNER & ABBOTT (1978), ZEIGLER & be preserved in the interests of stability. PORRECA (1978), PETUCH & SARGENT (1986). STATUS: objective synonym of spicata - emicator Meuschen (non binominal) for (Rôding, 1798). DAUTZENBERG (1927). ces aurata (p. 33, # 402) ORIGINAL DESCRIPTION: 402116 P. aurata. Die goldgelbe Dattel. Gmel. V.Oliva .sp. 17. 1St. DISCUSSION. Gmelin's sp. 17 is oliva, preoccupied by o/iva (L., 1758). This name 1s completely unidentifiable. STATUS: nomen nudum. Note. This is: - a variety of oliva (L., 1758) for DAUTZENBERG (1927). - oliva (L., 1758) according to BURCH & BURCH (1960). - a colour form of vidua (Rôding. 1798) for ZEIGLER & PORRECA (1969), WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). bulbosa (p. 37, # 459) Fig. 8. Oliva amethystina (Rüding, 1798). 8a: Martini, fig. 491. 8b: Martini, fig. 492. Scale 1:1. amoena (p.33, # 399) ORIGINAL DESCRIPTION: 399 | 13 P. Amoena. Die hübsche Dattel. Gmelin. V.Oliva. sp. 17. 3St. DISCUSSION. Gmelin's sp. 17 is oliva, preoccupied by oliva (L., 1758). This name 1s completely unidentifiable. STATUS: nomen nudum. arachnoïdea (p.36, # 450) ORIGINAL DESCRIPTION: 450|43 . P. arachnoidea. Die Spinneweben- Dattel. Gmel. V.Oliva a. sp. 17. Martini 2.t.48.f.509.10. 1St. Fig. 9. Oliva bulbosa (Rüding, 1798). 9a: DISCUSSION. Martinis figures were PO À MERE Tid: 50706: Metif 9 08 1e utilised by Rôding himself for spicata. As 1:1. 60 TURSCH, DUCHAMPS & GREIFENEDER ORIGINAL DESCRIPTION: 459|50 P. Bulbosa. Die knolligte Dattel. Gmel.sp. 17. Oliva &. Martini 2.t.47 .f.507.508. 2St. DISCUSSION. Martinr's figures 507 and 508 were previously utilised by Gmelin for oliva, a name preoccupied by oliva (L., 1758). The two figures are quite compatible with the present concept of ©. bulbosa, a name that should be preserved in the interests of stability. The drawing of the columella rather vaguely suggests the characteristic prominent columellar plications of the species and the adjective "knolligte" could be taken to refer to this peculiarity. STATUS: valid name. caerulea (p. 33, # 392) ORIGINAL DESCRIPTION: 39217 P. Caerulea. Die himmelblaue Dattel. Gmel.V.Oliva sp. 17.a Martini 24t.48.f. 518. Rumph. t.39. f.5. 13 St. DISCUSSION. This case has already been studied by KILBURN (1980). The figure of Martini was previously used for oliva var. of Gmelin. Gmelin's sp. 17 is oliva, preoccupied by oliva (L., 1758). The figure (designated as a type figure by KILBURN) shows a characteristic blue colour aperture and is quite compatible with the present concept of O. caerulea. STATUS: valid name. Fig. 10. Oliva caerulea (Rüding, 1798). Martini, fig. 518. Scale 1:1. The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 Note. This 1s: - episcopalis Lamarck, 1811 for DAUTZENBERG (1927), BURCH & BURCH (1960) and ZEIGLER & PORRECA (1969). cornea (p. 36, # 448) ORIGINAL DESCRIPTION: 448 | 41. P. cornea. Die hornfarbene Dattel. 1 St. DISCUSSION. This unidentifiable. species is completely STATUS: nomen nudum. cingulata (p. 34, # 415) ORIGINAL DESCRIPTION: 415121 P. Cingulata. Die gegürtelte Dattel. Gmel.V.Oliva sp. 17. 1St. DISCUSSION. preoccupied by oliva L., 1758. This name 1s Gmelin's sp. 17 1s oliva, completely unidentifiable. STATUS: nomen nudum Note. This is annulata (Gmelin, 1791) for WAGNER & ABBOTT (1978). coffea (p. 37, # 462) ORIGINAL DESCRIPTION: 462153 P. Coffea Die Kaffeebohne. Gmel.sp.24. V.carneolus. Martini 2.t.46. f.495,. 4St. DISCUSSION. Martini's previously utilised by Gmelin for carneola, cited by Rôding. STATUS: carneola Gmelin, 1791. figure 495 was objective junior synonym of Note. This 1s indeterminate, possibly o/iva (L., 1758) for WAGNER & ABBOTT (1978). conoiïdea (p. 35, # 430) ORIGINAL DESCRIPTION: 430|31 P. Conoïidea. Die kegelformige Dattel. 1St. DISCUSSION. This unidentifiable. name is completely STATUS: nomen nudum. 61 APEX 9(2/3): 51-78, juillet 1994 dealbata (p. 35, # 427) ORIGINAL DESCRIPTION: 427 | 29 P. Dealbata. Die schneeweisse Dattel. Knorr 6. t. 34. f. 4 5. 6 St. (07 LEE DISCUSSION. background) are compatible with an all-white Knorr’s figures (on a black specimen of the “O. oliva complex” shown to be composed of distinct, closely related species (TURSCH, MIssSA & BOUILLON, 1992) well separated by multivariate analysis but impossible to segregate on the basis of approximate illustrations. STATUS: nomen dubium. Note. This is a white form of oliva (L.. 1758) for WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). The name /asciata was used twice, for two different shells (sp. 387 and sp. 411). fasciata (p. 32, # 387) ORIGINAL DESCRIPTION: 387|2 P. Fasciata. Die gebandete Portobello- Dattel. Gmel. Voluta sp. 16 y. 1St. DISCUSSION. Gmelin's sp. 16 is porphyria, preoccupied by porphyria (L. 1758). One should note that Rôding also uses "Portobello- Dattel" 386). The adjective "“"gebandete" indicates that Rôding means a banded colour variation of this shell. for porphyria (species There is no indication whatsoever that such a variant would deserve specific or subspecific status. STATUS: porphyria (L., 1758). subjective junior synonym of fasciata (p. 34, # 411) ORIGINAL DESCRIPTION: 411|19 P. Fasciata. Die Band Dattel. Gmel.V.sp. Knorr 3.t.17 .f.3. 6St. DISCUSSION. Knorr's figure was previously utilised by Gmelin for oliva, preoccupied by oliva (L.,1758). The very fact that the name Jasciata was used twice (apparently for very different shells) casts a serious doubt upon 62 The pre-Lamarckian Oliva TURSCH, DUCHAMPS & GREIFENEDER Rôding's nomenclatural concepts. The figure of Knorr is easily recognisable as the dark form of nigrita Karsten, 1789, also described as vidua by Rôding. STATUS: junior homonym of fasciata (sp. 387 Rôding., 1798) to which we give page precedence. Note. This is: - oliva (L., 1758) for BURCH & BURCH (1960). - reticulata (Rôding, 1798) for WAGNER & ABBOTT (1978). fenestrata (p. 34, # 417) ORIGINAL DESCRIPTION: 417|22.P. Fenestrata. Die gegitterte Dattel. Gmel.V.Oliva sp. 17 8. Martini 2.t.47 .f.502. 2St. DISCUSSION. Martini's figure was previously used by Gmelin for oliva, preoccupied by oliva (> 21758) This (vaguely reminiscent of a golden form of vidua by the colour figure same author) presents a very strange cross-ruled pattern (never seen by us in an Oliva). It 1s not recognisable with any certainty. STATUS: nomen dubium. Note. This is a form of vidua (Rôding, 1798) according to DAUTZENBERG (1927), ZEIGLER & PORRECA (1969), WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). fimbriata (p.34, # 410) ORIGINAL DESCRIPTION: 410|18 P. Fimbriata. Eine schône rostfarbigte Dattel mit dem Saum. 1St. DISCUSSION. One should note that fimbriata was presented together with oliva under 410. This name is completely unidentifiable. STATUS: nomen nudum. fulgurator (p.36, # 453) ORIGINAL DESCRIPTION: 453|45 P. Fulgurator. Die Blitz-Dattel. Gmel.sp. 17.V.oliva @. Martini 2.t.51. f.562. 14St. DISCUSSION. Martinri's figure 562 previously utilised by Gmelin for oliva var., a Was name previously preoccupied by oliva (L. 1758). The figure is very recognisable and depicts a specimen of fulgurator in the presently accepted sense of the name. STATUS: valid name. Fig. 11. Oliva fulgurator (Rôding, 1798). Martini, fig. 562. Scale 1:1. griseola (p.35, # 424) ORIGINAL DESCRIPTION: 424126 P. Griseola. Die gräuliche Dattel. 3St. DISCUSSION. This name 1s completely unidentifiable. STATUS: nomen nudum. hepatica (p.33, # 400) ORIGINAL DESCRIPTION: 400114 P. Hepatica. Die lichtgraue Dattel. Gmel. V.Oliva. sp. 17. 2St. DISCUSSION. Gmelin's sp. 17 is oliva, preoccupied by oliva Linnaeus, 1758. This name is completely unidentifiable. STATUS: nomen nudum. isabella (p.33, # 401) ORIGINAL DESCRIPTION: 401|15 P. /sabella. Die isabellfarbene Dattel. Gmel. V.Oliva. sp. 17. 3St. DISCUSSION. Gmelin's sp. 17 is oliva, preoccupied by oliva (L., 1758). This name is completely unidentifiable. STATUS: nomen nudum. TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 ispida (p.35, # 431) ORIGINAL DESCRIPTION: 431132 P. /spida Die blaumündige Dattel. Gmel Voluta sp.23. Martini 2.t.49. f.524.25.30. Knorr 3.t.19.f.3. 21 St. 432| à Mart. 2. t.49. 1.535. 3St. 433| 8 Martini 2.t.49. f.522.23. 5St. 434| 4 St. 435| à 3 St. DISCUSSION. Martini's figures 522, 523, 524 as well as Knorr's fig. 19/3 were previously utilised by Gmelin for ispidula, a name preoccupied by ispidula Linnaeus, 1758. Martini's figures 525 , 530 and 535 were not previously utilised. All the figures are compatible with specimens of the "Oliva oliva complex" shown to be composed of distinct, closely related species (TURSCH, MIssA & BOUILLON, 1992) well separated by multivariate analysis but impossible to segregate on the basis of approximate illustrations. STATUS: nomen dubium. Note. This is: - a nomen nudum according to BURCH & BURCH (1960). - oliva (L., 1758) for ZEIGLER & PORRECA (1969), WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). labradoriensis (p.32, # 389) ORIGINAL DESCRIPTION: 389|4 P. Labradorensis. De Schiller-Dattel. Gmel. Voluta Oliva, sp. 17. Lister 731. f.20. 2 St. DISCUSSION. The figure of Lister was previously utilised by Gmelin in conjunction with oliva, a name preoccupied by oliva (L., 1758). It was later utilised by Lamarck for mustelina. The figure is not recognisable and possibly depicts an Agaronia. STATUS: nomen dubium. Note. This (misspelled "/abradorensis") is: a nomen nudum according to BURCH & BURCH (1960). 63 APEX 9(2/3): 51-78, juillet 1994 The pre-Lamarckian Oliva TURSCH, DUCHAMPS & GREIFENEDER - possibly funebralis Lamarck, 1811 for PETUCH & SARGENT (1986). litterata (p.36, # 452) ORIGINAL DESCRIPTION: 452144 P. Litterata. Die Buchstaben - Dattel. Gmel.sp. 17. V.oliva.yy. Martini 2.t.46. f.488. 14 St. DISCUSSION. Martins figure 488 was previously utilised by Gmelin for oliva var., a name preoccupied by oliva (L., 1758). It is not recognisable and could amongst others represent either spicata, reticularis or fulgurator. STATUS: nomen dubium. Note. This 1s spicata (Rôding, 1798) according to BURCH & BURCH (1960), WAGNER & ABBOTT (1978), ZEIGLER & PORRECA (1969). KEEN (1971) and PETUCH & SARGENT (1986). mica (p.35, # 436) ORIGINAL DESCRIPTION: 436133 P. Mica. Die blaugefleckte Dattel. Gmel. sp.23. V.ispidula. 23 St. 437| a Martini 2.t.49 f.527-529. 3 St. 438| B10St. 439| y 3St. DISCUSSION. Martini's figures were never utilised before. These figures represent an unidentifiable Olive, with a very strange lip. possibly a freak. STATUS: .nomen dubium. lote. This is oliva (L., 1758) according to WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). miniacea (p.33, # 391) ORIGINAL DESCRIPTION: 391|6 P. Miniacea. Das Morgenroth. Gmelin V. porphyria. sp.16 B. Martini 2.t.45. f.476,477. 9 St. DISCUSSION. Both figures of Martini were previously used for porphyria Gmelin, a name preoccupied by porphyria (L., 1758). The figures are highly recognisable and depict the 64 present, widely utilised concept of miniacea. The same figures were utilised later by Lamarck for erythrostoma. STATUS: valid name. Note. This is: erythrostoma Meuschen (rejected work) for DAUTZENBERG (1927). Fig. 12. Oliva miniacea (Rôding, 1798). 12a: Martini, fig. 476. 12b: Martini, fig. 477. Scale 1:1. TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 oculata (p.35, # 426) ORIGINAL DESCRIPTION: 426128 P. Oculata Die geängelte Dattel. 12 St. DISCUSSION. "Geängelte" is probalbly a misprint for "geäugelte". This name is completely unidentifiable. STATUS: nomen nudum. ornata (p.33, # 398) ORIGINAL DESCRIPTION: 398112 P. Ornata Die geschmüekte Dattel. Gmel. V.Oliva. sp. 17. 2 St. DISCUSSION. Gmelin's oliva is preoccupied by oliva (L., 1758). This species is completely indeterminate. STATUS: nomen nudum. Note. This is oliva (L. 1758) according to WAGNER & ABBOTT (1978). paleacea (p.36, # 442) ORIGINAL DESCRIPTION: 442 | 36 P. Paleacea. Die strohfarbene Dattel. 1 St. DISCUSSION. This name is completely unidentifiable. STATUS: nomen nudum. papyracea (p. 36, # 443) ORIGINAL DESCRIPTION: 443|37 P. Papyracea. Die buntpapierne Dattel. 4 St. DISCUSSION. This name is completely unidentifiable. STATUS: nomen nudum. punctata (p.33, # 397) ORIGINAL DESCRIPTION: 397|11 P. Punctata. Die gestippelte Dattel. Gmel. V.Oliva. sp. 17. 4 St. DISCUSSION. Gmelin's oliva is preoccupied by oliva (L., 1758). This name 1s completely unidentifiable. STATUS: nomen nudum. Note. This is: -spicata (Rôding, 1798) var. venulata Lamarck, 1811 according to BURCH & BURCH (1960). -oliva (L.. 1758) for WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986). - rejecta Burch & Burch, 1962 according to PETUCH & SARGENT (1986) quercina (p.34, # 418) ORIGINAL DESCRIPTION: 418|23 P. Quercina. Die Eichenholz - Dattel. Gmel. V.Oliva sp. 17. Knorr 5.tab.26. fig.4. 6 St. 419] P. 2 St. 420| B Knorr 5.t.27.fig.5. 1 St. DISCUSSION. Knorr's fig. 26/4 was previously utilised by Gmelin for o/iva var. and by Rôding himself for sepultura principis. Knorr's fig. 27/5 was previously utilised by Gmelin for oliva var. preoccupied by oliva (L., 1758). Many olives with a low spire have melanistic forms and these figures are ambiguous. STATUS: nomen dubium. Note. This could be reticulata (Rôding, 1798) for WAGNER & ABBOTT (1978). reticulata (p.33, # 396) ORIGINAL DESCRIPTION: 396 |10 P. Reticulata Die Netz-Dattel. Gmel.V.Oliva.sp. 17.œ. Martini 2.t.48. f.512.533. 9 St. DISCUSSION. Martini's figure 512 was previously utilised for o/iva as well as ispidula Gmelin (both names preoccupied by Linnaeus). It was later utilised for sanguinolenta Lamarck. Fig. 533 was not previously utilised. It was used later by Lamarck for fabagina. Fig. 533 is entirely unidentifiable (and "533" is probably a misprint for 513), but Fig. 512 (a dark brow- grey shell with an orange columella and lip) is not inconsistent with the current acceptation of the widely used name reticulata, which should be kept for the sake of stability. STATUS:.valid name. Note. This is sanguinolenta Lamarck, 1811 for BURCH & BURCH (1960). 65 APEX 9(2/3): 51-78, juillet 1994 Fig. 13. Oliva reticulata (Rôding, 1798). Martini, fig. 512. Scale 1:1. ruffina (p.32, # 388) ORIGINAL DESCRIPTION: 388|3 P. Auffina. Die rothe Dattel. Gmel.Vol. sp.49.y. 2 St. DISCUSSION. There is a previous l. ruffina of Gmelin (p. 3450, sp. 49) which is not cited by Rôding. It is based upon "Gualtieri t.54 f. G ?". This figure could depict vidua, but Gmelin's decription of ruffina "… V. testa integriuscula transversim rugosa: columella quadriplicata, labro crenulato" is not compatible with an Oliva. Rôding's ruffina 1s unidentifiable. STATUS: nomen dubium. completely Note. This is a nomen nudum for WAGNER & ABBOTT (1978). sepultura principis (p.33, # 403) ORIGINAL DESCRIPTION: 403|17 P. Sepultura Principis. Das Prinzen Begräbnisz. Gmel.V.Oliva.sp. 17.4 Rumpf t.39.f.4. a 2 St. 405| B Martini 2.t.51.f.563. Knorr 5.t.26.f.4. 4 St. 406| y Martini 2.t.45.f.480.481. Knorr 5..19.f.1. 3 St. 407| 54St. 408| e 2St. 409| E 1 St. 66 The pre-Lamarckian Oliva DISCUSSION. The figure of Rumphius was previously utilised by Gmelin for o/iva. Knorr's fig. 26/4 was previously utilised by Gmelin for oliva var. and was also utilised by Rôding himself for quercina. Knorr's fig. 19/1 was previously utilised by Gmelin for oliva var. and was also utilised by Rôding himself for variegata à. Martinr's figs. 45/480, 481 were previously utilised by Gmelin for oliva var. and are utilised later by Lamarck for funebralis. Martini's fig. 51/563 was previously utilised by Gmelin for oliva var. Gmelin's oliva 1s preoccupied by Knorr 5.t.19.f 1. is a Conus (possibly aulicus). Martini Linnaeus. 1758. 2.1.51.f.563 1s ambiguous, could be peruviana as well as vidua. Martini 2.t.45.f480.481 could be what we call Oliva funebralis. Rumpf. t.39.f4. 51264. is reminiscent of the shell now usually called is not recognisable. Knorr lignaria, but could be any dark olive with a low. calloused spire. All these figures are either unidentifiable or conflicting. STATUS: nomen dubium. Note. This could be funebralis Lamarck,1811 for WAGNER & ABBOTT (1978). sericea (p.33, # 390) ORIGINAL DESCRIPTION: 390|5 P. Sericea. Das Seidenzeug. Gmel. V.oliva sp. 17.8. Martini 2.t.51. f.559.561. 8 St. DISCUSSION. Martini fig.559 utilised for oliva var. Gmelin, later by Lamarck previously for textilina. Fig. 561 later utilised by Lamarck for both irisans and reticularis. Gmelin's sp. 17 is oliva, a name preoccupied by Linnaeus. The figures are not inconsistent with the current acceptation of the widely used name sericea, which should be kept for the sake of nomenclatural stability. STATUS: valid name. Note. This is: - textilina Lamarck, 1811 for DAUTZENBERG (1927) - textilina Lamarck, 1811 (pars) for BURCH & BURCH (1960) and ZEIGLER & PORRECA (1969). TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER 14a D 14b Fig. 14. Oliva sericea (Rôding, 1798). 14a: Martini, fig. 559. 14b: Martini, fig. 561. Scale ja à spicata (p.35, # 423) ORIGINAL DESCRIPTION: 423125 P. Spicata Die Kornähre. Gmel.V.oliva sp. 17. Martini 2.t.48. f.509.10. 7 St. DISCUSSION. Martini's figures were previously utilised by Gmelin for oliva, preoccupied by oliva (L., 1758). The same figures are utilised by Rôding himself for arachnoides. They were later utilised by Lamarck for araneosa. The figures are not inconsistent with the current acceptation and the widely used name spicata should be kept for the sake of stability. STATUS: valid name. Note. "O. intertincta Rôding, 1798" given in the synonymy of spicata by WAGNER & ABBOTT (1978) is not a name published by Rôding. Probably a typographic error for intercincta Carpenter, 1857. The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 15b Fig. 15. Oliva spicata (Rôding, 1798). 15a: Martini, fig. 509. 15b: Martini, fig. 510. Scale sen textilis (p.37, # 456) ORIGINAL DESCRIPTION: 456|47 P. Textilis. Die gewebte Dattel. 1 St. DISCUSSION. This name :is unidentifiable. completely STATUS: nomen nudum. tigris (p.36, # 441) ORIGINAL DESCRIPTION: 441135 P. Tigris. Die Tiger-Dattel. 1 St. DISCUSSION. This name :1s unidentifiable. completely STATUS: nomen nudum. 67 APEX 9(2/3): 51-78, juillet 1994 tuberosa (p.37, # 460) ORIGINAL DESCRIPTION: 460151 P. Tuberosa Die kanehlfarbene Dattel. Kammerer t.3.fig. 7.8. 3 St. DISCUSSION. Kammerer's figures are quite clear and unmistakably depict a form of ©. bulbosa (Rôding. 1798), a conclusion also reached by WAGNER & ABBOTT (1978), ZEIGLER & PORRECA (1969) and PETUCH & SARGENT (1986). The name bu/bosa has page precedence, is widely used and should be kept for the sake of stability. STATUS: subjective bulbosa (Rôding, 1798). junior synonym of tumida (p.37, # 455) ORIGINAL DESCRIPTION: 455|46 P. Tumida. Die aufgeblasene Dattel. Lister t.746.f.40. 1 St. DISCUSSION. Lister's figure 40 was previously utilised by Gmelin in his remarks on o/iva, à name preoccupied by oliva (L. 1758). The figure of Lister is unrecognizable and might even not depict an Oliva. This name :1s completely unidentifiable. STATUS: nomen nudum. Note. This is an indeterminate Ancilla for WAGNER & ABBOTT (1978). turgida (p.34, # 416) ORIGINAL DESCRIPTION: 416|21 * B. Turgida Die wulstige Dattel. Gmel. V.Oliva sp. 17. 1 St. DISCUSSION. We have no explanation for the “* B.” in the original description. It might be a typographical error. Gmelin's sp. 17 1s oliva, à name preoccupied by oliva (L. 1758). This name 1s completely unidentifiable. STATUS: nomen nudum. umbrosa (p.36, # 449) ORIGINAL DESCRIPTION: 449|42 P. Umbrosa. Die Licht- und Schatten- Dattel. Gmel.Voluta ispidula @. sp.23. Martini 2.t.49.f.537. Knorr 1.t.15. 1.7. 3 St. 68 The pre-Lamarckian Oliva DISCUSSION. Martinr's figure 537 was never utilised before. Knorr's figure 7 was utilised by Gmelin for oliva, a name preoccupied by oliva (L.. 1758). The figures of Martini and Knorr are compatible with a dark specimen of the "Oliva oliva complex" shown to be composed of distinct, closely related species (TURSCH, MISsA & BOUILLON, 1992), well separated by multivariate analysis but impossible to segregate on the basis of approximate illustrations. STATUS: nomen dubium. Note. This is oliva (L.. 1758) for WAGNER & ABBOTT (1978) and PETUCH & SARGENT (1986) undulata (p.35, # 425) ORIGINAL DESCRIPTION: 425127 P. Undulata Die wellenformige Dattel. 18 St. DISCUSSION. This name :1s unidentifiable. completely STATUS: nomen nudum. variabilis (p.33, # 395) ORIGINAL DESCRIPTION: 395|9 P. Variabilis. Die spielende Dattel. Gmel. V. Oliva.p.17. 7 St. DISCUSSION. Gmelin's sp. 17 1s oliva, a name preoccupied by ofliva (L., 1758). This name is completely unidentifiable. STATUS: nomen nudum. Note. This 1s: - sanguinolenta Lamarck, 1811 for BURCH & BURCH (1960). - reticulata (Rôding, 1798) for WAGNER & ABBOTT (1978), ZEIGLER & PORRECA (1969) and PETUCH & SARGENT (1986). variegata (p.33, # 393) ORIGINAL DESCRIPTION: 393|8 P. Variegata. Die schäckigte Dattel. Gmel.Voluta.Sp. 17.8. Martini 2t.45.f.478.479. 24 St. 394| a Martini 5.t.19.f.1. 5 St. 2.t.45.f.480.481. Knorr TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER DISCUSSION. Martini's fig. 478 and 479 were previously utilised by Gmelin for Martini's figs. 480 and 481 were utilised by Gmelin for a variety of o/iva, by Rôding himself for sepultura principis (a nomen dubium) and later by Lamarck for funebralis. Knorr's figure oliva. (possibly Conus aulicus) had already been utilised for oliva and oliva var. by Gmelin; and also by Rôding himself for sepultura principis. Gmelin's sp. 17 is oliva a name preoccupied by oliva (L., 1758). STATUS: nomen dubium. Note. This is: - sanguinolenta Lam., 1811 for DAUTZENBERG (1927). - reticulata (Rôding, 1798) for WAGNER & ABBOTT (1978). ZEIGLER & PORRECA (1969), PETUCH & SARGENT (1986). vellus-aureum (p.36, # 444) ORIGINAL DESCRIPTION: 444138 P. Vellus-aureum. Das goldne Vliesz. Gmel.V.Oliva sp. 17.var.ce. Martini 2.t.46.f.490. 3 St. DISCUSSION. Martini's fig. 490 was previously utilised by Gmelin for oliva var, a name preoccupied by oliva (L., 1758). The figure could represent reticularis as well as spicata or even another species. STATUS": nomen dubium. Note. This is probably oliva (L. 1758) for WAGNER & ABBOTT (1978) vidua (p.34, # 412) ORIGINAL DESCRIPTION: 412120 P. Vidua Die ungarische Wittwe. Gmel.V.sp. 17. Martini 2.t.45. f.472.473. 17 St. 413| à 1, St 414| BP 1 St. DISCUSSION. Martini's figures have been previously utilised by Gmelin for oliva, a name preoccupied by Linnaeus, 1758. They were later utilised by Lamarck for maura. As both cited figures were utilised to establish nigrita (Karsten, 1789) we are reluctantly compelled to give priority to Karsten's name. The pre-Lamarckian Oliva STATUS: objective junior synonym of nigrita Karsten, 1789. Note. This name is valid according to BURCH & BURCH (1960), ZEIGLER & PORRECA (1969) PETUCH & SARGENT (1986). ziczac (p.37, # 461) ORIGINAL DESCRIPTION: 461152 P. Ziczac. Die Ziczac Dattel. 4 St. DISCUSSION. This unidentifiable. STATUS: nomen nudum. name is completely 9, The Oliva of Link, 1807. In his Beschreibung, Link introduces the new names coerulea (not caerulea), fusca, miniata, taeniata and tentorium. The name aurata also has to be considered, since aurata (Rôding, 1798) is a nomen nudum and therefore remains available. The Oliva of Link are all grouped in the genus Porphyria, and in citations of these species, the name of the author should thus been enclosed in parentheses (Code, art. 51 c). aurata (p. 97) ORIGINAL DESCRIPTION: P. aurata Gelbmündige D. Mart.Conch. 2.t.46.f.491,492. Unterscheidet sich von allen übringen durch die gelbe Mündung; nähert sich sonst in allen Stücken sehr der vorigen (note: this refers to coerulea). DISCUSSION. The two Martini figures have already been used by Rôding for amethystina. STATUS: objective junior amethystina (Rôding, 1798). Note. This 1s a color form of bulbosa (Rüding, 1798) for PETUCH & SARGENT (1986) synonym of coerulea (p. 97) ORIGINAL DESCRIPTION: P. coerulea. Kamelot D. Bolt. Verz. p. 33. +. Oliva «. Linn. Gm. |. c. Mart. Conch. 2. t. 48. f. 515. 516. Unterscheidet sich von allen übrigen durch die blaue Mündung, auch ist viel Blau in 69 APEX 9(2/3): 51-78, juillet 1994 APEX 9(2/3): 51-78, juillet 1994 der Zeichnung. Die Grôsse ist nicht viel über ein Zoll; die kleinern weichen etwas in der Form ab. Die zweite Windung ist wenig oder gar nicht aufwärts getrümmt. Immer ist sie schmal. DISCUSSION. Although the cited figures of Martini reference to the Bolten Catalogue and the are not convincing, the explicit description point to caerulea (Rôding, 1798). STATUS: incorrect (Rôding, 1798). spelling for caerulea fusca. (p. 95) ORIGINAL DESCRIPTION: P. fusca. Zigeuner D. V.Oliva € Linn.Gmi.l.c. Mart. Conch.2.t.47. f.501. Gelblich braun; die zweite Mündung stark aufwärts gebogen. DISCUSSION. O. fusca (Link, 1807) is a homonym of fusca Fischer, 1807. We know the exact date of publication neither of the \useum Demidoff (Fischer, 1807) nor of the Beschreibung (Link. 1807) and we are thus unable to decide on priority. The Martini figure had already been used for a variety of ©. oliva by Gmelin. This figure depicts the orange variety of nigrita (Karsten, 1789) (see PL. 9, fig. 3 of Zeigler & Porreca, as vidua Rüding, 1798). STATUS: subjective junior synonym.of nigrita (Karsten, 1789). Note. This is: - oliva (L., 1758) for BURCH & BURCH (1960). - vidua (Rôding. 1798) for WAGNER & ABBOTT (1978), PETUCH & SARGENT (1986). miniata (p. 95) ORIGINAL DESCRIPTION: P. miniata. Morgenroth D. Bolt. Verz. p.33. Porphyria B Linn..c. Mar. Conch. 2. t.45. f.476.477. Kenntlich an der rothen Mündung. DISCUSSION. Both Martini figures had already been used by Rôding for miniacea. STATUS: objective junior synonym of miniacea (Rôding, 1798). 70 The pre-Lamarckian Oliva taeniata (p. 98) ORIGINAL DESCRIPTION: P. taeniata. Bandirte D. Mart. Conch. 2. t.49. f.530. Vielleicht nur eine Abänderung der vorigen (this is ispidula), von der sie sich allein durch das dunkle, einfarbige Querband am obern Ende der ersten Windung unterscheidet. DISCUSSION. The Martini figure had already been used by Rôding for ispida. STATUS: objective junior synonym of ispida (Rôding, 1798). Note. This 1s: - ispidula (L., 1758) according to BURCH & BURCH (1960). - a form of oliva (L. 1758) for WAGNER & ABBOTT (1978) and ZEIGLER & PORRECA (1969). - a subspecies of o/iva (L., 1758) for PETUCH & SARGENT (1986) tentorium (p. 95) ORIGINAL DESCRIPTION: P. tentorium. Portobello D. V.Porphyria Linn. Gm.p.3438. Mart. Conch.2 .t.46. f.485.486. Das türkische Zelt. DISCUSSION. One should note that Rôding also used the vernacular "Portobello-Dattel" for porphyria (species 386). Both Martini figures clearly depict porphyria (L., 1758) and had indeed been utilised in the original description by Linnaeus. STATUS: objective porphyria (L., 1758). junior synonym of Note. This is spelled "fentoria” by BURCH & BURCH (1960) and PETUCH & SARGENT (1986). 10. The Oliva of Fischer, 1807. G. Fischer von Waldheim described in the Museum Demidoff (in French) a collection given by Paul de Demidoff to the Imperial University of Moscow. This collection, long thought lost during the siege of Moscow by Napoleon, has been recently retrieved (IVANOV &KANTOR, 1991). TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 The species considered new by Fischer are marked by "m." or "mihi”. Fischer is the first author to have placed all his species in the genus Oliva. Fischer, a generally accepted author, appears however not to be consistently binominal, describing some species (e.g. the "Olive lettrée" only by vernacular names). fusca (p. 160, # 19-30) ORIGINAL DESCRIPTION: 19. 20. L'olive nègre, unie; la base de la spire recourbée, la columelle obliquement striée. Oliva fusca m. Voluta oliva Lin. Gm. 3439.17. Bosc.5.37. Martini 2.t.45.f.472-473. Knorr.5.t.28.f.6. - se touve dans la mer des /ndes. 21. 6. l' Olive à robe brune plus claire avec des raies transversales plus foncées. 22-25. y. l' Olive à robe brun - clair passant au rouge où au jaune. 26. à. l' Olive à robe brune avec une bande ou zone, au milieu, tachetée de noir. 27. 28. &. l' Olive brune à stries longitudinales plus foncées. 29. €. l' Olive blanchâtre avec des taches irrégulières couleur d'olive. 30. 8. l' Olive verdâtre avec des desseins en zigzag. DISCUSSION. O. fusca Fischer. 1807 is a homonym of fusca (Link, 1807). We know the exact date of publication neither of the Museum Demidoff (Fischer, 1807) nor of the Beschreibung (Link, 1807) and we are thus unable to decide on priority. Bosc refers to a vast assortment of entirely unrelated Olives. This species is represented in Moscow by a series. The lectotype selected by IVANOV & KANTOR (1991) was identified as vidua (Rôding, 1798), an objective junior synonym of nigrita (Karsten, 1789). STATUS: subjective junior synonym of nigrita (Karsten, 1789). Note. This is oliva (L., 1758) according to BURCH & BURCH (1960) and WAGNER & ABBOTT (1978). guttata (p.162, # 133) ORIGINAL DESCRIPTION: 133. Olive Girol, Alongée, lisse, la spire courte et lisse, la bouche très ouverte et violette. Oliva guttata mihi. Elle a été figurée par Lister 720.6. Martini 2. p. 161. t. 46. 493. 494. le Girol d' Adanson p. 61. PI. 4. f. 6. -La partie en est inconue. DISCUSSION. The specimen of guftata in the collection 1s said to have been lost before 1872 (IVANOV, DL. & Yu. KANTOR, 1991). The figure of Adanson 1s not clearly recognisable. Figs. 472-473 of Martini were utilised for olivacea (Karsten, 1789), an earlier name for tessellata Lamarck, 1811 (who indeed refers to the same figures). The figure of Lister, as well as the description agree with that identification. STATUS: objective junior synonym of olivacea (Karsten, 1789). Note. This is: - annulata (Gmelin, 1791) according to BURCH & BURCH (1960) and WAGNER & ABBOTT (1978). - a color form of annulata (Gmelin, 1791) for PETUCH & SARGENT (1986). plicata (p. 161, # 90-92) ORIGINAL DESCRIPTION: 90. L'olive bossue Ovale unie, le second tour de la spire enfoncé, trois plis distincts de la columelle, dont le premier très élevé. Oliva plicata mihi. Je n'en connois pas de figure. Elle est ovale, blanche ou verdâtre, ponctuée de brun de différente manière. La lèvre est épaisse, distante dans toute sa longueur. Elle est de la grandeur de l' Olive nègre. 91. Variété de la même, jaune ponctuée de brun. La patrie en est inconnue. DISCUSSION. The specimen of plicata in the collection 1s said to have been lost before 1872 (IVANOV & KANTOR, 1991). This name is unidentifiable and might even not apply to an Oliva. STATUS: nomen dubium. 71 APEX 9(2/3): 51-78, juillet 1994 One should also consider Fischer's ispidula, this name being still available for an Oliva since ispidula (L, 1758) has been shown not to belong to this genus). The specimen of ispidula in the collection 1s said to have been lost before 1872 (IVANOV & KANTOR, 1991). The argument used in the case of Born also applies here. The first reference for this species is "Voluta ispidula Gmel. 3442. n. 23". This is éspidula Land it can be concluded that Fischer refers to the species already described by Linnaeus. 11. The Ofiva of Montfort, 1808. In his Conchyliologie Systématique, Denys de Montfort did not describe species, but only genera. For the genus Oliva, he chose for type species Oliva panamensis. panamensis (p. 387) ORIGINAL DESCRIPTION. Olive de Panama. Oliva panamensis seu porphyrius. Voluta porphyria Linn. sp. 61...etc.. (follows a long list of references, previous illustrations, the names in different languages, and a description). DISCUSSION. The porphyria (L., 1758), the cited illustrations and explicit reference to the description leave no doubt whatsoever on the identity of the species. It is evident that the author himself considered that the two names porphyria and panamensis apply to the same animal. STATUS: Objective junior porphyria (L., 1758). synonym of 12. The Oliva of Perry, 1811. In his Conchology, Perry introduced three new names. The five Oliva illustrations of Perry are quite stylized and some features are obviously exaggerated. Perry gave no references and did not cite previous illustrations. 72 The pre-Lamarckian Oliva porphyracea (PI. 41) ORIGINAL DESCRIPTION: No. 2. OLIVA PORPHYRACEA. Shell dark purple and white, having three belts or circles enveloping the body; the spire also variegated with dark purple spots; the mouth red. From a shell in the Museum of Mr. Latham. DISCUSSION. The description are quite compatible with the Pacific form of ©. miniacea (Rôding, 1798). STATUS: subjective junior miniacea (Rôding, 1798). Note. This is porphyria (L., 1758) according to WAGNER & ABBOTT (1978), ZEIGLER & PORRECA (1978), PETUCH & SARGENT (1986). illustration and the synonym of leveriana (PI. 41) ORIGINAL DESCRIPTION: No. 3. OLIVA LEVERIANA. Shell of a pale purple and gray, richly studded and adorned with a close net pattern, inclosing the whole body; the columella covered with small branched flutings of a white colour, the general colour of the pattern a reddish pink, formed into angular marks. From a shell formerly in the Museum of the late Sir Ashton Lever, in honour of whose zeal for the promotion of natural history and the sciences, | have taken this opportunity of naming it. DISCUSSION. The drawing and the description leave no doubt that this is porphyria (L., 1758). STATUS: subjective porphyria (L., 1758). junior synonym of subviridis (PI. 41) ORIGINAL DESCRIPTION: No. 5. OLIVA SUBVIRIDIS. Shell of an olive green, interspersed with curious marks of dark brown, placed in the form of belts: the mouth gray, the girdle at the base of a rich brown colour. From a shell in the collection of Mr. Jennings of Chelsea, and supposed to be a native of the southern hemisphere. DISCUSSION. By a stretch of imagination, one could interpret the description and the rather TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva çcaricatural figure as depicting a specimen of either e/egans Lamarck, 1811, reticulata (Rôding, 1798) or even fricolor Lamarck, 1811. STATUS: nomen dubium. Note. This is a color form of fricolor Lamarck. 1811 according to PETUCH & SARGENT (1986) zigzag (PI. 41) ORIGINAL DESCRIPTION. No. 4. OLIVA ziGZAG. Shell pale yellow, thickly interspersed with brownish lines in an irregular and waving pattern, from whence its name; the mouth and girdle at the base of a strong orange colour. À native of Ceylon. DISCUSSION. The drawing and the description leave no doubt that this is rericulata (Rôding. 1798). STATUS: subjective junior synonym of reticulata (Rôding, 1798). Note. This 1s: - a variety of sanguinolenta Lamarck. 1811 for DAUTZENBERG (1927). - a color form of reticulata (Rüding. 1798) according to WAGNER & ABBOTT (1978), ZEIGLER & PORRECA (1969). PETUCH & SARGENT (1986). Acknowledgements. We are most indebted to Mrs. Kathie Way (Natural History Museum, London) for allowing us access to the Linnean types and providing us with photocopies of rare ancient works. We thank Dr. Jacky Van Goethem (IRSc.N.B.) for access to the books of the Dautzenberg Library, Dr. Patrick Grootaert (I.R.Sc.N.B.) for his kind collaboration, and we are especially grateful to Mr Antoine Lievrouw (I.R.Sc.N_B.) for his constant help. We thank Dr. Henry Coomans (Zoëülogisch Museum, Amsterdam) for his kind and valuable advice. Index to the names. valid names are in bold. amethystina (Rôding, 1798): valid. amoena (Rôding, 1798): nomen nudum. annulata (Gmelin. 1791): nomen dubium. arachnoidea (Rôding, 1798): objective synonym of spicata (Rôding, 1798). aurata (Rôding, 1798): nomen nudum. aurata (Link, 1807): objective Junior synonym of amethystina (Rôding, 1798). aurora (Solander in Dillwyn, 1817): manuscript name. baltheata (Solander in Düillwyn, 1817): manuscript name. bulbosa (Rôding, 1798): valid. caerulea (Rôding, 1798): valid. carneola (Gmelin, 1791): valid. carneolus (Gmelin, 1791): incorrect original spelling of carneola (Gmelin, 1791). cingulata (Rôding, 1798): nomen nudum. coerulea (Link, 1807): incorrect spelling for caerulea (Rôding. 1798). coffea (Rôding, 1798): objective Junior synonym of carneola Gmelin, 1791. conoidea (Rôding, 1798): nomen nudum. cornea (Rôding, 1798): nomen nudum. crassa (Gmelin, 1791): not an Oliva. cruenta (Solander in Dillwyn, 1817): manuscript name. dealbata (Rôding. 1798): nomen dubium. Jenestrata (Rôding, 1798): nomen dubium. Jasciata (sp. 387, Rôding. 1798): sub]. junior synonym of porphyria (L.. 1758). Jfasciata (sp. 411, Rôding, 1798): junior homonym of /asciata (sp. 387, Rôding, 1798). fimbriata (Rôding, 1798): nomen nudum. Julgurator (Rôding, 1798): valid. Jfusca Fischer, 1807: subjective junior synonym of nigrita (Karsten, 1789). Jfusca (Link, 1807): subjective jumior synonym.of nigrita (Karsten, 1789) griseola (Rôding, 1798): nomen nudum. guttata Fischer, 1807: objective junior synonym of olivacea (Karsten, 1789). 73 APEX 9(2/3): 51-78, juillet 1994 APEX 9(2/3): 51-78, juillet 1994 hepatica (Rôding, 1798): nomen nudum. incrassata (Lightfoot, 1786): valid. "O intertincta Rôding, 1798" (in Wagner & Abbott, 1978): not of Rôding. isabella (Rôding, 1798): nomen nudum. ispida (Rôding, 1798): nomen dubium. ispidula (L., 1758): not an Oliva. ispidula (Abel, 1787): non binominal. ispidula (Born, 1778): junior homonym of ispidula (L., 1758). ispidula (Schrôter, 1782): junior homonym of ispidula (L., 1758). ispidula (Gmelin, 1791): junior homonym of ispidula (L., 1758). ispidula Fischer, 1807: junior homonym of ispidula (L., 1758). labradoriensis (Rôding. 1798): nomen dubium. "leucophaea Gmelin" (in Môrch. 1850): not of Gmelin. leveriana Perry, 1811: synonym of porphyria (L., 1758). subjective junior litterata (Rôding. 1798): nomen dubium. maculata (Solander in Düillwyn. 1817): manuscript name. mauritana (Abel, 1789): non binominal. mica (Rôding, 1798): nomen dubium. miniacea (Rôding, 1798): valid. miniata (Link,1807): objective junior synonym of miniacea (Rôding. 1798). nigrita (Karsten, 1789): valid. oculata (Rôding, 1798): nomen nudum. oliva (L., 1758): valid. olivacea (Karsten. 1789): valid. ornata (Rôding, 1798): nomen nudum. paleacea (Rôding, 1798): nomen nudum. panamensis Montfort. 1808: objective junior synonym of porphyria (L., 1758). papyracea (Rôding, 1798): nomen nudum. plicata Fischer, 1807: nomen dubium. porphyracea Perry, 1811: sub}. junior syn. of ©. miniacea (Rôding, 1798) porphyria (L., 1758): valid. porphyrea (Abel, 1789): non binominal. punctata (Rôding, 1798): nomen nudum. quercina (Rôding, 1798): nomen nudum. reticulata (Abel, 1789): non binominal. reticulata (Rôding, 1798): valid. ruffina (Gmelin, 1791): not an Ofiva. 74 The pre-Lamarckian Oliva ruffina (Rôding, 1798): nomen nudum. sepultura principis (Abel, 1789): non binominal. sepultura principis (Rôding, 1798): nomen dubium. sericea (Rôding, 1798): valid. spicata (Rôding, 1798): valid. subviridis Perry, 1811: nomen dubium. taeniata (Link, 1807): objective junior synonym of ispida (Rôding, 1798). (Link, 1807): synonym of porphyria (L., 1758). tentorium objective junior tessellata Lamarck, 1811: objective junior synonym of olivacea (Karsten, 1789). textilis (Rôding. 1798): nomen nudum. tigris (Rôding, 1798): nomen nudum. tuberosa (Rôding, 1798): subjective Junior synonym of bulbosa (Rüding, 1798). tumida (Rôding, 1798): nomen nudum. turgida (Rôding, 1798): nomen nudum. umbrosa (Rôding, 1798): nomen dubium. undulata (Rôding, 1798): nomen nudum. variabilis (Rôding, 1798): nomen nudum. variegata (Rôding, 1798): nomen dubium. ventricosa (Solander in Dillwyn, 1817): manuscript name. vellus aureum (Abel, 1789): non binominal. vellus-aureum (Rôding. 1798): nomen dubium. vidua (Rôding., 1798): objective junior synonym of nigrita (Karsten, 1789). ziczac (Rôding, 1798): nomen nudum. zigzag Perry, 1811: subjective junior synonym of reticulata (Rôding. 1798). TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER Index to the cited figures The references to the figures are verbatim those used by the authors of the descriptions. ADANSON p. 61. PI. 4. f. 6: guttata Fischer, 1807. ARGENVILLE, d' t. 16. f. K: porphyria (Linnaeus, 1758). t. 16. f. R: oliva (Linnaeus, 1758). GUALTIERI t. 23. f B: oliva (Linnaeus, 1758). t. 24. f. O: porphyria (Linnaeus, 1758). t. 24. f. P: porphyria (Linnaeus, 1758). KAEMMERER t. 3. fig. 7: tuberosa (Rôding, 1798). t. 3. fig. 8: tuberosa (Rôding, 1798). KNORR .t. 15. f. 7: umbrosa (Rôding, 1798). t. 10. f6: amethystina (Rôding, 1798) t. 10. f 7: amethystina (Rôding. 1798). t 17. f. 3: fasciata (Rôding, 1798). t. 19. f 3: ispida (Rôding. 1798). t n SD D — 1798). 5. tab. 26. fig. 4: quercina (Rôding, 1798). 5. t 26. f. 4: sepultura principis B (Rôding, 1798). .t. 27. fig. 5: quercina 8 (Rôding, 1798). . t. 28. f. 6: fusca (Fischer, 1807). .t. 34. f. 4: dealbata (Rüding, 1798). 6.t. 34. f S: dealbata (Rôding, 1798). Un Un (o)} LISTER 4. f. 10. c. 1. t. 2: oliva (Linnaeus. 1758) (not seen) t. 664. f. 8: crassa (Gmelin. 1791). 720. 6: porphyria (Linnaeus, 1758). 731. f. 20: annulata (Gmelin, 1791) t. 746. f. 40: tumida (Rôding, 1798). MARTINI tab. 45. fig. 472: nigrita (Karsten, 1789). 2. t. 45. f. 472: fusca (Fischer, 1807). 2.t. 45. f. 472: vidua (Rôding, 1798). tab. 45. fig. 473: nigrita (Karsten, 1789). 2. t. 45. f. 473: vidua (Rôding, 1798). 2.t. 45. f. 473: fusca (Fischer, 1807). . 19. f 1: sepultura principis y (Rôding, The pre-Lamarckian Oliva APEX 9(2/3): 51-78, juillet 1994 .45. f 476: miniata (Link 1807). . 45. f 476: miniacea (Rôding, 1798). . 45. f. 477: miniata (Link. 1807). . 45. f. 477: miniacea (Rôding, 1798). . 45. f. 478: variegata (Rôding, 1798). . 45. f. 479: variegata (Rôding, 1798). . 45. f. 480: variegata à (Rôding, 1798). . 45. f. 480: sepultura principis y (Rôding, 1798). . 45. f. 481: variegata à (Rôding, 1798). . 45. f. 481: sepultura principis y (Rôding. 1798). 46. f. 485: tentorium (Link, 1807). 46. f. 486: tentorium (Link, 1807). 46. f 488: litterata (Rôding, 1798). 46. f. 490: vellus-aureum (Rôding, 1798). 46. f. 491: amethystina (Rôding, 1798). 46. f. 491: aurata (Link, 1807). 46. f. 492: amethystina (Rôding, 1798). . 46. f. 492: aurata (Link, 1807). 2. tab. 46. fig. 493: olivacea (Karsten, 1789). p. 161. t. 46. 493: guttata Fischer, 1807. 2. tab. 46. fig. 494: olivacea (Karsten, 1789). p. 161. t. 46. 494: guttata Fischer, 1807. 2. t. 46. f. 495: carneolus (Gmelin, 1791) 2. t. 46. f. 495: coffea (Rôding, 1798). Vol. IT- f 499: incrassata (Lightfoot, 1786). Vol. II- f 500: incrassata (Lightfoot, 1786). T. 2. tab. 47. fig. 501: nigrita (Karsten, 1789). 2.t. 47. f. SOL: fusca (Link, 1807). 2.147. f 502: fenestrata, (Rôding, 1798). 2.147. f 507: bulbosa (Rôding, 1798). 2.147. f 508: bulbosa (Rôding, 1798). 2.148. f. 509: spicata (Rôding, 1798) 2.t48. f. 509: arachnoidea (Rôding, 1798). 2 t. 48. f 510: spicata (Rôding, 1798). 2.t 48. f. 510: arachnoidea (Rôding, 1798). .t 48. f 512: reticulata (Rôding, 1798). .t. 48. f 515: coerulea (Link, 1807). . t. 48. f. 516: coerulea (Link, 1807). . 148. f 518: caerulea (Rôding, 1798). t. 49. f. 522: sspida Ê (Rôding, 1798). . 49. f 523: ispida Ê (Rôding, 1798). . 49. f. 524: ispida (Rôding, 1798). . 49. f. 525: ispida (Rôding, 1798). . 49 f. 527: mica var à (Rôding, 1798). . 49 f. 528: mica var à (Rôding, 1798). . 49 f. 529: mica var &« (Rôding, 1798). ENOMONO AO EN OS mm me D LE et ee me mt et OSSI CE LOS CEE D D D D D D D © D © D : tt tm 75 APEX 9(2/3): 51-78, juillet 1994 2. t. 49. f. 530: ispida (Rôding, 1798). 2. t. 49. f. 530: raeniata (Link. 1807). 2.t 48. f 533: reticulata (Rôding, 1798). (note: fig. 533 is in PI. 491). . 49. f. 535: ispida «à (Rôding, 1798). . 49. f. 537: umbrosa (Rôding, 1798). . 51. f. 559: sericea (Rôding, 1798). . 51. f 561: sericea (Rôding, 1798). . S1. f. 562: fulgurator (Rôding. 1798). . 51. f. 563: sepultura principis B (Rôding, 1798). 2.t. 51. f. 564: annulata (Gmelin. 1791) D © D D D D + et 7 7 et PETIVER. Gazophil., 1767. t. 59. f 8: ispidula (Linnaeus, 1758). REGENFUSS 2.t. I. f 2: oliva (Linnaeus, 1758). 8. t. 2. f. 15: porphyria (Linnaeus, 1758). RUMPHIUS. t. 39. f. 1: porphyria (Linnaeus, 1758). . 39. f 2: oliva (Linnaeus, 1758). 39. f. 3: oliva (Linnaeus. 1758). 39. f. 4: oliva (Linnaeus. 1758). 39. f. 4: sepultura principis (Rôding, 1798). 39. f. 5: oliva (Linnaeus. 1758). 39. f. 5: caerulea (Rôding. 1798). 39. f. 6: ispidula (Linnaeus. 1758). 39. f. 7: ispidula (Linnaeus, 1758). Ce EE ee EE REFERENCES. ABEL, J.C.A.M., 1787. Die Conchylien in dem Naturalkabinet Seiner Hochfürstlichen Gnaden des Herrn Fürsten und Bischofs von Konstanz …. Kaspar Graf, Bregenz. ADANSON, M. 1757. Histoire Naturelle du Sénégal. Coquillages. Paris. ARGENVILLE, A.JD. d', 1742. L ‘Histoire naturelle éclaircie dans une de ses parties principales. La Conchyliologie qui traite … Paris. BORN, I. 1778. Index Rerum Naturalium Musei Caesarei Vindobonensis. Vienna. BORN, I, 1780. Testacea musei Caesarei Vindobonensis. Vienna. BONNANI, U.A., 1681. Ricreatione dell' occhio e della mente .… Rome. 76 The pre-Lamarckian Oliva BONNANI, UA.. 1709. Musaeum Kircherianum, sive …. Rome. Bosc, L.A.G. 1802. Histoire naturelle des Coquilles, Paris. BURCH, J.Q. & RL. BURCH, 1960. Catalog of recent and fossil Olives. Minutes Conch. Club South. Calif. 196: 1-46. BURCH, J.Q. & RL. BURCH, 1967. The family Olividae. Pacific Science 21(4): 503-522. DALL, W. H. 1915. An Index to the Museum Boltenianum. Smithsonian Institution Publ. 2360: 1-64. DALL, W. H., 1921. Species named in the Portland Catalogue. I. American. Nautilus 34: 97-100, 124-132. DA MOTTA, B., 1985. Two new Oliva species. La Conchiglia 17(192-193): 8-9. DANCE, SP. 1962. The authorship of the Portland Catalogue. J. Soc. Bibl. nat. Hist. 4: 30-4. DAUTZENBERG, P., 1927. Olividés de Nouvelle Calédonie et de ses dépendances. J.Conchyl. 71: 1-72, 103-147. DILLWYN, L.W., 1817. À descriptive catalog of recent shells according to the Linnean method, with particular attention to the synonymy. Cornhill, London. DUCHAMPS, R. & B. TURSCH, 1994 A note on the Museum Leskeanum. Apex 9(1): 11-16. FISCHER von WALDHEIM, G., 1807. Museum Demidoff. ou Catalogue systématique et raisoné des Curiosités de la Nature et de l'Art. Données à l'Université Impériale de Moscou …. Moscow. GMELIN, JF. 1791. Caroli a Linné Systema Naturae per regna tria naturae. Editio decima tertia, aucta, reformata. Leipzig. GUALTIERI, N., 1742. Index Conchyliorum …. Florence. Testarum HANLEY, S., 1855. Zpsa Linnaei Conchylia. The shells of Linnaeus determined by his manuscripts and collection. Williams and Norgate, Londres. HOLM, A, 1957. Specimina Linnaeana :i Uppsala bevarade zoologiska samlingar fran Linnés tid. Acta Universitatis Upsalensis 6: 1- 67. IREDALE, T., 1916. Solander as a conchologist. Proc.Malac. Soc. Lond. 12(2-3): 85-93. TURSCH, DUCHAMPS & GREIFENEDER TURSCH, DUCHAMPS & GREIFENEDER The pre-Lamarckian Oliva IVANOV, DL. & Yu. KANTOR, 1991. Paul Demidoffs malacological collection in the Zoological Museum of Moscow University. Publishing House of Moscow State University. KÂMMERER, C.L. 1786. Die Conchylien im Cabinette des Herrn Erbprinzen von Schwarzburg-Rudolstadt. Rudolstadt. KARSTEN, G., 1789. Museum Leskeanum. Regnum Animale. Quod ordine systematico …. Müller, Lipsiae. KAY, E.A., 1965. The reverend John Lightfoot. Daniel Solander and the Potland Catalogue. Nautilus 79(1): 10-19. KEEN, M. 1971. Seashells of tropical West America. Stanford University Press. KILBURN, R.N. 1980. The genus Oiva in Southern Africa and Mozambique. Ann. Natal Mus. 24(1): 221-231. KLEIN, JT. 1753. Tentamen Methodi Ostracologicae, sive Dispositio Naturalis Cochlidium et Concharum, Lugduni Batavorum. KNORR, G.W.. 1760-73. Les délices des veux et de l' esprit, ou collection générale Nurenberg. LAMARCK. JB.P.A. 1810-11. Détermination des espèces de Mollusques testacés continuation du genre Ovule, Tarrière. Ancillaire et Olive. Ann. Mus. Hist. Nat. 16 (for 1810): 300-328. Paris (Jan.-Mar. 1811). LINK, DHEF., 1807. Beschreibung der Naturalien-Sammlung Universität zu Rostock. Adlers Erben. Rostock LIGHTFOOT, J. 1786. À catalogue of the Portland Museum, lately the property of the Duchess Dowager of Portland, deceased, which will be sold at auction, by Mr. Skinner and Co. …. London. LINNAEUS, C., 1758. Systema Naturae per regna tria Naturae. Editio decima. Reformata. Vol.1: Regnum Animale. Holmiae. LINNAEUS, (C. 1764. Museum Reginae Ludovicae Ulricae etc. Holmiae. LISTER, M., 1682-1695. Historiae siv. Synopsis Methodicae Conchyliorum … London. MARTINI, FH.W., & J.H. CHEMNITZ, 1769- 1795. Neues Systematisches Conchylien- Cabinet … Nurenberg. APEX 9(2/3) 51-78, juillet 1994 MONTFORT, D. de, 1808 Conchyliologie systématique, et classification méthodique des coquilles. Vol. 1. Paris. MGRCH. O AL. 1850 Catalogus Conchyliorum quae reliquit C.P. Kierulf …. Hafniae. OLSSON, A. A., et S. P. DANCE, 1966. The Linnean Olives. Bull. Am.Paleont. 50: 227. PERRY, G.. 1811. Conchology, or the natural history of shells. London. PETIVER, J., 1767. Opera Historiam Naturalem spectantia. Gazophylacii Naturae & Artis. John Millan, White-Hall. PETUCH, E.J. and D.M. SARGENT, 1986. Atlas of the living Olive shells of the world. CERF editions. Charlottesville, VA. REGENFUSS. FM. 1758. Auserlesne Schnecken Muscheln und anderre Schaalthiere …. Choix de Coquillages et de Crustacés … Copenhagen. REHDER. HA. 1945 A note on the Bolten Catalogue. Nautilus 59(2): 50-52. REHDER. H.A.. 1967. Valid zoological names of the Portland Catalogue. Proc. US. Nat. Mus. 121(3579): 1-47. RODING, PF. 1798. Museum Boltenianum sive Catalogus Cimeliorum . Pars Secunda. Hamburg. RUMPHIUS, GE. 1705. D' Amboinsche Rariteitkamer, Schaalvissen, Krabben, Kreeften, Hoorntjes en Schulpen Amsterdam. RUTSCH, REF. 1956. Remarks on I. Born's Index Rerum Naturalium Musei Caesarei Vindobonensis. 1778. Nautilus 69(3): 78-79. SEBA. À. 1734-65 Locupletissimi rerum naturalium theasauri accurata descriptio, …… Amsterdam. SCHRÔTER, JS. 1782. Musei Gottwaldiana. Testaceorum Stellarum Marinarum el Coralliorum quae supersunt Tabulae. Nurenberg. 77 APEX 9(2/3): 51-78, juillet 1994 The pre-Lamarckian Oliva SCHRÔTER, JS. 1783 Einleitung in die Cochylienkenntniss nach Linné. J.J. Gebauer. Halle. TURSCH, B.. L. GERMAIN and D. GREIFENEDER, 1986. Studies on Olividae. IV. Oliva annulata Gmelin, 1791 (of authors): a confusion of species. /ndo-Malayan Zoology 3: 189-216. TURSCH, B., O. MissA & J. BOUILLON, 1992. Studies Fon Olividae XIV. The taxonomic structure of Oliva oliva (auct.). Apex 7(1): 3-22. WAGNER, R.JL. & ABBOTT, RT. (1978). Standard Catalog of Shells. 3rd. Edition. American Malacologists Inc. Greenville, Delaware. WEINKAUFF, H.C., 1878 Systematisches Conchylien Cabinet. Bnd.V. Abtll. Die Gattung Oliva. Nurenberg. ZEIGLER, R.F. and H.C. PORRECA., 1969. Olive Shells of the World. Rochester Polychrome Press, Rochester, N.Y. 78 TURSCH, DUCHAMPS & GREIFENEDER PIZZINI, NOFRONI & OLIVERIO A new Caecum Contribution to the knowledge of the family Caecidae. 1. À new Caecum from Canary Islands (Caenogastropoda: Rissooïidea) * Mauro PIZZINI Largo della Caffarelletta 6. I1-00179 Roma, Italia. Italo NOFRONI Via Benedetto Croce 97. I-00142 Roma, Italia. Marco OLIVERIO Dipartimento di Biologia Animale e dell'Uomo, Università di Roma "La Sapienza”. Viale dell'Università 32. 1-00185 Roma, Italia. ABSTRACT. A very peculiar, new species of the meiobenthic gastropod family Caecidae is described and figured from the Canary Islands: Caecum lightfootae n.sp. It is compared with all the known congeneric species from this area. KEY WORDS: Caecum, new species, meiobenthic, North-East Atlantic, Recent, marine. INTRODUCTION The Caecidae are a worldwide distributed family of rissooidean gastropods with secondary uncoiled teleoconch, adapted to the meiobenthic life. Despite a good deal of efforts, past and recents, to the study of their taxonomy (see 1.e. DE FOLIN 1867-1876, 1867, 1875, 1880: CARPENTER 1858-1859: MOORE 1962; VAN AARTSEN 1977; LIGHTFOOT 1992a, b, c, 1993a, b), Caecidae are still one of the less-known prosobranch family. The use of classical morphology in their taxonomy is often hampered by the difficulty in individuating good characters in their simple, unusual tube- like shells. Even worse is the situation as for their supraspecific classification. Anatomical data-sets are still poor and insufficient to create a good phylogenetic scheme of the family. A revision of the North-East Atlantic members of the family, presently in progress by the authors, is revealing some important novelties as for their taxonomy. Preliminary to this revision, some still undescribed species need to be presented. Working on some samples from the Canary Islands, a very peculiar species of Caecum was sorted out. It is a new species, different from all the other species known from that area. SYSTEMATICS Superfamily RISSOOIDEA Gray J.E., 1847 Family CAECIDAE Gray ME. 1850 Genus Caecum Fleming, 1813 Caecum lightfootae n. sp. DESCRIPTION (holotype's measurements between parentheses ) Teleoconch uncoiled, tube-like. Shell very small for the genus, semitransparent, glossy. Tube very curved, regularly bent, slightly subconical for one third of the length (on the septum side), then nearly cylindrical for the rest (Figs. 1, 2). Surface seemingly smooth, with a microsculpture of only thin and irregular growth lines. Aperture regularly circular, with a more or less pronounced varix, always present (Fig. 3). Septum not retracted, with the lateral outline subtriangular and blunt. Mucro protruding, and right-handed when observed frontally (Figs. 4, 5). Protoconch and growth stages not identified amidst the available material. Shell whitish, with irregular * Work partly supported by CNR (Comitato Ambiente) funds. 19 APEX 9(2/3): 79-82, juillet 1994 APEX 9(2/3): 79-82, juillet 1994 wax-vitreous areas. Operculum thin, corneous, yellow-brownish; external surface with 4-5 concentrical ridges, the central and the outer part smooth. Soft parts not studied at present. Dimensions: lenght 1.15-1.30 mm (1.23); tube diameter at the septum level 0.24-0.26 mm (0.25); tube diameter at the aperture 0.34-0.36 mm (0.35). Material examined: Punta Blanca, Puerto Santiago (Is. Tenerife, Canary Islands) - 30 m. M.Oliverio leg. 12-IX-1992: 28 shells. Arigana (Is. Gran Canaria, Canary Islands) - 1 m, F.Gubbioli leg.: 1 specimen. The type material, all from the type locality. has the following location: Holotype and one paratype Museo Civico di Zoologia, Roma 2 paratypes Laboratorio di Malacologia. Università di Bologna 2 paratypes Muséum National d'Histoire Naturelle, Paris 2 paratypes Natural History Museum, London 2 paratypes History, Gainsville 2 paratypes Australian Museum, Sydney 2 paratypes Swedish Museum of Natural History, Stockholm 2 paratypes Museo Insular Naturales, Sta Cruz de Tenerife Other paratypes are stored in the private collection of M. Pizzini (2). I. Nofroni (2). M. Oliverio (2), C. Schander (2), L. Tringali (2). G. Ambrosiano (1), F. Gubbioli (1). Florida Museum Natural Ciencias Type locality. Punta Blanca, Puerto Santiago, Is. Tenerife, Canary Islands (Spain). At present the species is known only from Canary Islands, Is. Tenerife and Is. Gran Canaria. Etymology. The species is dedicated to a very keen american malacologist: the late Mrs. Joanne Lightfoot, known to the specialists of this family for her studies on the Caecidae of North America. The first author had the possibility to appreciate her scientific and human endowments, during a short but intense mail correspondence. Discussion. Although the morphological features of C. lightfootae are so peculiar to render it unic and easily separable from any other Canaric Caecidae, we prefer to compare it with all the other species known from this area. NORDSIECK & TALAVERA (1979: 84) recorded only five Caecum species from the Canary Islands: C. atlantidis Watson, 1897, C. 80 A new Caecum trachea (Montagu, 1805), C. glabrum (Montagu, 1803), C. vitreum Carpenter, 1858 and C. elegantissimum Carpenter, 1858. Evidentiy, they skipped C. clarkii Carpenter, 1858, though its type locality was just Is. Tenerife. To this list a seventh species should be added, namely C. armoricum DE FOLIN, 1869 recently recorded by HOEKSEMA & SEGERS (1993: 86) for the Canary Islands (Gran Canaria). C. lightfootae has à septum similar to that of C. clarkii and Cvitreum. It differs by its smaller size, by the lack of a longitudinal sculpture more (c/arkii) or less (vitreum) evident, by the presence of the annular varix at the aperture, and finally by its pronounced curvature. C. glabrum and C. armoricum are easily separable from the new species by a different septum (dome-shaped in the first, and nail- shaped in the second), and by their lack of an apertural varix. C._ atlantidis, C. trachea and C. elegantissimum are completely different and unrelated species, with an evident annular sculpture, completely absent in C. lightfootae. A further european species, namely C. auriculatum De Folin, 1868 (whose type series we have studied), needs a brief comparison, although it is not recorded for the Canary Islands. Its septum is nearly hemispheric, with a mucro hear-like, usually right-handed; the colour is uniformly white vitreous. Finally, C. auriculatum 1s more curved, has a larger mean size, and the apertural varix is more evident than in C. lightfootae. Acknowledgements. We are grateful to Dr. Christoffer Schander (University of Gôteborg, Sweden) for the numerous (too much!) SEM pictures he realized for us. Figs. 1-5 (opposite). Caecum lightfootae n.sp. holotype. Is. Tenerife. 1. General view (frontal). 2. General view (lateral). 3. Aperture. 4. Apex (frontal view). 5. Apex (lateral view). PIZZINI, NOFRONI & OLIVERIO APEX 9(2/3): 79-82, juillet 1994 A new Caecum © œ >. = © Ÿ Z. © e4 Lu © A Z N S [ew APEX 9(2/3): 79-82, juillet 1994 A new Caecum PIZZINI, NOFRONI & OLIVERIO REFERENCES AARTSEN J.J. VAN. 1977. Revision of the East Atlantic and Mediterranean Caecidae. Basteria 41: 7-19. CARPENTER P.P., 1858-1859. First steps towards a Monograph of the Caecidae, a family of Rostriferous Gasteropoda. Proc. Zool Soc. London (1858): 413-432 [1958], 433-444 [1859]. DE FOLN L., 1867-1876. Les fonds de la mer. Paris, vol.1: 326 pp, 36 pls + pl 21 bis; vol.2: 365 pp. 11 plis. DE FOLIN L., 1867. Descriptions d'espèces nouvelles de Caecidae. J. Conch., Paris, 15: 44- 58. DE FOLIN L., 1875. Monographie de la famille Caecidae. Bayon, 31 pp, 1 pl. DE FOLIN L., 1880. On the Mollusca of the HMS. "Challenger" Expedition - The Caecidae, comprising the Genera Parastrophia, Watsonia and Caecum. Proc. Zool. Soc. London (1879): 806-812. HOEKSEMA DF. & SEGERS W., 1993. On the systematics and distributions of the marine gastropod Caecum armoricum De Folin, 1869 (Prosobranchia, Caecidae). Gloria Maris 31(6): 79-88. LIGHTFOOT J.. 1992a. Caecidae of the Western Atlantic. Part one. Of Sea and Shore 14: 13-26. LIGHTFOOT J., 1992b. Caecidae of the Western Atlantic. Part two. Of Sea and Shore 15: 23-31. LIGHTFOOT J. 1992c. Hawaïian Caecidae. Hawaiian Shell News, 50 (7): 3-5. LIGHTFOOT J., 1993a. Caecidae of the Panamic Province. Part one. Of Sea and Shore 16: 1-13. LIGHTFOOT J., 1993b. Caecidae of the Panamic Province. Part two. Of Sea and Shore 16: 75-87. MOORE D R., 1962. The systematic position of the Family Caecidae (Mollusca: Gastropoda). Bull. Mar. Sci. Gulf Caribb., 12 (4): 695-701. NORDSIECK F. & TALAVERA FG. 1979. Moluscos marinos de Canarias y Madeira (Gastropoda). Aula de Cultura de Tenerife, 208 pp. lam. XLVI. 82 MERLE Muricidae de l'Eocène APEX 9(2/3): 83-91, juillet 1994 Révision des Muricidae de l'Eocène de la falaise de la Côte des Basques à Biarritz (Pyrénées-Atlantiques, France) Didier MERLE Laboratoire de Paléontologie du Muséum national d'Histoire naturelle de Paris 8, rue Buffon F - 75005 PARIS RESUME: Cinq espèces de l'Eocène supérieur de Biarritz (Priabonien de la Côte des Basques, Les Bains) sont décrites. Deux sont nouvelles: Siphonochelus (Laevityphis) biarritzensis et S. (Trubatsa) evainae. Pterynotus (Pterynotus) cf. consobrinus (d'Orbigny, 1852), du Stampien de Gaas (Landes) est cité pour la première fois dans le Priabonien de Biarritz. Murex (Pteronotus) subfiligrana Tournouër in de Bouillé, 1876 est révisé. ABSTRACT: Five species from the Priabonian of Biarritz (Côte des Basques, Les Bains, Pyrénées-Atlantiques, France) are described. Two species are new: Siphonochelus (Laevityphis) biarritzensis and S. (Trubatsa) evainae. Pterynotus (Pterynotus) cf. consobrinus (d'Orbigny, 1852) 1s recorded for the first time in the Priabonian of Biarritz. Previously, the species was only known in the Stampian of Gaas (Aquitaine basin). Murex (Pteronotus) subfiligrana Tournouër in de Bouillé, 1876 1s revised. MOTS-CLES: Muricidae, systématique, phylogénie. Eocène supérieur, Priabonien, Bassin d'Aquitaine, KEY-WORDS: Muricidae, Upper Eocene, Priabonian, Aquitaine Basin, systematics, phylogeny. Cadre stratigraphique: Le Paléogène des falaises de Biarritz a été révisé par Mathelin (1988). Mathelin situe les Marnes des Bains. épaisses de 350 m, au-dessus des Marnes à Turbinolia calcar (-Flabellum calcar). D'après le nannoplancton calcaire, il place les Marnes des Bains dans les biozones NP19 (Zsthmolithus radians) et NP20 (Sphenolithus pararadians), INTRODUCTION Les Muricidae de l'Eocène de la Côte des Basques (Biarritz) sont connus par deux espèces citées par Tournouër in de Bouillé (1876) qui proviennent des Marnes des Bains. A la suite de recherches à l'Université de Bordeaux I, j'ai retrouvé plusieurs Muricidae provenant de cette localité dans la collection Neuville. Cette découverte m'a incité à rechercher dans d'autres collections des Muricidae de l'Eocène de la Côte des Basques. Les collections Castex (Muséum d'Histoire naturelle de Biarritz), Cossmann (Muséum national d'Histoire naturelle de Paris - Laboratoire de Paléontologie) et Tournouër (Institut de Géologie A. de Lapparent) ont livré d'autres exemplaires de cette famille. L'ensemble rassemblé constitue peu d'individus, mais complète nos connaissances sur les Muricidae de l'Eocène supérieur d'Aquitaine. Situation géographique de la Côte des Basques (Les Bains): (Carte géologique 1/50.000, feuille XII-44 (Bayonne), x — 284.125, y = 3138.300. définies par MARTINI (1971). STEURBAUT (in NOLF, 1988) place également les Marnes des Bains dans NP 20. Ces deux biozones permettent de dater le Priabonien terminal. Abréviations utilisées: Collections: UBX: Université Bordeaux I, Talence: MNAN-LP: Muséum national d'Histoire naturelle de Paris (Laboratoire de Paléontologie), BIA: Muséum d'Histoire naturelle de Biarritz; IGAL: Institut de Géologie A. de Lapparent, Cergy. Mensurations: H: hauteur: D: diamètre; HO: hauteur de l'ouverture; LO: largeur de l'ouverture; G: chez les Typhinae, angle du tube par rapport à l'axe (selon Houart, 1991). 83 APEX 9(2/3): 83-91, juillet 1994 Muricidae de l'Eocène MERLE ES À tt Figures 1-2 (échelle: 1 mm) 1. Protoconque de Chicoreus (?) subfiligrana (Tournouër in de Bouillé, 1876), néotype, collection Neuville (UBX, n° TYFIPAL 57.21 ). 2. Protoconque de Pterynotus (Pterynotus) crenulatus (Rôding, 1798), collection Neuville (UBX, n'TYFIPAL 57.2.2). Figures 3-6 (échelle: 1 mm) 3. Profil de l'ouverture et dernière côte de Siphonochelus (Laevityphis) biarritzensis nov sp, paratype, collection Cossmann (MNHN LP n°J00003). 4. Profil de l'ouverture et dernière côte de S. (Trubatsa) evainae nov. sp., paratype, collection Castex (BIA). 5. Profil de l'ouverture et dernière côte de S. (T.) gaasensis (d'Orbigny, 1852), collection Merle (localité: Gaas). 6. Protoconque de S. (L.) biarritzensis nov sp, holotype, collection Neuville (UBX, n° TYFIPAL 57-2:3) 84 MERLE Muricidae de l'Eocène ETUDE SYSTEMATIQUE Famille MURICIDAE Rafinesque, 1815 Sous-famille MURICINAE Rafinesque, 1815 Genre Chicoreus Montfort, 1810 Espece-type: Murex ramosus Linné, 1758 par désignation originale. Chicoreus (?) subfiligrana (Tournouër in de Bouillé, 1876) (Figs. 1, 7,8) Murex (Pteronotus) subfiligrana Tournouër in de Bouillé, 1876: p. 58, pLlIil, fig.7. Localité- type: Biarritz (Côte des Basques, Les Bains). Niveau-type: Marnes à Zurbinolia calcar. Priabonien de Biarritz. Materiel-type: Je n'ai pas retrouvé l'holotype dans la collection Tournouër déposée à l'IGAL, où 1l existe des fossiles de la Côte des Basques. L'holotype a le labre cassé et ne présente que les quatre derniers tours. D'après Tournouër, sa hauteur est de 35 mm et son diamètre de 18 mm. Un spécimen à spire complète, mais avec le canal siphonal et l'ouverture cassés a été retrouvé dans la collection Neuville (UBX, n°TYFIPAL 57.21). Il est désigné ici comme néotype. Autre matériel: 1 fragment de labre, collection Cossmann n° 16453a (MNHN-LP n°J00001). Description: Coquille d'environ 30 mm de hauteur (hauteur estimée) et de 15 mm de diamètre. Protoconque conique composée de 3,5 tours. Téléoconque composée de 6 tours étagés. Sculpture axiale composée de 3 côtes principales subépineuses dans l'intervalle desquelles s'intercale une côte intermédiaire. Sculpture spirale composée de 3 à 4 cordons spiraux sur la spire et de 10 cordons sur le dernier tour. Microsculpture formée de fins cordonnets spiraux croisés par les stries d'accroissement grossières, l'ensemble donnant l'aspect d'une surface écailleuse. Bord columellaire lisse. Labre droit, liré. Canal siphonal court (d'après la figure de l'holotype). Dimensions: Néotype, H: 15 mm (base incomplète); D: 11 mm (ouverture incomplète). Comparaisons: TOURNOUER in DE BOUILLE, 1876 compare ce Muricidae à Murex filigrana von Koenen, 1867, du Lattorfien d'Allemagne. Cette comparaison se justifie par une microsculpture développée et une symétrie à APEX 9(2/3): 83-91, juillet 1994 trois côtes principales. Toutefois, les côtes principales de M. filigrana sont plus développées et plus foliacées. Ses cordons spiraux sont plus fins et son ouverture porte deux cals sur le bas de la columelle. Enfin, les anciens canaux siphonaux sont nettement abaxiaux alors que ceux du M. subfiligrana sont droits. Les caractères de M. filigrana rapellent ceux des Pterynotus actuels de l'Océan Pacifique tels que P. (P.) pellucida (Reeve, 1845) ou P. (P.) tripterus (Born, 1778). A l'instar de Tournouër, je considère les M. filigrana et M. (P.) subfiligrana comme deux epèces distinctes. Discussion sur la position générique:. Les caractères du Murex subfiligrana (symétrie à trois côtes principales, côtes subépineuses, présence d'une microsculpture et bord columellaire lisse, voir HOUART, 1992) permettent un rapprochement avec Chicoreus Montfort, 1810. D'après HOUART (1992), Chicoreus (Chicoreus) se distingue C. (Triplex) par la présence de dent labiale. Le mauvais état de l'ouverture de notre échantillon le plus complet empêche de faire cette distinction. C. (T.) tateiwai Hatai & Kotaka, 1952 , du Miocène inférieur de Corée, ressemble à l'espèce de Biarritz, mais sa microsculpture est moins développée. Murex subfiligrana ne peut se classer dans Phyllonotus Swainson 1833 (que VOKES (1990) admet comme sous-genre de Chicoreus), qui présente souvent une spire plus basse, une sculpture spirale plus grossière et fréquemment plusieurs cals ou une série de rides columellaires. C. (Phyllonotus) initialis Vokes, 1990, de l'Eocène moyen du Texas possède des côtes principales plus foliacées avec deux côtes intermédiaires, et se remarque par ses cordons spiraux plus épais. Genre Pterynotus Swainson, 1833 Espece-type: Murex pinnatus Swainson, 1822 par désignation originale. Sous-genre Pterynotus Swainson, 1833 Pterynotus (Pterynotus) crenulatus (s1.) (Rôding, 1798) (Figs. 2, 9) Purpura crenulata Rôding, 1798: Museum Boltenianum: 144 (d'après Brander, 1766, pl.3, figs 77, 79). 85 APEX 9(2/3): 83-91, juillet 1994 Synonymie: Murex tricarinatus Lamarck, 1803: Ann. Mus. natn. Hist. nat. Paris, v.Il : 223. Murex (Pteropurpura) palensis Magne, 1940: J. de Conch., v. 84: 372. Réference concernant Biarritz: Murex trigonus -Tournouër in de Bouillé, 1876: 50; non Rouault, 1850: 493 , pl.XVII, fig. 17. Materiel examiné: 1 spécimen, collection Neuville (UBX, = N°TYFIPAL" 57.22); "1 spécimen cassé au niveau du labre, collection Cossmann n° 16978 (MNHN-LP). Description: Coquille trapue de 9 mm de hauteur et de 5 mm de diamètre. Protoconque conique, multispirale dont 2,5 tours sont conservés, le sommet étant cassé. Téléoconque composée de cinq tours. Côtes foliacées non épineuses au sommet. Présence d'une côte intermédiaire entre les côtes foliacées. Quatre cordons sur la spire et 18 cordons sur le dernier tour. Ouverture subcirculaire à bord columellaire distinct. Sept dents internes. Canal siphonal court, légèrement orienté à gauche. Dimensions: H: 9,5 mm: D: 5.6 mm, H/D: 1,71; HO: 5,3 mm, LO: 2,8 mm; HO/LO: 1,89. Considérations nomenclaturales sur l'espèce: En accord avec VOKES (1971), LE RENARD (1992) considère Murex tricarinatus Lamarck. 1803 comme un synonyme postérieur de Purpura crenulata Rôding, 1798. MERLE (1990) admet pour l'espèce fricarinatus quatre transiants. Ceci m'amène à proposer les combinaisons nomenclaturales suivantes: 1°) Pterynotus (P.) crenulatus trigonus (Rouault, 1850) pour Murex trigonus Rouault, 1850 du Cuisien du Béarn. 2°) P. (P.) crenulatus crenulatus (Rôding. 1798) du Lutétien anglo-franco-belge. 3°) P. (P.) crenulatus tricuspidatus (Deshayes, 1865) pour Murex tricuspidatus Deshayes, 1865, du Lutétien supérieur anglais, de l'Auversien du Bassin de Paris et du Bartonien anglais. 4°) P. (P.) crenulatus brevicauda (Hébert. 1849) pour Murex brevicauda Hébert, 1849, du Lattorfien et de l'Oligocène d'Allemagne. Comparaisons avec des populations conspécifiques: Les deux spécimens, dont celui de la collection Neuville est cité par MAGNE (1940, p.372), sont très comparables aux jeunes individus de P. (P.) crenulatus trigonus, de l'Eocène du Béarn. A la différence des spécimens de Gan qui portent 2 à 3 côtes intermédiaires entre les côtes foliacées, ceux de Biarritz n'en portent qu'une seule. De plus, le sommet des côtes foliacées n'est pas épineux. Ce caractère les distingue de P. (P.) crenulatus crenulatus et de P. (P.) crenulatus 86 Muricidae de l'Eocène tricuspidatus. En revanche, il les rapproche de P. (P.) crenulatus brevicauda du Lattorfien. Comme il s'agit de formes juvéniles, ce rapprochement avec P. (P) crenulatus brevicauda ne peut être certain. En revanche, l'attribution à l'espèce crenulatus ne fait pas de doute. Pterynotus (P.) cf consobrinus (d'Orbigny, 1852) (Figs. 10, 11) Murex consobrinus d'Orbigny, 1852: Prodr.. t. IT, n°1346. Autres références: Murex (Pteropurpura) consobrinus - MAGNE, 1940: J. de Conch., v.84: 370-376, pl. IL, fig. 3, 4. Localité-type: Gaas (Landes). Etage-type: Holotype, Université de Bordeaux Il: Matériel de Biarritz. 1 spécimen incomplet, collection Cossmann n°16453 a (MNHN-LP n°J00002); 1 spécimen très jeune de 2 mm de hauteur, collection Tournouër (IGAL). Description: Coquille à spire élancée, Protoconque lisse, paucispirale composée de 1,5 tour. Côtes foliacées assez planes et non épineuses sur la spire. Une côte intermédiaire bien marquée entre les côtes foliacées. Deux cordons spiraux sur la spire et cinq visibles Jusqu'à la partie pariétale du dernier tour. Dimensions: H (jusqu'à la partie pariétale): 12 mm, D: 9 mm. Comparaisons: Cette espèce ne peut être confondue avec le P. (P.) crenulatus de Biarritz. plus trapu, dont les côtes principales sont plus fortes et dont les cordons spiraux sont plus nombreux (4 sur la spire au lieu de 2). Je compare ces deux spécimens à deux petits spécimens de P. (P.) consobrinus de Gaas provenant de ma collection. La sculpture est analogue puisqu'elle est formée de deux cordons spiraux sur la spire et d'une côte intermédiaire bien marquée (Fig.12). D'autre part, la protoconque lisse et paucispirale est analogue chez les spécimens de Biarritz et de Gaas. Pour ces raisons, Je rapproche les individus de Biarritz de P. (P.) consobrinus, mais la mauvaise qualité du matériel m'empêche d'en être totalement certain. MERLE MERLE Muricidae de l'Eocène Sous-famille TYPHINAE Cossmann, 1903 Genre Siphonochelus Jousseaume, 1880 Sous-genre Laevityphis Cossmann, 1903 Espèce-type: Typhis coronarius Deshayes . 1865 (=Zyphis muticus J. Sowerby, 1834) par désignation originale. Siphonochelus (Laevityphis) biarritzensis nOV. SP. (Figs 3, 13:14, 15, 16) Matériel-type: Holotype, collection Neuville (UBX, n°TYFIPAL 57.23); paratype, collection Cossmann (MMNHN-LP n°J00003). Localité-type: Biarritz (Les Bains). Niveau-type: Marnes à Turbinolia calcar (Priabonien de Biarritz). Etymologie: biarritzensis: de Biarritz. Dimensions de l'holotype: H: 6,2 mm, D: 3,2 mm; HO: 1,4 mm, LO: 0,9 mm; G: 55°. Dimensions du paratype: H: 8,6 mm, D: 38 mm, HO: 1,8 mm; LO: 1,1 mm. G: 30° Diagnose: Petite coquille de 6 mm à 8 mm de hauteur et de 3,5 mm à 3,8 mm de diamètre. Protoconque paucispirale et globuleuse composée de 3/4 de tour. Téléoconque composée de quatre tours étagés. Dernier tour égal à la moitié de la hauteur totale. Suture bien visible et profonde. Sculpture spirale obsolète sur le dernier tour. Scupture axiale composée de quatre stades de développement par tour comprenant chacun une côte large dissociée du tube. Côtes lisses. Tubes s'écartant de l'axe de 37° en moyenne. Ouverture subcirculaire à péristome marqué. Canal siphonal clos. court. Comparaisons: Siphonochelus (L.) muticus, du Cuisien du Bassin de Paris se distingue par sa protoconque composée de 3,5 tours. Ses côtes sont aussi nettement plus fines et son ouverture plus ovale. Sa spire est élancée alors que celle de S. (Z.) biarritzensis est étagée. Siphonochelus (L.) gracilis (Conrad, 1833), du Claibornien des Etats-Unis (Eocène moyen) présente une morphologie voisine de S. (L.) muticus et n'en diffère que par sa spire encore plus élancée. Murex alternata Lea, 1833 est synonyme de cette espèce (PALMER, 1937). Siphonochelus (L.) thagus (Olsson, 1930), de l'Eocène supérieur du Pérou (Talara Formation, Yasila) présente une spire élancée et n'a pas les tours étagés de S (L.) biarritzensis. En revanche, ses côtes sont plus épaisses que celles de S. (L.) muticus et de S. (T.) gracilis. APEX 9(2/3): 83-91, juillet 1994 Position générique. Cette espèce peut se classer dans le sous-genre S.(Laevityphis) en raison de sa petite taille et de ses côtes lisses, nettement dissociées des tubes. Sous-genre 7rubatsa Dall, 1889 Espèce-type: Typhis (Trubatsa) longicornis D'Orbigny, 1850, par désignation originale. Siphonochelus (Trubatsa) evainae NOV. SP. (Figs. 4, 17, 18, 19, 20)) Matériel: Holotype: collection Neuville (UBX, n TYFIPAL 57.24), paratype n 67.04.03, collection Castex (BIA); autres spécimens: 2 Spm: collection Cossmann (MNHN-LP n J00004). Localité-type: Biarritz, Côte des Basques. Niveau-type: Marnes à Zurbinolia calcar (Priabonien de Biarritz). Etymologie: evainae: d'Evaine, ma fille nouvelle née. Dimensions de l'holotype: H: 9,6 mm; D: 5,6 mn 0:20 mm; "207 1,3 "ons ‘CG:063". Dimensions du paratype: H: 7,4 mm, D: 4,1 mm: HO: 1,7 mm; LO: 1,2 mm. G: 65. Dimensions des specimens de la collection Cossmann: H: 7,6 mm, 6,9 mm; D: 3,9 mm, 39m 40" 17 on, 1/4/mm; LO:"12 mm, 0,9 mm; G: 56’, 53. Diagnose: Coquille trapue de 7 à 9 mm de haut et de 4 à 5 mm de diamètre. Protoconque conique, multispirale, composée de 3,5 tours. Téléoconque composée de quatre tours. Dernier tour égal au 2/3 de la hauteur totale. Suture visible. Sculpture spirale ornée de un ou deux cordons obsolètes sur la partie centrale du dernier tour. Sculpture axiale composée de quatre côtes totalement associées aux tubes. Présence d'un cal partant de la base du tube, s'appliquant à la base de la côte du tour précédent et recouvrant une partie de la suture. Tube anal s'écartant de l'axe de 59° en moyenne. Ouverture ovale à péristome marqué. Canal siphonal clos, large. Comparaisons: Les deux spécimens de la collection Cossmann se trouvaient mélangés avec un spécimen de S. (Laevityphis) biarritzensis décrit ci-dessus. L'ensemble avait originellement été identifié comme 7Zyphis pungens Solander, puis Cossmann a remplacé, sur l'étiquette, ce nom par celui de Typhis newtoni. Dans un autre tube portant 87 APEX 9(2/3): 83-91, juillet 1994 originellement l'étiquette 7: newioni, se trouvaient six fossiles de Barton parmi lesquels on peut reconnaître cinq Zyphis (T.) pungens et un $S. (Trubatsa) parisiensis. Dans ce tube, Cossmann a écrit :"spire beaucoup plus courte que ZT. fistulosus". Cossmann fait très probablement allusion à S. (T.) parisiensis [car S. (Siphonochelus) fistulosus (Brocchi, 1814) n'existe pas à Barton]. T°: newtoni paraît être un nom manuscrit. À mon avis, il désignait plutôt les individus de Barton, que ceux de Biarritz. Siphonochelus (T.) parisiensis (d'Orbigny, 1850), de l'Eocène d'Europe diffère de S. (T.) evainae par sa grande taille (de 16 à 23 mm de hauteur), par sa spire plus élancée, par sa protoconque globuleuse de 1,5 à 2 tours et par ses côtes ne portant pas de cal recouvrant la suture du tour précédent. Siphonochelus (T.) gaasensis (Tounouër in Benoist, 1880), du Stampien d'Aquitaine diffère par sa taille dépassant 26 mm (pour un individu de 5 à 6 tours) et par des sillons spiraux sur la partie externe des côtes. Un cal partant du sommet des côtes et débordant sur la suture est un caractère commun avec S.(1.) evainae (fig.5). On ne peut confondre S. (T.) evainae avec le jeune de S. (T°) gaasensis qui porte déjà des sillons spiraux marqués et dont la spire est bien plus courte. Siphonochelus (T.) hortensis (Oppenheim, 1900), du Priabonien d'Italie est aussi de grande taille. Il porte comme S.(T) gaasensis des sillons spiraux sur la partie externe des côtes. Le sommet des côtes ne porte pas de cal. Siphonochelus (T.) nigeriensis Arua, 1981, de l'Ameki Formation (Eocène moyen du Niger), ressemble à S. (T.) evainae par sa taille (de 6 à 9,5 mm) et par la présence au sommet des côtes d'un cal débordant sur la suture du tour précédent et par ses côtes dépourvues de crénelures. Il se distingue par sa spire plus étagée, par l'absence de cordons spiraux et par Sa protoconque de 1,5 tours. Considérations phylogénétiques: Sipho- nochelus parisiensis, S. hortensis et S. gaasensis semblent constituer une même lignée de Trubatsa paléogènes. Elles ont en commun une grande taille (20 à 30 mm), et le port de crénelures sur les côtes. La protoconque, observée chez S. (T.) parisiensis est de type paucispiral. L'évolution de S. (T.) parisiensis conduit très probablement à S. (T.) hortensis au Priabonien. La sculpture spirale développée chez S. (T.) hortensis et chez S. (T.) gaasensis constitue à mon sens une synapomorphie, témoin d'un ancêtre commun. Siphonochelus (T.) evainae et S (T.) nige- riensis, par leur petite taille (6 à 9 mm) et par 88 Murnicidae de l'Eocène l'absence de crénelure sur les côtes, semblent appartenir un groupe différent de celui de S. (T.) parisiensis. Le cal porté sur les côtes, qu'on retrouve chez $. (T.) gaasensis et chez S. (T.) nigeriensis résulterait d'un parallèlisme. Position générique: L'ensemble de ces espèces porte des côtes totalement associées aux tubes. Pour cette raison, elles sont très voisines de S. (T.) longicornis, espèce-type du sous-genre Trubatsa Dall, 1889. Remerciements: Je tiens à exprimer ma gratitude à L. Rousselle et B. Cahuzac de l'Université de Bordeaux I, Talence, à J. Harambillet du Muséum d'Histoire naturelle de Biarritz et à L.Gauthier de l'IGAL, qui m'ont permis d'emprunter les si rares Muricidae récoltés à Biarritz. Mes remerciements vont aussi à R. Houart et à P. Bouchet qui ont relu ce manuscrit. Les clichés photographiques sont de L. Merlette et les clichés au MEB sont de C. Chantograin. La planche photographique a été montée par F. Pilard. MERLE MERLE BIBLIOGRAPHIE ARUA, IL, 1981. First record of typhine gastropods from the Eocene of southern Nigeria. Geologie en Mijnbouw, 60 (2): 277-288, fig. 1-4. BOUILLE (de), R., 1876. Paléontologie de Biarritz et de quelques autres localités des Basses-Pyrénées, Veronese édit, Pau, 71 pp, 3 pis. BRANDER, G., 1766. Fossilia Hantoniensia collecta, et in Musaeo Britannico deposita. London, vi, 43 p.,9 pl. HOUART, R., 1991. Mollusca Gastropoda: The Typhinae (Muricidae) from the New- Caledonian region with description of five new species. /n A. Crosnier & P. Bouchet édit. Résultats des campagnes de Musorstom, vol.7, Mém Mus. natn. Hist. nat., Paris. A, 150: 223- 241. HOUART, R., 1992. The genus Chicoreus and related genera (Gastropoda: Muricidae) in the Indo-West Pacific. Mém. Mus. natn. Hist. nat. Paris, 154: 1-188. KOENEN, (von) A. 1889. Das norddeutsche Unter-Oligocän und seine Molluskenfauna. Abh. geol. Specialkarte preuss. und Thür. Staaten, Berlin, 10 (1): 1-280, pl.24-39. LAMARCK, J.B.. 1803. Mémoires sur les fossiles des environs de Paris (suite 1). Annales du Muséum d'Histoire naturelle. Paris, t. II, 1803: 57-64; 163-169; 217-227: 315-321: 385-391. LE RENARD, J., 1992. Note de nomenclature sur quelques mollusques caractéristiques de l'Eocène du Bassin de Paris, Cossmanniana, Paris, 1 (2-4): 1-14. MAGNE, A., 1940. Les Pteropurpura tertiaires du Bassin d'Aquitaine. J. de Conch., Paris, 84: 370-376. MARTINI, 1971. Standard Tertiary and Quaternary calcareous nannoplancplon zonation: Proc.ll Planktonic Conf. Rome, Edizioni Technoscienza, Roma: 739-783. Muricidae de l'Eocène APEX 9(2/3): 83-91, juillet 1994 MATHELIN, J.C., 1988. Le Paléogène des falaises de Biarritz Révision biostratigraphique. Paléoenvironnement et diapirisme. Thèse de 3ème cycle, Université de Paris VI, Paris: 160 pp. 20 pls, figs. MERLE, D. 1990. Révision des Muricidae du Cuisien de Gan et de Bos d'Arros (Bassin d'Aquitaine, France). Bull. Mus. natn. Hist. nat., Paris, 4è sér., 11, 1989, C (3): 145-185, 4 pis. 7 figs, 1 tab. NOLF, D. 1988. Les otolithes de téléostéens d'Aquitaine et leur intérêt stratigraphique. Mém. Acad. roy. de Belgique, Bruxelles, Coll. in 40, sér.3, 19: 1-147, 14 pls, 9 figs, 3 tabls. OLSSON, A., 1930. Contributions to the Tertiary paleontology of northern Peru. Bull. Amer. Paleont., 17 (62): 1-96, pl. 1-12. OPPENHEIM, P, 1900. Die Priabonaschichten und ihre fauna, Paleontographica, 47: 1-348, 21 pls. PALMER, K.E.V., 1937. The Claibornian Scaphopoda, Gastropoda and dibranchiate Cephalopoda of the southern United States, Bull. Amer. Paleont., 17 (62), part I: 1-548, 90 pis. RADWIN, G.E. & A. D'ATTILIO. 1976, Murex Shélls of the World. Stanford édit, Stanford, 283 pp, 32 pis. RÔDING, P.F., 1798. Museum Boltenianum sive Catalogus cimeliorum e tribus regnis naturae quae olim collegrat Joa. Fried Bolten, M.D.p.d. Hamburg. vi, 199 p. VOKES, EH., 1971. Catalogue of the genus Murex Linné (Mollusca, Gastropoda); Muricinae, Ocenebrinae, Bull. amer. Paleont.. 61 (268), 141 pp. VOKES.E.H., 1990. Cenozoic Muricidae of the western Atlantic région. Part VIII. Murex (s.s.), Haustellum, Chicoreus and Hexaplex; additions and corrections. Zulane stud. geol. paleont. Tulane, 23 (1-3): 1-96, 12 pls. 89 APEX 9(2/3): 83-91, juillet 1994 Muricidae de l'Eocène MERLE Figures 7-20 (ci-contre). 7 - Chicoreus (?) subfiligrana (Tournouër in de Bouillé, 1876), néotype, collection Neuville (UBX, n TYFIPAL 572.1), H: 15 mm. 8 - C. (?) subfiligrana (Tournouër in de Bouillé, 1876), collection Cossmann n°16453a (MNAN-LP n°J00001), détail de la microsculpture du labre, X 20. 9 - Pterynotus (Pterynotus) crenulatus (Rôding, 1798), collection Neuville (UBX, n° TYFIPAL 57.2.2), H: 9,55 mm. 10 - Pterynotus (Pterynotus) cf. consobrinus (d'Orbigny, 1852), collection Tournouër (IGAL), très jeune individu montrant une protoconque paucispirale, H: 3,3 mm. 11- P. (P.) cf. consobrinus (d'Orbigny, 1852), collection Cossmann n°16453a (MNHN-LP n'J00002), individu plus âgé, H: 11,5mm. 12 - P. (P.) consobrinus (d'Orbigny, 1852), collection Merle, spécimen de Gaas (Stampien, Landes), H: 18,0 mm. 13 - Siphonochelus (Laevityphis) biarritzensis nov. sp., collection Neuville (UBX, n TYFIPAL 57.2.3), holotype, vue montrant la protoconque et les premiers tours, H: 6,20 mm 14,15 -S. (L.) biarritzensis nov. sp., collection Neuville (UBX, n° TYFIPAL 57.2.3), holotype, H: 6,2 mm. 16 - S. (L.) biarritzensis nov sp. collection Cossmann n°14276 (MNHN-LP n°J00003), paratype, H: 8,6 mm. 17,18 - S. (Trubatsa) evainae nov. sp., collection Neuville (UBX, n° TYFIPAL 57.2.4), holotype, H: 9,65 mm. 19 - S. (T.) evainae nov. sp., collection Castex (BIA), paratype, H: 7,45 mm. .20 - S (T.) evainae nov. sp., collection Cossmann n°14276a (MNHN-LP n°J00004), protoconque X 45. 90 MERLE Muricidae de l'Eocène APEX 9(2/3): 83-91, juillet 1994 91 CAPENSIS SEA SHELLS SPECIMEN SHELLS OF THE SOUTHERN AFRICAN REGION Bruno de Bruüin P.O. 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For the layout authors will refer to a recently published article and take the Editorial Board remarks into account. Off print: members or subscribers. Thirty off prints representing a maximum of 200 pages, will be sent free of charge to the authors. More copies can be ordered when the proof sheets are returned. Those as well as all postcharges will be billed to the author. Non membeïs: Off prints are available to the authors. In this case there is an obligation to order at least 50 copies when the proof sheets are retumed. They will be available at cost. Manuscripts have to be sent to M. R. Houart, St Jobsstraat, 8, 3400 Lan- den (Ezemaal), Belgium. MCZ LIBRARY JAN 18 1995 HARVARD UNIVMERSITY Société Belge de Malacologie association sans but lucratif DECEMBRE 1994 SOMMAIRE J. Vidal A review of the genus Fulvia Gray, 1853 (Mollusca, Cardiidae) R. Houart The Muricidae (Gastropoda) from Madeira with the A. D. Abreu description of a new species of Ocenebra (Ocinebrina) (Muricidae: Ocenebrinae) B. Tursch The scale of sympatry in the genus Oliva (Gastropoda, Olividae) A. Panigrahi Noradrenaline and adrenaline in the cerebral ganglia S.K. Raut of the giant African land snaïl Achatina fulica Bowdich. Périodique trimestriel Bureau de dépôt 1180 Bruxelles 18. Comité d'édition : Editeur responsable : R. Duchamps Dr. Y. Finet L. Germain R, Houart Dr. CI. Massin Prof, B. Tursch Dr. J. Van Goethem Les articles et textes présentés dans cette revue réflètent l'opinion personnelle de leur(s) auteur(s), et non pas nécessairement celle de la Société ou de l'éditeur responsable. Tous droits de reproduction, de traduction et d'adaptation des articles publiés dans ce bulletin, réservés pour tous pays. All rights of reproduction are reserved without the written permission of the board. Belgique - Belgium [avec le service des bulletins) Membre effectif nb NanniREr Membre éldiant 2 500 BEF (sans le service des bulletins) Personne appartenant à la famille d'un membre effectif et ayant la même résidence RAR Te 400 BEF Versements à effectuer au C.C.P. n° 0000974225:54 de la Société Belge de Malacologie c/o M. J. Buyle, Av. M. Maeterlinck, 56, 1030 Bruxelles. Etranger - Foreign Abonnement aux revues APEX & ARION Subscription to APEX & ARION Le SERA PM PP 1400 BEF Versement à effectuer par mandat postal international ou ar chèque bancaire en francs belges uniquement. ee s international money order, or by bank check in Belgian Francs only. au nom de at name of : M. J. Buyle Av. Maurice Maeterlinck, 56, bte 8 B-1030 Bruxelles. CONSEIL D'ADMINISTRATION DE LA SOCIETE BELGE DE MALACOLOGIE ° Président : MR. Duchamps, Av. Mozart, 52, 1 190 Bruxelles © : (02) 344.15.47 + Vice-présidents Dr. Y. Finet, 16 Chemin des Clochettes, CH-1206, Genève (Suisse) M.R. Houart, St. Jobsstraat, 8, 3400 Landen [Ezemaal) ® : A1-2246.77.95 © : (01é) 78.86.16 e Secrétaire Mme J. Masson, Rue du Merlo, 10, 1180 Bruxelles © : (02) 376.62.25 e Trésorier M. J. Buyle, Av. Maeterlinck, 56, bte 8, 1030 Bruxelles © : (02) 216.68.21 e Bibliothécaire : Mme ML Buyle, Av. Maeterlinck, 56, bte 8, 1030 Bruxelles © : (02) 216.68.21 + Relations publiques e Administrateurs M.G. Geeraerts, Dorpsstraat, 125, 3078 Meerbeek Mme ML. Bresson, Place Guy d'Arezzo, 7, 1060 Bruxelles M. L. Germain, Rue de Linthout, 77, 1040 Bruxelles : Mme À. Langleit, Av. Cicéron, 27, bte 92, 1 140 Bruxelles M. C. Van Osselaer, Chée de Waterloo, 512, 1060 Bruxelles M. E. Waiengnier, Rue C. Wolles, 42, 1030 Bruxelles @ : (02]757.07.47 ® : (02) 343.62.38 ® : (02) 34.80.11 ® : (02) 720.41.61 ® : (02) 347.36.84 © : (02) 241.51.80 VIDAL The genus Fulvia APEX 9(4): 93-118, déc. 1994. A review of the genus Fulvia Gray, 1853 (Mollusca, Cardiidae). Jacques VIDAL Attaché au Museum National d'Histoire Naturelle de Paris, Laboratoire de Malacologie, 55 rue Buffon, 75005 Paris, France. KEY WORDS. Mollusca, Bivalvia, Cardiidae, Fulvia, review. ABSTRACT. The Cardiid genus Fulvia is revised based on examination of type material and over 900 lots in relevant museums. Characters of shell sculpture are regarded as taxonomically more reliable than shape and colour, hitherto the base of specific taxonomy in the genus. It is established that Fw/via shares with Vepricardium numerous characters, including the presence of ocular organs on siphonal tentacles, and is included here in the tribe VEPRICARDIINI. Zaevifulvia subgen. nov. [Type species: F undatopicta Pilsbry, 1904] is segregated from Fulvia s.s., based on the lack of periostracal insertions on the ribs. Sixteen Indo-Pacific species of recent Fulvia are recognized, of which six are new. The name F. papyracea, hitherto used for a common Indo-West Pacific species 1s shown to represent a rare and restricted species. The name 7° fragilis (Forsskal in Niebuhr, 1775) is to be used for most usages of F. papyracea of authors. RESUME. Révision du genre Fulvia, famille CARDIIDAE, basée sur l'examen du matériel type et de plus de 900 lots de différents muséums. Les caractères de l'ornementation de la coquille sont considérés comme taxonomiquement plus fiables que la forme générale ou la couleur, sur lesquelles, jusqu'à present, les séparations spécifiques sont basées. Il est constaté que les Fulvia partagent avec les Vepricardium de nombreux caractères, en particulier la présence d'organes oculaires à l'extrémité des tentacules siphonaux, et sont donc placées ici dans la tribu des VEPRICARDIINI. On sépare de Fulvia s.s. un sous genre ZLaevifulvia [Espèce type F. undatopicta Pilsbry, 1904], séparation basée chez ce dernier sur l'absence d'insertions périostracales sur les côtes. Il est reconnu seize espèces récentes de Fulvia dans l'Indo-Pacifique, dont six sont nouvelles. On montre que le nom Fulvia papyracea, utilisé jusqu'ici pour désigner une espèce commune de l'Ouest de l'Indo-Pacifique, représente en réalité une espèce rare et d'extension limitée. Le nom F. fragilis (Forsskal in Niebuhr, 1775) doit êre utilisé, la plupart du temps, à la place de F. papyracea utilisé par les auteurs. INTRODUCTION very polymorphic and, although there are twenty five nominal species, the most recent This general study originates from an authors have been prudently inclined to attempt to identify some species of the genus Fulvia found around New Caledonia. For these identifications I both consulted literature and compared identified specimens in museums. The result was very confusing and I remained perplexed about the true identity of many forms at the specific level, the generic level always being easy to determine. The literature on this subject is of little help because the descriptions are always too succint, the effective comparisons quasi-absent, and opinions often vary from an author to another. In the museums the identifications are also often rather disconcerting. I then presumed that the reason of this confusion could be the inconsistency of the characters in the species of this genus. As a matter of fact, they are generally reputed to be consider only a small number of species (about four or five) in this group, and FISCHER-PIETTE (1977) admits the existence of only two species of recent Fulvia! MATERIAL AND METHODS In the literature, the criteria for description and species separation are scarce and rather vague and subjective. They mainly concern the general shape of the shells, the colours and the strength of the ribs, characters which, at first sight, look variable in a same species and may be common to several. The review of the genus Fulvia consisted of examining systematically in each specimen the largest number of characters, beside shape and colours, in order to find 98 APEX 9(4): 93-118 , déc. 1994. characters with importance. significant taxonomic The type specimens are very rarely cited and can often be supposed as not having been taken into account by the authors. Almost all the extant types specimens have been examined. For this study about 905 lots have been examined and ,in almost half this number, at least one, often several specimens, have been measured. In this material, about 588 lots originate from ORSTOM's dredging program conducted by B. Richer de Forges in the peripheral lagoon of New Caledonia and in the shallows of Chesterfield Islands, from 1984 to 1992. This material is in the MNEN. This material comes from the following museums: AMS: Australian Museum - Sydney. ANSP: Academy of Natural Sciences - Philadelphia. BM(NH): British Museum (Natural History) - London. IRSNB: Institut Royal des Sciences Naturelles de Belgique - Brussels. MANG: Museum d'Histoire Naturelle de Genève - Geneva. MNAN: Museum National d'Histoire Naturelle - Paris. NM: Natal Museum - Pietermaritzburg. QM: Queensland Museum - Brisbane. WAM: Western Australian Museum - Perth. ZMA: Zoologisch Museum - Amsterdam. ZMUC: Zoologisk Museum - Copenhagen. DEFINITIONS In order to simplify the descriptions, some expressions are used, defined as follows: A = Angle A: In the right valve, the angle between two lines joining the main cardinal tooth to the laterals. AT= Anterior Third of the shell in a radial division, excluding the lunule sensu stricto and the sublunule (pl. 1, fig. 1). When the sublunule is long this "third" 1s reduced. D = Ratio D: In the left valve, the ratio between two distances: the distance from the extremity of the posterior lateral to the tip of the umbo and the distance from this point to the middle of the anterior lateral. 94 The genus Fulvia VIDAL Granulation, granules (pl. 1, figs. 4 and 8; pl. 2, fig. 10; pl. 3, fig. 7): Minute tubercles very often present on the external surface of the shell. They can be concentrically aligned, sometimes on long distances (several ribs and interstices). They are usually situated on the anterior half of the shell but can be present everywhere. These small tubercles have an hexagonal symmetry in most species. IMR = Internal Marginal Ribbing: The reverse of the external ribbing extending onto the internal side of the shell at the margin and in à certain distance from it. This "ribbing" is always well discernible and the comparison of the width between ribs and interstices (which is an important taxonomic character) is always easier to observe in the IMR. Intersticial riblets: In the posterior third of the shell, more or less numerous small ribs. often irregular, can appear in the interstices between the main ribs (pl. 1, fig. 8). Last flat: On the external posterior part of the left valve, the area, often relatively wide, between the last rib and the dorsal margin, bearing the posterior lateral on its inner side. This zone is generally flat or slightly concave (pl. 1, fig. 8). Last fold: On the external posterior part of the right valve, the area, often relatively wide. between the last rib and the dorsal margin, bearing the posterior lateral on its inner side. This zone is generally convex. When the shell is closed, the last fold partially overlaps the last flat (pl. 1, fig. 8). Lunule: It is not a true lunule (present only in FRAGINAE) but the most anterior part of the shell, always without ribs and periostracal insertions, often separated from the rest of the shell by the perilunular furrow (see this word). The smooth zone can extend farther backwards (sublunule), but, in the descriptions, the lunule sensu stricto Will be limited at the level of the extremity of the basement of the anterior teeth, even when a furrow is not present, and its "length" along the margin will be measured from this point to the extremity of the umbo (pl. L'fig ol) Lunular heart: The anterior dorsal margin of the shell just beside the umbo, exteriorly arising and spreading on both sides, forming like a "heart" (pl. 1, figs.2 and 3). Ocular organs: Situated on the tip of tentacles bordering the two siphonal apertures: they are generally darker. MT = Medial Third in a radial division (pl. 1, fig. 1). VIDAL Perilunular furrow: The lunule sensu stricto is sometimes limited by a more or less large and deep furrow, a slight and open groove corresponding internally to the ridge supporting the anterior tooth. Periostracal insertions: In most species, periostracal lamellae are perpendicularly implanted in the shell on or along the ribs. These insertions consist of a succession of elementary components formed with short concentric lamellae and longer radial ones, forming "nodes" at their intersection, with a simple (pl. 1 fig. 5) or complex arrangement (pl. 1, fig. 7) according to the different species. Joined end to end these basic elements form finely undulating or zigzagging palisades on or along the ribs. These fragile palisades often disappear when the shell dries, but the traces of the insertions are always visible on the surface of the shell. Sometimes the nodes become calcified and form tubercles or spines on the ribs (pl. 1, fig. 6). Posterior groove: Most species have a distinct more or less pronounced radial groove in the middle of the posterior third of the shell forming a notch in the margin and corresponding interiorly to a raised radial oblique ridge. PT = Posterior Third of the shell in a radial division, often divided into two parts by the axis of the posterior groove (see above): PT1: the anterior part, close to the MT and PT2: the posterior part, ending beside the last flat or the last fold. Often, these two parts of the PT have distinct ribs (pl. 1, fig. 1). Sublunule: In some species the lunule sensu stricto is followed by a more or less developped area without ribs nor IMR, but with some possible periostracal insertions. This area is sometimes separated from the lunule by a “perilunular furrow" (see this word). The length of the sublunule along the margin will be estimated by comparison with the length of the lunule (pl. 1, fig. 1). TAXONOMY OF THE GENUS FULVTA GRAY, 1853 The taxon Fulvia was created as a monotypic genus of the family "CARDIADAE" by GRAY (1853: 40), with a type species: Fulvia aperta Bruguière, from the Indo-Pacific. Also Gray cites another genus: Papyridea Swainson, 1840, for which he had designated the type species (1847: 185): Papyridea soleniforme. from the American Atlantic. In spite of these The genus Fulvia APEX 9(4): 93-118, déc. 1994. clear, logical and unambiguous definitions, and in spite of the fact that these two species, the gaping excepted, are extremely different, the majority of the subsequent authors did not follow Gray and, for a long time, Fulvia was erroncously considered as a subgenus of Papyridea. In 1951, KEEN justly separated both taxa: Papyridea was placed, as a genus, into the subfamily TRACHYCARDIINAE and Fulvia, as a subgenus of Zaevicardium, into the subfamily LAEVICARDIINAE Keen, 1951. [The erection of LAEVICARDIINAE by KEEN (1936: 367) does not comply with ICZN art.13,(a),(1), and is nomenclaturally unavail- able]. KAFANOV & Popov (1977: 310) consider Fulvia and Laevicardium as two genera of the tribe LAEVICARDIINI, subfamily CARDI- INAE. Finally, SCHNEIDER (1992: 146), again places Fulvia and Laevicardium into the subfamily LAE VICARDIINAE. Nevertheless, several authors, for example WILSON & STEVENSON (1977: 52), have noted that "the species of Fulvia do not agree with the diagnosis of Laevicardium" for many reasons. This study confirms the veracity of this opinion and it can be remarked also that the species of Fulvia have numerous affinities with the species of V’epricardium Iredale, 1929. These similari- ues are as follows: 1)- Shape comparable, with length more or less equivalent to height, often equilateral with rounded margins and no or limited posterior trunçation. 2)- Hinge similar with, in particular, a large angle A. 3)- Several other features are comparable in both groups: large lunule with a lunular heart, well developped last fold overlapping a last flat te: 4)- Young specimens of Vepricardium are also radially divided into three parts, with a smoother median one, and can look very much like Fulvia. 5)- Young specimens of some lepricardium bear also "periostracal insertions", which turn into calcified palisades or spines in the adults, with equivalent detailed structures. 6)- In some species of Fulvia, the nodes of the periostracal insertions change into calcified spines, like those of V’epricardium. 7)- The external surface of the shell, in both groups, bears numerous minute granules, often concentrically aligned. 8)-The last but maybe the most significant element of similarity: the presence in both groups of ocular organs (see above). As far as I know, this character does not exist in the other 95 APEX 9(4): 93-118 , déc. 1994. CARDIIDAE, except in the two species of the European genus Cerastoderma. Because of these numerous affinities, I think that Fulvia and Vepricardium must be placed into a same suprageneric group and logically into the tribe VEPRICARDIINI subfamily CARDIINAE (see later). As far as Laevicardium s.s. is concerned, the species of this genus remain similar to Acrosterigma Dall, 1900, as far as certain characters are concerned, and should be placed into the tribe TRACHYCARDIINI Stewart, 1930. Nevertheless, it 1is true that some characters of Laevicardium are present in some evolved species of Fulvia (particularly in the subgenus Laevifulvia). 1 think this is due to a phenomenon of convergence. In brief the taxonomy of Fulvia is considered here as follows. Family CARDIIDAE Lamarck, 1809, Subfamily CARDIINAE Lamarck, 1809, Tribe VEPRICARDIINI Kafanov & Starobogatov (in Kafanov & Popov, 1977). Genus Fulvia Gray, 1853. Type species: Fulvia aperta (Bruguière). General characters: Shell often thin and fragile, more or less translucent, but sometimes moderately thick and opaque. Dimensions variable from small (10 mm) to large (100 mm) in length. Shape: Rarely noticeably elongated (length generaly nearly equivalent to height), varying from subcircular and equilateral to inequilateral, with the posterior side more or less transversally to obliquely expanded. Sometimes slightly truncated on the posterior margin, the anterior and ventral margins being always rounded. Variably but rather moderately inflated, with a regular general concentric curvature of the shell (no or exceptional medio- posterior accentuation of the curvature nor rostration). Often, presence of a "posterior groove" (see above). The shells are generally practically closed in the posterior margin, only two species noticeably gaping. Other external features: (See above for certain definitions). Umbo generally prosogyrate without exaggeration. "Lunule" generally broad, with a variably developed "lunular heart”. "Sublunule" from nonexistant to extremely well developped. Presence of a "last flat" and a "last fold", both of variable extention and profile. 96 The genus Fulvia Colours: Exterior of the shells of variable colour: white, beige or yellowish to orange, brown purple, with often, mainly in the young shells, more or less developed darker irregular concentric stripes or zigzags dark yellow, brown to pink or purple. The tip of the umbo is often coloured purple, and also sometimes the sublunule and part of the posterior third. Interior generally white, beige to orange often with more or less developed irregular concentric stripes corresponding to those in the exterior and also one or two thin radial lines crossing the above stripes. Posterior zone often coloured purple. Sometimes the interior is almost entirely coloured pink, orange, brown or purple, and sometimes only in the umbonal cavity. Sculpture: a)- Number of ribs: This number is extremely variable between the species, and within some species according to populations and individuals. It ranges between 33 and 68. b)- Ornamentation: The young shells (and the earliest formed part of the adult shells) are smooth, except for some large ribs in the PTI. When the shell grows, ribs and interstices appear progressively in the other parts. They are always the strongest and the largest in the PT and the weakest in the MT, where they can sometimes be hardly perceptible. Nevertheless the internal marginal features (IMR, which have a significant taxonomic interest) are obvious everywhere.In the PT some irregular interticial riblets can appear in the interstices. In some species the ribbing and the IMR disappear in the most anterior part of the shell, behind the lunule (sublunule more or less developed). The profile of the ribs and interstices varies according to the species and to the position on the shell. The ribs are often flat, with rounded edges, but can be triangular or rounded. They are generally smooth, without any ornaments, but in some rare species tubercles can appear. The interstices are generally flat and of very variable width with regard to the ribs. They are generally smooth but they can be striated in the PIE In most species the ribs bear "periostracal insertions". They are generally located on the posterior limit of the ribs, but they can be placed in à variable position from this limit to the middle of the rib. When they exist, these insertions are present on all the ribs of the shell. They sometimes remain also in the sublunule, although the ribs and IMR have disappeared. VIDAL VIDAL The genus Fulvia APEX 9(4): 93-118, déc. 1994. In some species, one, sometimes two, periostracal insertions develop on the last flat or the last fold. The external surface of the shell is almost always ornamented with minute tubercular granules, often not perceptible with the naked eyes. These microscopic pustules mostly develop in the anterior part of the shell, but they can be present everywhere. They are often concentrically aligned. They can be limited to the interstices, or to the ribs, or to be placed on both. Hinge: Hinge plate narrow, thin, moderately curved (angle A varying from 150° to 120°), occasionally angled in its anterior part, rarely excessively assymmetric (ratio D generally varying from 10 to 1.2, but sometimes reaching more extreme values: 0.8 to 2.0). Cardinals generally unequal and separated in the left valve, approximately equal and touching at their base in the right valve. Posterior lateral in the left valve very low. elongated, blade-like, often hardly separated from the ventral margin, in the right valve this tooth is higher and triangular. The main anterior lateral in the right valve is thin, pointed and triangular, it 1s prolonged towards the umbonal area below the hinge plate, forming a very regularly arced well delineated ridge, flattened against the internal surface of the shell. In the left valve the anterior lateral is roughly triangular and is prolonged by a raised narrow ridge, more or less elevated, separated from the dorsal margin by a deep elongated cleft which reaches the cardinal area. The ventral border of the above internal ridges limiting the anterior laterals corresponds exactly with the limits of the lunule. Soft parts: The softs parts of ten species (of sixteen) have been observed, plus one literature record. All have two siphonal apertures bordered by tentacles, and some of these tentacles always bear dark ocular organs on their tip. The genus Fulvia will here be separated into two subgenera: Subgenus Fulvia Type species: Fu/via aperta (Bruguière). Diagnosis: See general diagnosis of the genus Fulvia, periostracal insertions present. Subgenus Laevifulvia new subgen. Type species: Fulvia undatopicta (Pilsbry). Diagnosis: See general diagnosis of the genus Fulvia, but no periostracal insertions. SPECIES LIST AND IDENTIFICATION KEY A)- Fulvia (Fulvia): Periostracal insertions. la: Presence of a rather long sublunule 2 1b: Very short or no sublunule 3 2a: Sublunule as long as the lunule: Fulvia (F.) dulcis (Deshayes, 1863): Small, equilateral, large lunular heart, irregularly coloured, ribs in AT and MT hardly perceptible, ribs strong in PT. 2b: Sublunule shorter than the lunule: Fulvia (F) scalata sp. nov: Small, equilateral,, very large lunular heart, uniformly coloured, ribs strong in PT and AT, perceptible in MT. Granules in AT and PT aligned like rungs of ladder in the interstices. Triangular nymph. 3a: In MT, ribs and interstices of unequal width 4 3b: In MT, ribs and interst. of about the same width 6 4a: In MT and part of AT, ribs very much wider than interstices: Fulvia (F.) mutica (Reeve, 1844): Large to v. large, uniformly coloured, ribs in MT very weak but IMR well marked. Periostr. insertions strong, always placed in the middle of the ribs. No granulation in adult specimens. 4b: In MT and part of AT, ribs very much narrower than interstices 5 Sa: Shell equilateral, not gaping: Fulvia (F.) papyracea (Bruguière, 1789): Large, ribbing very weak in MT, numerous granules in AT, hinge regularly curved, ratio D about 1.2. Sb: Shell inequilateral, gaping: Fulvia (F) aperta (Bruguière, 1789): Large, lunule large, ribbing very weak in MT, granulation in AT, hinge anteriorly angled, ratio D about 1.0 or less. 6a: Ribs and interstices not flat (rounded or triangular) fl 6b: Ribs and interstices flat 9 7a: Shell fragile, gaping, dark colour, discretely ribbed: Fulvia (F) natalensis (Krauss, 1848): Medium to large, inequilateral, lunule very small, ribbing regular on the whole shell, no granulation, hinge not angled, ratio D very high (1.4 to 1.9). 7b: Shell solid, closed, light colour, markedly ribbed 8 97 APEX 9(4): 93-118 , déc. 1994, 8a: Rüibs flatly rounded to asymmetrically triangular in MT, rather rounded in AT: Fulvia (F) fragilis (Forsskal, 1775): Medium to large, last fold high and regularly rounded, 41 ribs (34-52), calcareous spines in PT2, granules only in young shells very anteriorly. 8b: Ribs symmetrically triangular in MT, rather triangular in AT: Fulvia (F.) tenuicostata (Lamarck, 1819): Medium to large, last fold very large with two parts (round and flat), 51 ribs (44-59), No calcareous spines in PT2. Granules rather rare in AT. 9a: Elongated calcified tubercles in PT: Fulvia (F.) boholensis sp. nov.: Medium, inequilateral, generally orange, IMR in MT often almost indistinct. 9b:No calcified tubercles in PT 10 10a: Shell small, roundly angled between MT and PT: Fulvia (F.) fragiformis sp. nov. 10b: Shell medium, with regular concentrical curvature: Fulvia (F.) australis (Sowerby, 1834): More often than not obliquely inequilateral, in MT and AT: identical weak flat regular ribbing, strongly constrasting with the one in PT, much wider and stronger and rather irregular. B)- Fulvia (Laevifulvia): No periostracal insertions: la: Shell elongated when adult, higher than long 2 1b: Shell not elongated. 3 2a: Sublunule about as long as lunule: Fulvia (L.) undatopicta (Pilsbry, 1904): Small, equilateral, often 4 ‘"crushed" spots, numerous aligned granules in AT. 2b: Sublunule much longer than lunule: Fulvia (L.) lineonotata sp. nov: Small, inequilateral, coloured by small aligned triangles, thin granules in AT. 3a: Shell longer than high: Fulvia (L.) hungerfordi (Sow, 1901): Small, asymmetric, "polygonal" aspect, granules on the whole shell, ribbing and IMR in MT variable. 3b: Shell as long as high. À 4a: Shell small, very inequilateral: Fulvia (L.) prashadi sp. nov.: Ridged and strongly separated sharpened PT, No ribbing nor IMR in MT, strong in PT, very rare or no granules. 4b: Shell medium, almost equilateral: 98 The genus Fulvia Fulvia (L.) ballieni sp. nov.: Regular weak ribbing and IMR in MT and AT, larger ribbing in PT. Granules on the umbo. SPECIES DESCRIPTIONS. Fulvia (Fulvia) aperta (Bruguière, 1789) pl. E, figs. 9a-b; pl. 2, fig. 7). Cardium apertum sive hians, testa tenui etc. Chemnitz, 1782 :189, pl.18, figs. 181-183 (Not binominal). Cardium apertum Chemnitz: Bruguière, 1789: 226. Synonyms: Cardium "rogata" (sic) Gronovius, 1781: 266 and Index No 1125, pl. 18, fig. 5 (Not binominal). Cardium hians Spengler, 1799: 39. Cardium rugatum "Gronovius": Dillwyn, 1817: 125; Questionable synonyms: Cardium virgineum Linn,, 1758: 682. (See Dodge 1952: 67). Cardium bullatum Linn,, 1758:673. (See Dodge 1952: 37, 68). Types: C. apertum: Bruguière's 1789 description is based only on Gronovius and Chemnitz, and the only shells cited are the ones figured by these authors. Gronovius' specimen has not been traced, but Chemnitz's specimen is still in the ZMUC, Spengler collection, Ref. BIV 42. It is erroneously located from Jamaica. Dimensions: H=48.6, L=48.1, W=30.5. D=1.0. With 44 ribs. This shell is here selected as lectotype of Cardium apertum (pl. 1, figs. 9a-b). C. rugatum: The specimen figured by Gronovius was not located. C. hians: Spengler had two syntypes in his own collection of this nominal species, now in the ZMUC: the specimen BIV 42 above cited, and another specimen BIV 43. Dimensions of the latter: H=—43.6, L=44.4, W=29.3. D=0.92. With 44 ribs. The specimen BIV 42 is here selected also as lectotype of Cardium hians, so this taxon becomes an objective synonym of C. apertum. C. virgineum and C. bullatum: no type specimens located. Description: Shell of medium size, up to 48 mm in length. Shape relatively constant, inequilateral, with its anterior side short and rounded and its posterior side transversaly expanded, often VIDAL slightly truncated and always raised at its extremity, causing a significant gaping (pl. 2, fig. 7). Sometimes, the flattening of the lunular area sharpens the anterior side a little. No or very weak posterior groove. Length always a little greater than eight. Lunule very large, with a significant perilunular furrow. No lunular heart. No sublunule. Last flat present, but small, in the young shells only; when the shells become adult it decreases progressively in width, then disappears. Last fold rounded, high but narrow, without periostracal insertions. External colour variable, uniformly whitish or beige or with irregular concentric zones of purple red to brown, sometimes only spotted with these colours. The density of the coloured zones generally decreases with the growth of the shell. Umbo often purple. Internal colours generally the same by transparency except in the posterior area, almost always purple coloured. Sometimes, presence of a thin radial red strip in the umbonal cavity. Mean number of ribs 43 (range 39-48). In the whole AT and MT zones, the ribs are triangular, very narrow, and the interstices flat and comparatively very large. . In the PTI the ribs become flatly triangular, increasing in width, and in the PT2 they become equivalent to the interstices. The IMR extends far from the margin. The periostracal insertions are always situated in the posterior side of the ribs, and form complex arrangements. Granules very numerous, more or less aligned on short distances, present only on the AT or the anterior half of the shell. The hinge line has two specific characteristics: First, and paradoxically, its posterior side is relatively short ( ratio D = approximatively 1 or less). Secondly, it is appreciably angled in the middle of its anterior part (and not at the level of the umbo, like in the other species). Ocular organs observed. Material examined and distribution: The type specimens listed- Other lots: OMAN: Gulf of Oman: 1 MNHN- MAURITIUS: 1 ZMUC- THAILAND: Phuket: 1 ANSP: 2 MNHN- THAILAND: Gulf of Thailand: 1 MHNG- MALAYA: Srait of Malacca: 1 WAM, Gulf of Thailand: 1 WAM- PHILIPPINES: 2 MNAN, 6 ANSP- HONG-KONG: 1 MNEHN;: 1 BM(NH)- CHINA: Hainan: 1 MNHN- JAPAN: 2 MNEN- INDONESIA: Java: 1 MNHN- MALAYA: N. Borneo: 1 ANSP- PAPUA: 1 ANSP- AUSTRALIA: 1 MNHN- Queensland: 2 MNHN;: The genus Fulvia APEX 9(4): 93-118, déc. 1994. 3 ANSP- W. Australia: 1 MNHN; 3 ANSP: 1 ZMUC- NEW CALEDONIA: 9 MNHN, 3 ANSP- SOLOMON: 1 ANSP. The lots from the Eastern Indian Ocean are rather scarce, more numerous from Western Indian Ocean and Western Pacific. Habitat: In New Caledonia it prefers littoral muddy organic environments. Observations: Æulvia aperta is easily distinguished from the other species of the genus mainly by its gaping which is exceptionally absent, but also by its ribbing and the characters of its hinge. Fulvia (Fulvia) papyracea (Bruguière, 1789) (pl. 1, figs. 7, 10a-b and 11a-b) Cardium papyraceum, testa cordata, fragili etc. Chemnitz, 1782: 190, pl. 18, fig. 184 (Not binominal). Cardium papyracea Schrôter, 1788: 82 (Not binominal). Cardium papyraceum Chemnitz: Bruguière, 1789, Vers, I: 231. Synonym: Fulvia voskuili Healy & Lamprell, 1992: 89-91, pl. 4, figs. a-d. [= Æ sp. : Lamprell et al., 1992, n° 226]. Questionable synonym: Cardium pallidum Reeve, 1845: Sp 92, fig.92. Types: Cardium papyraceum: The description of Bruguière is just a translation of Chemnitz and the only shell cited is the one figured by the latter (pl. 18, fig. 184). This holotype 1s still preserved in the ZMUC, No BIV 44, Spengler collection. The old labels indicate "Jamaica", but Chemnitz gives East Indies as locality. Dimensions: H=41.0, L=37.4, W=25.0. Angle A=125°, ratio D=1.4. With 42 ribs (pl. 1, figs. 11a-b). Fulvia voskuili: Holotype in the QM reg. M0O32906 (pl. 1, Figs. 10a-b), from Kelso Reef (N. Queensland), H=34.3, L=33.8, W=22.0, angle A= 130°, ratio D= 1.3, with 33 ribs. - Paratype n° 1 in the AM (reg. C166907), from Dinpo "Beach (N" Old), 5310, E-310 W=20.6. - Paratype n° 2 in the Museum of Victoria (reg. No F60471), also from Dingo Beach, H=28.0, L= 28.4, W=17.7. Cardium pallidum: In the BM(NH), two shells are registered n° 1978.134 as syntypes of C. pallidum, considered as coming from the Bay of Manila, Philippines, Cuming collection. But, 99 APEX 9(4): 93-118 , déc. 1994, as mentioned on the label: "none agree with figure and locality. Has been crossed out". These two shells look very much like FÆ tenuicostata, since the description, figure and locality of Reeve's specimen strongly suggest it is À. papyracea. Description: Shell of medium size, up to 45 mm in height. Almost perfectly equilateral, rarely a little expanded backwards like the holotype. Rather depressed. No or very slight posterior truncation. Generally almost circular with L/H ratio very close to 1, the holotype being exceptionally a little elongated (L/H=0.91). No or very weak posterior groove. Lunule large. No sublunule. Lunular heart small. Last flat relatively wide and flat. Last fold wide and flatly rounded. External colour of the adult rather uniform, variable but almost always light, whitish to light brown, the young shell being more coloured with darker concentric stripes. Umbo often purple. Internal colour whitish with more or less numerous brown-purple concentric stripes and often a thin radial purple line in the umbonal cavity. Almost always purple coloured in the posterior zone, this coloration being sometimes also perceptible exteriorly. Mean number of ribs: 43 (range 37-48). With the exception of the four or five last ribs of the PT2, the ribs are always much narrower than the interstices, which are wide and flat. In the AT, the ribs are very thin and triangular and become flatly rounded and hardly perceptible in the MT, then more perceptible in the PT1. In the PT2 the ribs are closer one from another with almost always well marked intersticial riblets. The rib features of the AT and MT are discernible internally, but at the margin itself they are of variable distinctness. Periostracal insertions, always located in the posterior side of the ribs, form complex arrangements (pl. 1, fig. 7). Granulation always present on the anterior half of the shell. Hinge and dorsal margin looking symmetrical on both sides, with a mean angle A of 134° and a ratio D of about 1.3. Ocular organs: no data. Material examined and distribution: The holotype of C. papyraceum, and the holotype of F. voskuili- Other lots: INDIA: Gulf of Manaar: 2 ANSP- MALAYA: Strait of Malacca: 1 ANSP- SINGAPORE: 1 MNHN: 1 ANSP- PHILIPPINES: 2 ANSP; Manila: 1 QM: Subic Bay. Luzon: 1 100 The genus Fu/via MNHN- INDONESIA: Amboina: 1 MHNG: Yapen Island: 1 ANSP; Schouten Island: 1 ANSP- AUSTRALIA: N. QIld, Shelburne Bay: 1 QM, Gulf of Carpentaria: 1 QM. Other lots: In MNHN, one specimen in a lot of F australis labelled from Mauritius (old collection, mixing ?). In ANSP, two lots whithout locality (54159,54198). Observed also Lamarck's specimen in the MANG (see below). Observations: As mentioned by Chemnitz, F. papyracea 1s close to F aperta in appearance: both have equivalent thinness, same colours and also approximately a same structure of ribs and interstices. But F. papyracea never gapes and has some other characters to distinguish it from F. aperta: symmetry of the shell and of the hinge, ratio D distinctly higher, lunular heart more developped, last fold and last flat wider, intersticial riblets in the PT2 more frequent. F. papyracea is distinguished also from F. mutica which is larger and of different coloration, has a smaller lunule, much wider ribs with axial periostracal insertions, very different and typical IMR, and from F. australis mainly by its shape and its ribbing. The name papyracea is particularly erroneously used for a form of Red Sea, Western Indian Ocean and Persian Gulf, here described as F. fragilis, which is incontestably different (see the description of this species). F. voskuili has all the characters of F papyracea, except for the rib number of the holotype (33) which seems too low. Nevertheless three other specimens examined have: 42, 44, 47 ribs, in agreement with the range of }. papyracea. Although the name papyracea is the most used in the literature for the different species of the genus Fulvia (23% of all the citations according to the chresonymy of Fischer-Piette (1977:76, 78, 79), F. aperta and its synonyms excluded), Fulvia papyracea is a rather rare species with a limited area of distribution. In the literature, the subsequent citations corresponding undoubtedly to the species in question are also extremely rare. These are: 1)- LAMARCK 1819: 6. Locality: East Indies. The shell referred to is now in the MNHG. reg. No 1085/40. It is represented by a typical right valve. 2)- REEVE 1844, Sp 9, fig. 9. Locality: East end of the Island of Luzon (Philippines). Good description and figure. 3)- HIDALGO 1903: 344 Locality: Subic Bay, Island of Luzon (Philippines). The description is excellent. No illustration given. VIDAL VIDAL Fulvia (Fulvia) mutica (Reeve, 1844) (pl. 1, figs. 5 and 12a-b) Cardium muticum Reeve, 1844, Sp 32, fig. 32. Synonyms: Cardium japonicum Dunker, 1860: 223, 1861: 28, pl. 3, figs. 1-6. Cardium annae Pilsbry, 1904: 557, pl. 40, fig. 20. Cardium tcheliense Debeaux in Pilsbry, 1904: 558. Types: Cardium muticum: The only type is the specimen 67 mm long, from Cuming collection, without locality, figured by Reeve, not traced. Cardium japonicum: The type material of Dunker is now in the Senckenberg Museum in Frankfurt (see JANSSEN, 1993). It is a shell 73 mm long, from Japan. Cardium annae: The type specimens are two valves in the ANSP (reg. No. 86319): one left valve H= 20 mm, broken, and one right valve H=25.3, L=27.2. This right valve is figured by PILSBRY, pl. 40, fig .20 and here in pl. 1, figs. 12a-b). Cardium tcheliense: The specimen mentioned by Pilsbry, from China, is in the ANSP (reg. n° 54302). Its dimensions are: 24.4x26.3x17.7. Description: Fulvia mutica is the largest of all the known living Fulvia, reaching 100 mm in length. Its shape is relatively constant, roughly "diamond-shaped", almost equilateral, often with the posterior side a little longer. Not truncated but rather a little sharpened. The length is always a little larger than the height. Shell slightly gaping in the posterior side. Lunule small, without perilunular furrow. No sublunule. Lunular heart extremely small. Last flat narrow and a little grooved. Last fold rounded, high and well separated, bearing or not a periostracal insertion. External colour uniform, more or less dark beige. Internal colour white to purplish pink by zones. Mean number of ribs: 47 (range 43-54). The ribs are weak but clearly discermible in the AT and the PTI, where they are flatly rounded and about equivalent in width to the interstices. In the PT2 they are very numerous. Towards the MT, the ribs tend to be wider and flatly triangular, and become more and more indiscernible. On the other hand, in the IMR zone (which does not extend very far from the margin), the "ribs" are well and finely marked, the "interstices" being very wide. The genus Fulvia APEX 9(4): 93-118, déc. 1994. The periostracal insertions are thick and persistent, they are generally placed on the middle of the ribs. The insertion corresponds on the other side, in the IMR , to a very thin furrow on the middle of the "interstice" (very specific character). The insertions form a simple arrangement (pl. 1, fig. 5). Granulation often present on the lunule and the AT in the young specimens (less than 10-15 mm), but never in the adults. Hinge indistinguished, symmetric, with an obtuse angle A (range 135-150°). The ratio D varies from 1.2 to 1.4. Ocular organs observed. Material examined and distribution: The type specimens of C. annae and C. tcheliense- Other lots: (CHINA: 1 ANSP: Kingdao: 1 MNHN- HONG-KONG: 1 ANSP- KOREA: 1 ANSP- JAPAN: 11 MNHN,; 10 ANSP. According to KIRA (1962: 210) the species extends from South-China to Alaska. REID & SHIN (1983: 281) indicate a southern limit around Yamaguchi (Japan). Habitat: "Muddy bottom of sheltered waters" (KIRA, 1962: 210). Kira indicates also that " This is also one of the important edible shells in Japan”. Observations: ÆFulvia mutica can easily be distinguished from the other Fulvia by its particular "diamond-shaped" shape, its colours, but above all by its very particular ribbing and IMR. Fulvia (Fulvia) tenuicostata (Lamarck, 1819) (pl2, figs. la-b) Cardium tenuicostatum Lamarck, 1819: 5, No d: Synonyms: Cardium racketti Donovan, 1826: pl.124. Cardium radiatum Reeve, 1845: Sp 89, pl.89. Fulvia fagea Voskuil & Onverwagt, 1992: 42 (nomen novum for the latter). Types: Cardium tenuicostatum: There are three lots labelled by Lamarck in the MNHN. The first lot of two specimens 50.5 and 47 mm high, from Timor and New Holland. The second and the third lots with two and one specimen respectively, are somewhat different from the first lot (smaller and more equilateral) and respectively hand-labelled by Lamarck as "var." and "individual very young”. I select the largest shell from the first lot as lectotype: its 101 APEX 9(4) 93-118 , déc. 1994. dimensions are : H=50.5, L=540, W=307; number of ribs: 47, angle A= 138°, ratio D= 1.4. (pl. 2 , figs. la-b). The reference to Timor is very probably erroneous. In Lamarck's collection of the MHNG, there is another specimen comparable to the main syntypes of the MNHN, but not hand-labelled by Lamarck, reg. n° 1085/38. It is 43.5x44.9x29.4, A= 140°, D= 1.2. This specimen, figured by DELESSERT (1841, pl.11, figs. 6a-c), must also be considered as a paralectotype. Cardium racketti: The specimen figured by Donovan has not been traced. Cardium radiatum: Holotype in the BM(NH), reg. n° 1912.64. from Hanley collection, locality unknown. H=32.3, L=34.3, W=23.8. Angle A=130°, ratio D=1.31. Number of ribs: 52. Description: Shell of medium size up to 56 mm in length, generally thick and solid in the adult stage. Shape variable from almost equilateral to transversely and even obliquely expanded, often slightly truncated in the posterior side, which sometimes can be somewhat sharpened. Length always a little larger than height. No posterior groove. Lunule small with no perilunular furrow. No sublunule. Lunular heart small. Last flat large. Last fold very large with two parts often separated by a small furrow: a rounded posterior part and a flat anterior one. External colour light and uniform whitish to yellowish, rarely more or less concentrically mottled with darker colours. Umbo often purple coloured. Some specimens, a little differently coloured than usually, but in other respects typical, have been considered as specifically different (C. radiatum Reeve). Internal colour white, sometimes with a red-purplish stain in the umbonal cavity and more rarely on the posterior ventral edge. Mean number of ribs 51 (range 44-59). In the adult shells the ribs of the MT are as strongly marked as in the other two thirds. In the whole shell the ribs are equivalent in width with the interstices, or slightly wider.The ribs are generally more or less roundly triangular, often symmetrically. Rounded ribs can be present, but only in non adult shells. According to the symmetry of the ribs, the periostracal insertions can be placed in the axis or more or less in the posterior part of the ribs. No periostracal insertion on the last fold. The nodes of the periostracal insertions never become calcified and spiny in the PT. 102 The genus Fulvia The granules are never abundant. They are present mainly in the young shells, in the AT. No particularity in the hinge: mean A= 138°, mean D= 1.3. Ocular organs observed. Material examined and distribution: The type specimens of C. tenuicostatum and C. radiatum- Other lots: AUSTRALIA: 2 MNHN;: 2 ZMUC:; S.Eastern Australia: 3 MNHN: 5 AMS,; 2 BM(NH);, Southern Australia: 2 ANSP: 4 ZMUC; S.Western Australia: 4 MNEHN,; 2 AMS, 1 QM; 3 BM(NH), Tasmania: 2 MNHN; 2 QM, 2 BM(NH). Southern Australia only. According to Wilson and Stevenson (1977: 55), "from the vicinity of Sydney on the East Coast to Fremantle in W.A". Habitat: "Lives gregariously in sand and mud from two to 30m depth of water"(Ludbrook 1984:176). Observations: F tenuicostata differs from all the others (except F fragilis) by having, when adult, a solid and thick shell with strong ribs in the MT. It is apparently close to F° fragilis, but both can easily be separated (see this species). Fulvia fragilis (Forsskal in Niebuhr, 1775) (pl. 1, fig. 6; pl. 3, figs. la-b) Cardium fragile Forsskal in Niebur, 1775: 31. Types: Forsskal [in Niebuhr, 1775: 31] introduced the name Cardium fragile with the diagnosis: "striatum transverse; tantum ad cardines laeve". Although brief, this diagnosis qualifies as a description and the name C. fragile is nomenclaturally available. This opinion is shared by YARON et al.(1986: 95) who regard it as "eligible for standing as valid [taxon]". In ZMUC., there is a lot comprising 3 bivalves and one valve, originating from the Forsskal expedition to the Red Sea. This lot is without label and it is not absolutely certain that it corresponds to C. fragile. However, rather than naming a new species and leaving /ragile forever in limbo, it seems preferable to stabilize this name by designating one of the Copenhagen's putative syntypes as neotype. I designate here the shell figured by YARON et al. (1986:194, fig 42) as neotype of Cardium fragile. This shell: H=23.6, L=23.0, W=15.9, with 44 ribs, angle A=120°, ratio D=1.6. Description: Shell of medium to large size up to 75 mm in height. VIDAL Shape generally approximately symmetri- cal, often with slight posterior truncation, but sometimes with the posterior part more or less transversally or obliquely expanded. Mean L/H a little higher than 1. Posterior groove weak to non existent. Sometimes very slightly gaping posteriorly. Lunule large without perilunular furrow. No sublunule. Lunular heart variable but never large. Last flat wide and flat. Last fold very large and regularly rounded, bearing a periostracal insertion only in the young stage and in 50% of the specimens. External colour generally uniform whitish, beige to yellowish, with a purple stain only on the umbo and sometimes on the lunular heart and on the last flat and fold. Internally white, except on the posterior third which is almost always purple, and sometimes the umbonal cavity. Mean number of ribs 41 (range 34-52). In the young shell, up to about 2 or 3 cm in height, the ribs of the MT are much less well marked than in the other thirds, like in the majority of the Fulvia, but in the fully adult shells they become as strong as the others.In the whole shell the ribs are equivalent in width with the interstices, or slightly wider. In the AT the ribs are generally rounded, but can sometimes become more or less symmetrically triangular. In the MT they become progressively asymmetrically triangular (posterior slope shorter and steeper), and can become practically flatly rounded. They remain more rounded in the PTI, but without transition they become more or less symmetrically triangular in the PT2 where the last interstices often bear intersticial riblets. In this latter area, the periostracal insertions become calcareous at the nodes, and produce tubercles or spines (pl. 1, fig. 6). The periostracal insertions are always on the /crest, of” the ribs äin the AT; they progressively migrate to the posterior part of the ribs in the MT and also in the PTI: they migrate again on the crest of the ribs in the PT2. They form simple arrangements. No granulation in the adult shells, but sometimes someones very dense on the lunule and the two or three first ribs of young specimens. Indistinguished hinge: mean A= 138°, mean D rather high = 1.5. Ocular organs observed. Material examined and distribution: The type material- Other lots: EGYPT: Mansaleh lake: 1 MNEN,; Timsah lake: 3 BM(NH), Great lake: 1 The genus Fulvia APEX 9(4): 93-118, déc. 1994. ANSP: Suez canal: i MNHN: Gulf of Suez: 11 MNHN- ISRAEL: Elat: 1 MNHN,; 1 ZMUC- Red Sea (EGYPT, SUDAN, ERITREA, ARABIA, YEMEN): 6 MNAHN,; 2 AMS- DyIBOUTI: Gulf of Tadjoura: 5 MNHN- YEMEN: Aden: 2 MNHN- OMAN: Gulf of Oman: 2 MNHN- South Persian Gulf (TRUCIAL COAST, QATAR, BARAIN): 3 MNEN, 3 AMS, 1 ANSP: 1 BM(NH), 5 ZMUC- N. SOMALIA: 3 MNHN- KENYA: 1 MNEAN- ZANZIBAR: 1 MNHN- MOZAMBIQUE: 1 MNHN, 1 ANSP: SOUTH AFRICA: Natal: 2 MNEN, 2 BM(NH)), 2 NM- ANGOLA: Luanda: 4 MNHN- CONGO: Pointe Noire: 4 MNHN- GHANA: Accra: 1 NM- COTE D'IVOIRE: 1 MNEAN. The species migrated through the Suez Canal and is present in the Mediterranean coast of Israel, together with F australis (see Barash & Danin 1992: 275). The status of the populations found in the Atlantic Ocean will be discussed by Dr. von Cosel in a separate paper. Habitat: Littoral to shallow water. Observations: The first figuration of this species was in SAVIGNY's plates (1805-1812: pl. 13, fig. 9). AUDOUIN (1827: 200) named these figures only: Cardium....Subsequently Fulvia Jfragilis has generally erroneously been named F. papyracea, and sometimes F. tenuicostata. It differs from the former mainly by the thickness of the shell and strength of the sculpture, the larger width and different profile of the ribs and the absence of granulation, and from the latter by the lower number of ribs, the presence of rounded ribs in section, periostracal insertions situated on the posterior side of the ribs, rarely observed in the Australian species, and by some characters of the lunule and posterior flat and fold. It can be separated from F. australis, with which it is sympatric (except in the Atlantic Ocean), by its less oblique shape, its rounded and large instead of triangular and smaller last fold, its fewer ribs, the presence of well rounded ribs never observed in }. australis. The distinction from the other species can also be bhelped, particularly with young specimens, by the presence in Æ /fragilis of "spines" in the PT2 and sometimes à periostracal insertion on the last fold. In other respects the young specimens seems to be variable as far as shape, proportions and colours are concerned and can look very different, from one to another and from the adults, which 1s often a little disconcerting. 103 APEX 9(4) 93-118 , déc. 1994. Fulvia (Fulvia) natalensis (Krauss. 1848) (pl. 2, figs. 2a-b and 2c). Cardium natalense Krauss, 1848: 12, pl. 1, fig.9. Cardium natalense var. adjacens Turton, 1932: 243, No 1710, pl. 65, fig. 8. Types: Cardium natalense: Krauss had only three small specimens and cited and figured one of them: H=14.6, L=16.6, W=9.0. These syntypes could be in the Stuttgart Museum. Cardium natalense adjacens: Turton had only two tiny valves to define his subspecies, of which he selected the largest (6x8 mm!) as type. This holotype could be in the Oxford University Museum. Remark: The present study is based on the examination of much larger and representative specimens, particularly from the lot n° 9576 Natal Museum, Kilburn collection, with numerous large specimens from Port Elizabeth (pl. 2, figs. 2a-b and fig. 2c). Description: Shell of medium size up to 43 mm in length. Very inequilateral, with the posterior zone transversally expanded. rarely obliquely. Always longer than high: mean L/H= 1.12 (range 1.04-1.21). Anteriorly rounded and also posteriorly (never truncated). No posterior groove. Large posterior gape. Lunule generally small to very small. No perilunular furrow. Lunular heart very little developed. Last flat narrow, concave. Last fold medium, flatly triangular, with always one, sometimes two, periostracal insertions. External colour generally beige to light brown, more or less mottled with brown-purple, rather uniform, a little darker on the umbo. Interior uniformly or irregularly dark brown-purple. Mean number of ribs 45 (range 41-50). Ribbing rather regular, of the same strength on all the shell, the width of the ribs being everywhere equivalent to the one of the interstices, which is particularly visible in the IMR. The ribs are generally triangular and the interstices rounded. In the AT the ribs are symmetrically triangular and they become progressively asymmetric in the MT (posterior side shorter) and can become almost flat. In the PT the ribs are a little more rounded. No granulation. Periostracal insertions well marked, generally with predominance of the radial components, the concentric components often 104 The genus Fulvia being absent or very small. The insertions are situated on the crest of the triangular ribs in the AT, migrating towards the posterior side in the MT, then coming back on the crest in the PT. Hinge line slightly arced (A range 140- 150°). without any angular break (difference with aperta). Left anterior lateral almost confused with the margin and much more distant from the umbo than the anterior lateral (ratio D range 1.5-2.0). Ocular organs: no data. Material examined and distribution: S. MOZAMBIQUE: Inhambane Bay: 2 MNHN- SOUTH AFRICA: St Lucia Bay: 1 MNHN,; Port Alfred: 1 NM; Jeffrey's Bay: 1 NM; Plettenberg Bay: 2 MNAHN,; Knysna Lagoon: 2 MNHN; 1 NM, Still Bay: 1 NM; Capetown: 1 ZMUC. Endemic of the south-Eastern coast of South Africa, from Port Alfred to False Bay. Habitat: Mainly found in estuaries, in calm littoral waters of variable salinity, probably in muddy organic terrigenous facies. Observations: ÆFulvia natalensis, often erroncously named as papyracea, has also been confused with F aperta because of its gape. Nevertheless the latter differs in its less transversally expanded and less rounded shape, its lighter colour, its ribbing (ribs narrower than the interstices), its very much smaller ratio D, its larger lunule and its high and narrow last fold without periostracal insertions. Fulvia (Fulvia) australis (Sowerby, 1834) (pl. 2, figs. 3a-b, 4, 5 and 6a-b) Cardium australe Sowerby, 1834: fig. 12 and 1840: 105. Synonym: Cardium striatum Spengler, 1799: 45. Questionable synonyms: Cardium varium Sowerby 1834, fig 19. Cardium pulchrum Reeve 1845, Sp 98, fig 98. Types: Cardium australe, varium, pulchrum: No type specimen referred to these three nominal species. Cardium striatum: Two syntypes in ZMUC, Spengler collection, from South Seas. N° 1: 24.4x25.1x17.0, 50 ribs. N° 2: 23.8x23.8x15.5, 55 ribs. Remark: The name Cardium striatum has priority over C. australe, but australe is used because: (a) striatum has never been used subsequently to Spengler, other than FISCHER- VIDAL VIDAL PIETTE (1977: 160) in a list of “uncertain species"; (b) australis has been used over 50 times since its description, and is currently the second most widely used specific name in the genus Fulvia. (An application to conserve the name }#. australe will be presented to the International Commission on Zoological Nomenclature). Description: Shell of medium size, up to 35 mm in height. The shells are generally thin, but rarely excessively, and are relatively solid. Shape sometimes almost equilateral but generally obliquely ovate, inequilateral, with the dorsal anterior part inflated and the posterior part more or less obliquely expanded. Almost always slightly truncated in the PT2. Posterior groove generally well developed. Lunule of medium size with a small lunular heart. Perilunular groove sometimes present. Sublunule often present but always short (equivalent of two or three ribs in length). This zone is very often red coloured. and bears periostracal insertions (it is therefore more easily discernible in the IMR). Last flat medium, sometimes concave. Last fold markedly triangular, mainly in the young shells, with no periostracal insertions. Exterior colour very variable from pure white to dark brown purple. The young specimens (and the early formed part of the adult shells), are often more coloured, with generally darker, irregular, more or less concentrically arranged stains of variable density. The intestices are not stained, or are lighter coloured than the ribs. Interior generally white to yellow or light pink, with the exterior darker stripes transparently visible and also some radial rays. Rarely presence of a slight reddish coloration in the posterior side. The mean number of ribs is 49, but this number is extremely variable (range 34-68) between areas and populations (see below). The ribs are similar and homogeneous in AT and MT, weakly marked, equivalent in width to the interstices or a little narrower, flat or very slightly rounded; interstices flat. In these two zones the IMR is also homogeneous and well marked. In the PTI the ribs and interstices become irregular and the interstices become very wide with regard to the ribs which take a triangular profile. Intersticial riblets in the PT2 often present, but more or less well developed. The periostracal insertions are always situated on the posterior limit of the ribs in the The genus Fulvia APEX 9(4): 93-118, déc. 1994. AT and MT, but tend to migrate to the crest of the triangular ribs in the PT. Granulation of variable density, on individuals and populations, sometimes lacking, generally limited to the anterior half of the shell, on the ribs as well as in the interstices, sometimes aligned but always on short distances. Hinge line and dorsal margin caracteristically asymmetric, with a receding posterior side and a raised and inflated anterior one , forming a vague angle like a hump. Angle A relatively "acute", mean value 133° (range 130-140), and ratio D rather high: mean 1,3 (range 1.2-1.5). Ocular organs observed. Material examined and distribution: The syntypes of C. striatum- Other lots: EGYPT: Gulf of Suez: 5 MNHN,; Marsa Thlemel: 1 MNHN- ISRAEL: Elat: 1 ZMUC- DyIBOUTI: 2 MNHN- N. SOMALIA: 1 MNHN- OMAN: Gulf of Oman: 1 MNHN- ARABIA: S. Persian Gulf: 1 MNHN- KENYA: 1 MNHN- ZANZIBAR: 1 MNHN- TANGANYIKA: Dar-es-Salaam: Il AMS- MOZAMBIQUE: Vilanculos: l MNAN- MADAGASCAR: E. coast: 5 MNHN; Dredging Tulear: 51 MNHN - MAURITIUS: 9 MNAN; 1 ZMUC- SEYCHELLES: 1 MNHN- INDIA: Gulf of Manaar: 2 ANSP- W. THAILAND: Phuket: 2 MNEAN, 1 ZMUC,; 1 ANSP- MALAYA: Tocal: 1 ZMUC- VIETNAM: 1 MNAHN,; Poulo Condor: 2 MNAN, Natrang Bay: 1 ZMUC- CHINA: 2 MNEHN- JAPAN: 1 MNHN- PHILIPPINES: Manila 1 MNAN,; Luzon: 1 WAM- AMER. PAC. TRUST TERR: Nukuoro Atoll: 1 WAM- INDONESIA: Molucas, Cayeli: 1 MNHN- PAPUA: Hansa Bay: 2 IRSNB: Madang: 1 WAM- SOLOMON: 1! ANSP: 1 QM.; 1 WAM- GILBERT: Kiribati: 1 AMS- TONGA: 1 MHNG; 1 WAM- Fi: 3 MNAN,; 1 WAM- VANUATU: 2 MNHN-NEW CALEDONIA: 113 MNHN,; 2 ANSP- AUSTRALIA: 1 MNHN,: Queensland: 2 MNHN:; 2 AMS: 5 QM. Found in the Suez canal and in the Mediterranean coast of Israel and Palestine, together with F fragilis (see BARASH & DANIN, 1992: 275-276). Habitat: littoral to shallow water. Seems to prefer clean reefal or perireefal facies. Observations: Fulvia australis 1s the most common and widespread species of Fulvia and can be sympatric with almost all the others. It can easily be differentiated mainly by its very characteristic oblique asymmetry and the strong contrast between the ribbing in the AT and MT, dense, serried, regular and homogeneous, and the ribbing in PT, loose and irregular. As à 105 APEX 9(4) 93-118 , déc. 1994, matter of fact it has not often been confused with the other species, unless in very young specimens. Nevertheless, in spite of its constant characters allowing always an accurate identification, this species can be variable in shape, colour and number of ribs. À good illustration of this variability can be found in New Caledonia, where very numerous lots of this species are available, from several environments. It is possible to distinguish three forms: 1)- Forms comparable to the types of Sowerby and Reeve: shells of rather small size, shightly asymmetric, rather depressed (mean W/L = 0.65), strongly coloured outside and inside, with a small number of ribs (34 to 44). They are found mainly in the lagoon of the Ouvea atoll, at water depths from 10 to 20 meters. Such forms (with more ribs) exist locally in Australia, and sporadically elsewhere as individual variants. A specimen from a lot close to this form, from Shelburne Bay, North Queensland, lot QM No 45517, is figured here in pl. 2, figs. 6a-b: H=26.5, L=24.8, W=17.0, with 55 ribs. 2)- (pl. 2, fig. 5), shells of average size, mainly asymmetric, fairly depressed (mean W/L = 0.70), coloured by irregular concentric stripes of brown or purple (ribs only coloured, not interstices), and frequently with the umbo, the sublunule and the lunular heart purple coloured, an average number of ribs (40-55). Mainly in littoral zones. This form is the most common form of the species, found everywhere, more or less coloured, sometimes entirely white (see also pl. 2, fig. 4 an intermediate form). 3)- (pl. 2, figs. 3a-b), shells of rather large size for the species, almost symmetric, more globular (W/L up to 0.80), lightly coloured with or without darker stripes, with a large number of ribs (50-68), and no or rare granules. They are found mainly in the lagoon at water depths from 15 to 80 meters and exceptionally in the littoral zones. Comparable forms are also present in the coast of East Africa, islands of the Indian Ocean, Melanesia, China. This differentiation in New Caledonia suggests the existence of a significant influence of the environment on certain characters of this species. Nevertheless, the above three forms cannot be considered as subspecies because of their geographical dispersion and the existence, even at the populational level, of many intermediate forms. 106 The genus Fulvia Fulvia (Fulvia) fragiformis sp. nov. (pl. 3, figs. 2a-b) Types: The only known four shells, in MNEN, from New Caledonia. Holotype: Found alive near Abore reef (22°47' S-166°40' E), Depth 15m, H=16.5, E=15.7, W=11.2, with 52 ribs. Angle À =150°, ratio D= 1.4. (pl. 3, figs. 2a-b). Paratype n° 1: Dredged in the SE lagoon, sta.731 (21°17'2 S-165°52'0 E), depth 40m, H=12.1, L= 110, W= 90, with 49 ribs, Paratype n° 2: Dredged in the S lagoon, sta.589 (22°31'7 S-167°23'0 E), depth 31m H=11.1, L= 106, W=7.4, with 45 ribs. Paratype No 3: A left valve dredged in the SW lagoon, sta.21 (22°22'1 S-166°23'4 E), depth 16m, H=16.6, L=16.8, 1/2W=5.3, with 46 ribs. Description: Shell small, the paratype No 3 being the largest, thin and fragile. Very special shape comparable to some FRAGINAE, with the PT in a different plane and separated by a rounded angle . This PT is largely reduced. In consequence of these particular features, the shell has an uncommon asymmetry, with the anterior half longer than the higher posterior half. Posterior groove present, well marked in the interior of the shell. Lunule small, without perilunular furrow. Lunular heart very small. No trace of coloured sublunule. Last flat small, slightly concave. Last fold large, almost flat. External colour white to yellowish, with more or less zigzagging irregular brown-purple concentric stripes. À brown spot on the umbo. Internal colour similar by transparency. Number of ribs 45 to 52. In the AT and the MT the ribs are mainly flat rounded and roughly equivalent in width between themselves, the interstices being a little wider than the ribs, particularly in the AT. In the PTI the ribs and interstices are equivalent and wider, the ribs being rounded to flatly triangular. In the PT2 the ribs become more triangular with, in the smallest specimen, interstices wider and flat. In all specimens the IMR is very little marked in the PT2. Periostracal insertions strong, situated on the posterior side of the ribs in the AT and MT, then on the middle of the triangular ribs in the PT, with the nodes sometimes a little calcified. Granulation present, but not very abundant in the AT and sometimes in the PT. Hinge thin with small teeth. Ocular organs observed. VIDAL Material examined and distribution: The type material. Habitat: Shallow water in clean non terrigenous facies. Observations: Fulvia fragiformis is very close to F. australis, except for its very characteristic shape, never observed in the latter. There are also slight differences in the ribbing and the IMR (a little less marked), in the thicker periostracal lamellae, and in the stronger granules in Æ fragiformis. Fulvia (Fulvia) boholensis sp. nov. (pl. 3, figs. 6a-b) Types: The type series in the MNHN. Holotype: from Panglao, Bohol Island, Philippines. It is said to have been recovered at 80 fms. Dimensions: H=36.0, L=35.2, W=21.6. Angle A=130°, ratio D=1.2, 55 ribs (pl.3 .figs. 6a-b). Paratype n° 1: From Philippines, without precise data, H=33.5, L=34.0, W=21.1. Angle A=130°, ratio D=1.2, 54 ribs. Paratype n° 2: From Sulu Sea, Philippines. H=33.3, L=35.1, W=20.8. Angle A=130°, ratio D=1.1, 52 ribs. Paratype n° 3: From Philippines. without precise data, H=28.0, L=27.4, W=17.2. Angle A=130°, ratio D=1.2, 50 ribs. Description: Shell of medium size, up to 36 mm in length, relatively thin and fragile. Always significantly asymmetric, obliquely expanded backwards, truncated in the PT2, rather depressed (mean W/L=0.62). Posterior groove well developed. Lunule medium with à small lunular heart. Sublunule often present but always very short (ke F australis). Last flat medium. Last fold flatly triangular with no periostracal insertions. Colour almost always orange or yellowish- orange internally and externally, the umbo and margins often being darker. The lighter medial zone of the shell often bears a little darker irregular concentric stripes. Some rare specimens (like paratype No 3) have "normal" colours: beige, light brown, purple, a part of the internal PT only being orange. The only observed specimen from outside of the Philippines (Western Australia) is yellow- orange with white margins. Mean number of ribs 51 (range 47-55). Ribs a little triangular in the AT, with wider interstices, flat or flatly rounded in the MT, with interstices about equivalent in width, The genus Fulvia APEX 9(4): 93-118, déc. 1994. slightly triangular with wider interstices in the PT2. IMR extending far from the margin, often not perfectly marked in the MT which has a thin and fragile margin. Periostracal insertions of rather complex arrangement, situated on the posterior side or on the crest of the ribs. One of the radial components of these insertions become calcified and form, at each node, a thin oblique often elongated tubercle, in the PT. Granules numerous in the anterior half of the shell, often aligned on short distances. Ocular organs: no data. Material examined and distribution: The type specimens and four corresponding lots- Other lots: PHILIPPINES: Bohol: 1 QM- AUSTRALIA: Western Australia, Dampier: 1 QM. Observations: Fulvia boholensis seems to be an intermediate form between Æ australis and F. papyracea. It differs from the former by its nibbing, particularly its IMR which 1s smoother, from the latter by its ribbing and its asymmetry. It differs from both by its particular colours, and also from all FÆulvia by the particular arrangement of the periostracal insertions in the PT forming elongated carbonaceous tubercles. (The equivalent tubercles of Æ /ragilis are different and not elongated). Fulvia (Fulvia) dulcis (Deshayes, 1863) (pl. 1, fig. 8; pl. 2, figs. 8a-b) Cardium dulce Deshayes, 1863: 12, pl. 29, figs. 3-5. Types: Four syntypes in the MNHN from Réunion, 12.0 to 7.1 mm in eight. The largest: H=12.0, L=11.5, W=8.7, with 41 ribs, 1s selected here as lectotype (pl. 2, figs. 8a-b). Description: Shell small, the above lectotype being the largest specimen observed. Shell roughly equilateral, with the anterior margin rounded and the posterior one a little straightened. Almost circular, with L/H close to 1. Presence of a weak posterior groove corresponding inside to a well marked ridge. Lunule of medium size, without perilunular furrow. Lunular heart rather large. Presence of a long sublunule, about equivalent in length to the lunule. Last flat small. Last fold very large, a little rounded to flatly triangular (pl. 1, fig. 8). 107 APEX 9(4): 93-118 , déc. 1994. External colours: in the Indian Ocean the shells are whitish with a pink coloration in the umbo, diminishing towards the margin, forming a few irregular concentric stripes more or less zigzagging. The AT and PT zones are also light pink. Some specimens are entirely white.In the Pacific, same coloration but light brown instead of pink. Interior with same colours by transparency. Number of ribs: 41 to 47. The ribs of the AT and MT zones are equivalent in width with the interstices. They are flatly rounded, low, and hardly perceptible with the naked eyes, with no difference between these two zones. The IMR of the above zones is also regular, well perceptible, extending far from the margin which is also distinctly serrated. In the PTI the ribs become suddenly wider and stronger, with equivalent interstices. They are also well marked, but smaller and more numerous, in the PT2 with possibly some intersticial riblets. In the lunule and sublunule, the growth lines form very thin concentric folds. Periostracal insertions (3 to 5) without ribs in the sublunule. Present also in the AT and the MT and situated on the posterior limit of the ribs. The insertions migrate to the axial part of the ribs in the first ribs of the PT, then can disappear: in that case some ribs of the PT do not bear any insertion, or rudimentary ones. Granules often numerous, but never aligned, on the posterior side of the shell, even on the posterior flat and fold (pl. 1, fig. 8); they are rarer on the anterior part. They can be totally absent from the whole shell. Hinge with no particularities. with both sides almost symmetric. Ocular organs observed. Material examined and distribution: The type specimens- Other lots: SOUTH AFRICA: Zululand: 10 NM- PAPUA: Hansa Bay: 1 QM- NEW CALEDONIA: 33 MNAN. Habitat: In Réunion, according to DRIVAS & JAY (1988: 16), it is "found alive in muddy black sand at 12 to 30 m depth". In New Caledonia also it is exclusively present in shallow terrigenous facies, East of the island, between 20 and 50m. In South Africa it was dredged in fine sandy facies between 45 and 50 m depth. Observations: F: dulcis is easily distinguished from several small forms of ÆFulvia by its periostracal insertions, and from the young specimens of F. australis by its more equilateral and globular shape and by its longer sublunule. 108 The genus Fulvia Fulvia (Fulvia) scalata sp. nov. (pl. 1, figs.3 and 4; pl. 3.figs. 5a-b and 7) Types: Types from the eastern lagoon of New Caledonia. Holotype: In the MNHN, sta.833 (20° 27'6- 164° 44'3), depth 27m, H=19.1, L=19.3, W=14.8, with 46 ribs, (p1.3, figs. 5a-b). Paratype n° 1: In the MNHN, same sta. as holotype, H=17.6, L=18.4, W=12.7, with 43 ribs. Paratype n° 2: In the MNHN, sta.883 (20°276 S-164°44'3 E), depth 27m, H=17.1, L=17.2, W=12.4, with 42 ribs. Paratype n° 3: In the MNAN, sta.873 (20°38'55 S-164°46'2 E), depth 27m, H=15.0, L=15.2, W=11.5, with 37 ribs. Paratype n° 4: In the MNHN, sta.847 (20°37'65 S-165°13'4 E), depth 28m, H=11.0, L=11.1, W=8.3, with 38 ribs. Paratype n° 5: In the AMS, same sta. as holotype, H=166, L=16.8, W=11.1, with 48 ribs. Paratype n° 6: In the ANSP, same sta. H=17.7, L=17.8, W=13.0), with 51 ribs. Paratype n° 7: In the NMNZ, same sta, H=17.3, L=16.7, W=12.7, with 45 ribs. Description: Shell small, the holotype being the largest specimen observed. Roughly equilateral, with the anterior margin rounded and the posterior one a little straightened. Almost circular, with L/H very close to 1.0. Presence of a very weak posterior groove, only in adult shells. Lunule rather small, without perilunular furrow. Lunular heart extremely large, the largest of all the Fulvia (pl. 1, fig. 3). Presence of a sublunule of rather variable length, but never as long as the lunule (varying between 1/2 and 3/4 of the lunule), with no periostracal insertions but sometimes with a rudimentary rib in its middle. Last flat extremely large, sometimes divided into two differently tilted zones. Last fold also extremely large with a posterior triangular part and a flat large anterior one. External colour generally white to light brown, rarely with concentric irregular darker stripes. Umbo and posterior part often darker coloured. Internally, the umbonal cavity is often yellow to pinkish and the posterior part slightly purple. Number of ribs about 40 (range 36-44). With the exception of the PT2, the nbs are strictly equivalent in width to the interstices. The ribs, well rounded in the AT, then becoming more or less flatly rounded or flat in the MT, are well delimited, contrasting with the VIDAL flat interstices. The ribs are weaker in the MT but always well perceptible in the adult shells. In the PTI the ribs and interstices are a little wider and stronger than in the other parts of the shell. In the PT2 the interstices are very narrow, the two last ribs being very large and flat, and the others narrow and a little triangular. The IMR is very well marked and extended. In the lunule and sublunule, the growth lines form thin concentric folds. Periostracal insertions situated on the posterior limit of the ribs in the AT and MT, then migrating progressively to the top of the ribs in the PT. Granulation almost always present both in the anterior half, including the lunule, and in the posterior half, including the last flat and fold. In the AT and PTI, the granules are located in the interstices, aligned and placed at more or less regular intervals, like rungs in a ladder (scalata). In the very young shells, granulation can also be developped in the MT, forming irregular concentric lines (pl. 1, figs. 4 and pl. 3, fig. 7). Hinge typical of the genus, very asymmetric as far as laterals are concerned (D= 1.7 to 1.8). with a particular triangular nymph strongly projecting exteriorly beyond the margin of the shell. Ocular organs observed. Material examined and distribution: The type series- other lots: OMAN: Mascate : 1 MNHN- MADAGASCAR: Tulear: 4 MNHN- PHILIPPINES: Visayas and Luzon: 2 MNHN, 1 WAM- INDONESIA: Macassar Strait: 1 MNHN; Moluccas 1 WAM- PAPUA: Hansa Bay: 3 IRSNB- NEW CALEDONIA: 75 MNHN- AUSTRALIA: Queensland: 7 AMS; Carpentaria: 1 QM, Western Australia: Dampier: 1 QM: 1 WAM. Habitat: In New Caledonia only found in shallow water from 20 to 50 m (never littoral), and in the terrigenous sandy facies of the Eastern coast (rarely in clean perireefal facies). Fulvia (Laevifulvia) undatopicta (Pilsbry, 1904) (pl.1, figs.1 and 2: pl. 2, figs.9a-b, 10 and 11) Cardium hungerfordi undatopictum Pilsbry. 1904: 556, pl. 40, figs. 14-15. Cardium undatopictum Pilsbry: Nomura & Niino, 1940: 55. Synomym: Cardium hungerfordi stigmaticum 1904: 556, pl. 41, figs. 13-14. Pilsbry. The genus Fulvia APEX 9(4): 93-118, déc. 1994. Types: Cardium hungerfordi undatopictum: Syntypes in the ANSP, reg. n° 80521: from Hirado, Hizen, Japan (Hirase collection, 1901), four paired specimens and one right valve, from 11.6 to 18.3 mm in height. The largest (H=18.3, L=17.5, W=10.5), with 48 ribs, is figured by Pilsbry and here in pl. 2, figs. 9a-b. Cardium hungerfordi stigmaticum: Syntypes in the ANSP, reg. n° 86.279: also from Hirado, Hizen, Japan, three paired specimens from 10.5 to 11.7 mm in height. The largest (H=11.7, L= 11.0, W= 7.5) is figured by Pilsbry and here in pl. 2, fig. 11. Description: Shell small. The first syntype above cited being the largest observed. Generally roughly equilateral, rarely appreciably obliquely inequilateral (like some syntypes). Often slightly truncated posteriorly, sometimes with a small open indentation in the margin corresponding with the posterior groove which is more or less developed. Shell a little higher than long: L/H range 0.88-0.93. Lunule of medium size with a rather small lunular heart (pl. 1, fig 2), both not coloured. Sublunule long, length equal to the lunule (pl. 1, fig. 1). External colour very variable from pure white to almost entirely dark brown purple. Often whitish to beige with more or less developed irregular brown red stripes or zigzags. Sometimes only coloured by small spots. Four different red stains can be present, often together, forming a cross: one stain on the umbo, another one radially elongated in the middle of the shell (the variety stigmaticum of Pilsbry is a pure white form with only these two stains, see pl. 2, fig. 11), a third one on the sublunule, and the last one on the posterior margin on the PT2 zone. In the syntypes of undatopicta, these stains are not present, or extremely pale. The internal colours are the same, by transparency. Mean number of ribs about 48 (range 46- 53). with 10 in the PT. From the medio- posterior part of the MT, where the ribs are present but not easily discernible, they become progressively a little more developed and wider towards the AT where, however, they remain of moderate development. Towards the PT, the ribs evolve in the same way, but they become quickly well marked and wider in this zone.They are generally flatly rounded, except in the PT2 where they become somewhat triangular, and equivalent in width with the 109 APEX 9(4) 93-118 , déc. 1994. interstices. The IMR is always clearly discernible and shows the same evolutions as the external ribbing. Intersticial riblets present in the PT2, where the ribs tend to be very wide close to the posterior margin. In the sublunule and the posterior part of the lunule, the growth lines tend to form thin concentric folds. No periostracal insertions. Granules more or less abundant, but always present, concentrically aligned on long distances in the AT (pl. 2, fig. 10), rare to absent in the MT, more or less abundant and not aligned in the PT. Hinge characteristic of the genus, regularly curved. Nymphal plate narrow, not triangular. Ocular organs observed. Material examined and distribution: The type material- Other lots: Red Sea: 4 MNHN- EGYPT: Gulf of Suez: 2 MNHN- DJIBOUTI: 1 MNHN- YEMEN: Aden: 1 MNAN- MADAGASCAR: Dredging Tulear: 8 MNHN: Nosy Bé,: 1 MNHN- THAILAND: Kut Is: 1 AMS- PHILIPPINES: Visayas: 1 MNHN- JAPAN: Tosa: 1 MNHN- PApUA: Hansa Bay: 1 IRSNB;: 1 QM- NEW CALEDONIA: 232 MNEHN - AUSTRALIA: Queensland: 5 AMS: Gulf of Carpentaria: 1 AMS. Habitat: Only shallow water, rarely littoral. In New Caledonia, lives in reefal or perireefal calcareous facies, between 10 and 35m. Fulvia (Laevifulvia) lineonotata sp. nov. (pl. 3, figs.4a-b) Types: Types from New Caledonia. Holotype: In MNEAN, sta.992 (20°16'1 S-163°56'6 E), depth 28 m, H=13.6, L=11.1, W=8.2, with 52 ribs (pl. 3, figs. 4a-b). Paratype n° 1: In MNHN, same sta. as holotype, H=8.9, L= 8.0, W= 52. Paratype n° 2: In MNHN, Chesterfield lagoon staDW 31 (19°24'86 S-158°45'03 E), depth 57m, H=10.2, L= 9.6, W=63. Paratype n° 3 in MNEHN, Chesterfield lagoon sta DW 50 (19°18'30 S-158°33'57 E), depth 50m, H=102, L=9.6, W=6.1. Paratype n° 4 In MNEN, Chesterfield lagoon sta. DW40 (19°29'46 S- 158°3527NE); "depth "58m HI NE 87 W=6.0. Paratype n° 5: In AMS, sta.1174 (19°21'3 S-163°14"2 E), depth 53m, H=102, L=9.2, W=6.1. Paratype n° 6: In ANSP, same sta’ as” holotype, H=9.0;" E-82,-W=S3; Paratype n° 7: In NMNZ, sta.542 (19°06'4 S- 163°10'0 E), depth 5Om, H=100, L=91; W=6.0. 110 The genus Fulvia Description: Shell small about 10mm in height, the holotype of 13.6 mm being the largest observed. Always a little elongated (L/H between 0.84 and 0.94). Asymmetric with the anterior margin rounded and the posterior margin a little obliquely expanded, with a small depression at the extremity of the posterior groove, which 1s weakly but always developed. Lunule rather small without lunular heart and with no or very small perilunular furrow, limited by a coloured line formed by triangular spots. Presence of a very long sublunule (1.5 times or more the lunule), the longest of all the Fulvia. External coloration constituted by juxtaposed or imbricated differently coloured (white to brown purple) triangles, with more or less diffuse limits. The posterior part of the MT is marked by a radial alignment of darker coloured triangles , alternating with white ones. This coloured "line", almost always perceptible, gave its name to the species. Umbo and often posterior margin stained red purple. Internal coloration the same as the external one by transparency. Ribs almost indiscernible in the AT and MT, but the IMR, limited to a small band, is clearly discernible and allows counting of about 40 ribs in these two zones. The PT bears about 13-15 well marked ribs, flatly triangular, with narrow interstices. The narrowest ribs (about four in number) are located in the anterior part of the PT2. These ribs are often poorly marked and can disappear. Except in the PT, the growth lines form very thin concentric folds. No periostracal insertions. Thin granules, often concentrically aligned, on all the anterior half of the shell including the lunule. Hinge very thin, particularly the nymphal plate, and well arced. Angle A of medium value for the genus, and ratio D higher than 1.0. Ocular organs observed. Material examined and distribution: The type material- Other lots: MAURITIUS: Port Louis: 1 NM- INDIA: Maldives, Milandu Atoll: 1 BM(NH)- NEW CALEDONIA: 45 MNHN. Habitat: in New Caledonia exclusively shallow water. in clean calcareous facies from 25 to 60m. Observations: Fulvia lineonotata is easily separated from the other small Laevifulvia by its elongated oblique shape, its particular colour pattern, its long sublunule. VIDAL VIDAL Some small specimens of Zaevicardium lobulatum Deshayes, 1855, which are sometimes sympatric in New Caledonia, can have approximately the same shape and colours, and easily be confused. Nevertheless, the latter has still less perceptible ribs in the AT and MT and no ribs in the PT2, many more "ribs" in the IMR (about 85 versus 55), a shorter sublunule and no granules. In addition the umbo of this species bears a special reticulated ornamentation, never observed in any Fulvia, and it never has ocular organs. Fulvia (Laevifulvia) hungerfordi Sowerby, 1901 (pl. 2, figs. 12a-b, 13, 14 and 15) Cardium (Papyridia) hungerfordi Sowerby, 1901: 103, pl. 9 fig 5. Types: Three syntypes from Japan, preserved in the BM(NH), reg. 1902.5.28.5.7, respectively 9.0, 8.4 and 8.0 mm in length. The two latter are figured here pl. 2, figs. 12a and 12b. Description: Shell small up to 14 mm in length. Always longer than high (L/H range 1.02 to 1.16). Generally inequilateral with the posterior part expanded, but never obliquely; nevertheless Indonesian forms are quasi-perfectly equilateral. Posterior margin a little truncated in the PT2, and PTI1 often lengthened, overstepping a little the margin of the shell: some slight straightenings in the ventral and anterior margins can give the shell like a vague “"polygonal" aspect, with a sharpening in the posterior margin. Posterior groove weak but well marked in the early stage of growth, disappearing in the adult shell. Lunule medium, flat or a little concave, limited by a significant rounded ridge, clearly delineated also in the early stage of growth. Lunular heart small. Sublunule of variable length, according to the different populations (between one half and almost one and a half times the length of the lunule). Last flat large, a little concave. Last fold large, flatly triangular. Colour of the shells, external and internal, uniform, from beige to brown or red purple. becoming lighter towards the margins. About 10 ribs in the AT, 12 ribs in the PT and 20-22 ribs in the MT when this third 1s ribbed. The ribs are generally equal in width with the interstices, except in the PT2 where the interstices are very thin. In the AT the ribs have The genus Fulvia APEX 9(4): 93-118, déc. 1994. a rounded or slightly flatly rounded profile and the interstices are flat. In the PTI1 the ribs and the interstices, the largest of all the shell, both have a rounded profile. In the PT2 the ribs flatten. As far as the the MT is concerned, it can be entirely ribbed like the AT, or partially ribbed, or not ribbed, according to populations (see later).In all the forms, the growth lines form, in the adult shells, more or less strong concentric folds, except in the PT. No periostracal insertions. Granulation of variable density (sometimes absent) on the whole shell, sometimes perfectly concentrically aligned in the MT. Hinge with no particularities. Siphonal eyes present according to REID & SHIN, 1983: 277, fig. 1. Material examined, distribution and observations : Observed specimens from four areas: JAPAN: Syntypes and specimens of the ANSP cited by PILSBRY (1904: 555, pl. 41, fig. 17). Inequilateral forms with no ribbing in the MT. Found in littoral and shallow water in organic muddy facies. Recorded also from Hong Kong (REID & SHIN, 1983: 275), but with no data about the shell itself. PHILIPPINES: One lot from Musorstom Campaign 1985, sta. DR 140, Jintololo channel, North of Panay Island, depth 93-99m in clean muddy and shelly facies (thanatocenosis). Inequilateral forms with partial variable development of nbs in the MT (pl. 2, fig. 13). INDONESIA! Two lots from Siboga Expedition: sta.53, North of Sumba Island, depth 36m in coral sand and sta.174, East of Ceram Island, in very shallow reef facies. In both areas the shells are almost equilateral with ribs in the MT (pl. 2, fig. 14). NEW CALEDONIA: Only in one locality: Gatope Beach, near Voh, in the North-Western coast, littoral in deltaic argillaceous sandy facies. Asymmetric forms with ribs in the MT (pl. 2, fig. 15). Fulvia (Laevifulvia) prashadi sp. nov. (pl. 3, figs. 8a-b) Types: In the ZMA, from Siboga expedition, sta.71, South of Macassar Strait, Indonesia, depth 32m, in mud, sand with mud and coral. Identified as Cardium hungerfordi Sow by PRASHAD (1932: 281). Holotype: A right valve H=10.2, L=10.2, 1/2W=04 (pl, figs. 8a-b). Paratype n° 1- À left valve H=9.3, L=9.3, 111 APEX 9(4): 93-118 , déc. 1994. 1/2W=0.37. Paratype n° 2- A right valve H= 9.0, L=9.0, 1/2W=0.35. Paratype n° 3- A left valve H=7.5, L=7.7, 1/2W=0.3. Description: Shell small, the holotype being the largest known specimen. Shape relatively constant, inequilateral, with the anterior side short and slightly trunçated and the posterior one not obliquely expanded. Often the MT margin is straightened in its posterior part. The PT is sharpened and clearly isolated from the rest of the shell by a significant notch in the margin, the PT forming a ridge making the margin pointed. Lunule medium, a little concave, limited by a rounded ridge well delineated, even in the earliest stage of growth. Lunular heart small. Sublunule equivalent to the lunule in length, or a little longer. Last flat large, a little concave. Last fold large, flatly triangular. External colour uniformly light beige. Internally yellow, lighter in the axis of the umbonal cavity, with the posterior part a little purplish. About ten ribs in the AT equivalent in width to the interstices, with a well marked IMR. No ribs nor IMR in the MT. About ten strong ribs in the PT, equivalent in width to the interstices in the PTI, wider in the PT2, corresponding to a strong IMR. In the whole shell, presence of well marked rounded concentric folds, corresponding to the growth stages. No periostracal insertions. Never seen any granulation on the few specimens observed. Hinge with no particularities. Ocular organs: no data. Material examined and distribution: Found only by the Siboga expedition in Indonesia in five stations: sta. 71, South of Macassar Strait near Sulawesi coast: eleven valves (type series) sta.4, East Java, depth 9m in coarse sand: 1 specimen and one valve - sta.33, East Lombok Island, depth 22m in mud, coral and coral sand: one specimen - sta.164, South of Salawati Island, North East New Guinea, depth 32m in mud, sand and coral: one specimen - sta.114, North coast of Sulawesi, depth 75m in very fine hard sand: one specimen. Observations: Æulvia prashadi is close to certain forms of F hungerfordi and could be considered as a special form or a subspecies of the latter. The problem needs more material to be solved. In favour of a specific separation 1s the fact that the only two known sites of F 112 The genus Fulvia hungerfordi in Indonesia (Siboga sta. 53 and 174) are close to sites of the former, and contain quite different symmetrical specimens (pl. 2, fig. 14), constant in characters. It is the same for F. prashadi which shows very constant characters in the five sites where it is found in Indonesia. Anyway, I think that these very special forms merit a new name (species or subspecies). Fulvia (Laevifulvia) ballieni sp. nov. (pl. 3, figs. 3a-b) Type: Holotype in the MNHN labelled "Sandwich Islands, Mr Ballien 1876". It may seem improbable that such a large and remarkable Hawaïian bivalve has remained unknown to this date. Indeed, that it is known from a single shell, collected in the 19th century, in a European Museum, is suspect. No Fulvia or Laevicardium is known from Hawaii, and there is no material in the Bishop Museum (KRASLOWITZ, pers. comm.). | cannot exclude the possibility that the shell of F: ballieni has been erroneously labelled and in fact it does not originates from Hawaï. However, there are indications in the catalogue of the MNHN that a Mr Ballien did send repeatedly shells from Honolulu between 1872 and 1878. Fulvia ballieni, be it Hawaïian or not, must be a rare species, as evidenced by the fact that no other specimen has shown up since Ballien's collect in 1876. Description: Shell of medium size: H = 48.0, L = 49,3, W = 343. Practically equilateral, but with a very small double-truncation making the PT slightly sharpened. No gaping. Lunule not separated from the sublunule, which is a little shorter than the lunule. Lunular heart very small. Last flat hardly differentiated, a little concave. Last fold narrow, triangular. External colour uniformly light beige, shining. Interior whitish, with the margin yellowish. Number of ribs 50. The ribs and interstices are hardly marked on the whole shell, a little more in the PTI. No ribs in the PT2 which is smooth, externally and internally. The IMR, which extends rather far from the margin, is very better marked than the ribbing and shows "ribs" a little narrower than "interstices". No periostracal insertions. VIDAL VIDAL Granules more or less aligned present only on the umbonal part up to about one centimetre from the umbo. Hinge typical of the genus. Angle A = 130°. Ratio D = 1.32. Ocular organs: no data. Material examined and distribution: Only the holotype, maybe from Hawaii. Observations: Shining apart, Fulvia ballieni has approximately the same appearence as F. mutica from which it differs by many other characters (no periostracal insertions, rather long sublunule, etc...). It differs mainly from the other species of the subgenus Laevifulvia by the size, the homogeneity of the ribbing and IMR in the AT and MT, by the presence of a smooth PT2. Because of this latter character it approaches the genus Laevicardium (convergence). ACKNOWLEDGEMENTS: I! am especially grateful to Dr Philippe Bouchet and Dr Bernard Métivier, in the MNHN of Paris, for their help in many ways, and to Dr Rudo von Cosel for its advice at manuscript stage. I am also indebted to Dr Bertrand Richer de Forges (ORSTOM, Nouméa) for his assistance. I thank also very much the following people for loans and information: Mr Ian Loch (AMS, Sydney), Dr Gary Rosenberg (ANSP, Philadelphia), Ms Kathie Way (BM(NH), London), Dr J. van Goethem and Dr Claude Massin (IRSNB, Bruxelles); Dr Yves Finet (MNHG, Genève), Dr R. N. Kilburn (NM. Pietermaritzburg);, Mr Kevin Lamprell and Dr John Stanisic (QM, Brisbane), Dr Robert Moolenbeek (ZMA, Amsterdam), Dr Tom Schiotte (ZMUC, Copenhagen). REFERENCES: AUDOUIN, V., 1827 - Explication sommaire des planches de Savigny. Panckoucke, Paris. BARASH, A. & Z. DANIN, 1992 - Annotated list of Mediterranean Mollusks of Israel and Sinaï. Fauna Palaestina. Mollusca 1. The Israel Acad. of Sciences and Humanities. Jerusalem. BRUGUIERE, J.C., 1789 - Encyclopédie Métho- dique, Histoire Naturelle des Vers, 1. Panc- koucke, Paris. CHEMNITZ. J. H., 1782 - Neues Systematisches Conchylien Cabinet, 6. Raspe, Nürnberg. DELESSERT, B., 1841 - Recueil des coquilles décrites par Lamarck et non encore figurées. Fortin, Masson et Cie, Paris. The genus Fulvia APEX 9(4): 93-118, déc. 1994. DESHAYES, G. P., 1863 - Catalogue des Mol- lusques de l'île de la Réunion (Bourbon). Dentu, Paris. DILLWIN, L. W., 1817 - Descriptive Catalogue of Recent Shells.. I.J. & A. Arch, London. DODGE, H., 1952 - A Historical Review of the Molluscs of Linnaeus, Part1,The Class Loricata and Pelecypoda. Bull. Amer. Mus. Nat. Hist, 100, art.I. DONOVAN, E., 1826 - Naturalist's Repository, 4. DRIVAS, J. & M. JAY, 1988 - Shells from Réunion, 18 Fam. Cardiidae. La Conchiglia, 20 (232,233): 13-17. DUNKER, (G., 1860 - Neue Japanische Mollusken. Malacozool. Blätt., 6: 221-240. DUNKER, G., 1861 - Mollusca Japonica Des- cripta et Tabulis Tribus Iconum. Stuttgartiae. FISCHER-PIETTE, E., 1977 - Révision des Cardiidae (Mollusques Lamellibranches). Mém. Mus. Hist. Nat. Paris, N.S.A, 101. GRAY, J. E., 1847 - A list of the Genera of Recent Mollusca, their synonyma and types. Proc. Zool. Soc. London, Part 15: 129-219. GRAY, J. E., 1853 - A revision of the genera of some of the families of Conchyfera or Bivalve shells. Ann. Mag. Nat. Hist., (2), 11: 33-34. GRONOVIUS, L. T., 1781 - Zoophilacium Gronovianum. Th. Haak & al. Leiden. HEALY, J. & K. LAMPRELL, 1992 - New species of Veneridae, Cardiidae,… from Australia (Veneroida, Bivalvia, Mollusca). J. Malac. Soc. Australia, 13:75-97. HIDALGO, G., 1903 - Estudios preliminares sobre las Faunas Malacologicas de las Islas Filipinas. Mem. Acad. Cienc. Madrid, 21: 207- 306. IREDALE, T., 1929 - Strange molluscs in Sydney harbour. Australian Zool. 5, part 4. JANSSEN, R., 1993 - Die Typen der von Dunker 1860/1861 beschriebenen Japanischen Meeresmollusken. Arch. für Molluskenk. 112: 403-435. KAFANOV, A. I. & S. V. POPOV, 1977 - On the system of the Cenozoic Cardioidea (Bivalvia). Paleontological Journal, 11 (3): 307-314. KEEN, À. M. 1936 - Revision of Cardiids Pelecypods. Proc. Geol. Soc America for 1935, Preliminary abstracts: 367. KEEN, À. M., 1951 - Outline of a proposed classification of the Pelecypod family Cardndae. Conchological Club of Southern California, Minutes 111: 6-8. 113 APEX 9(4): 93-118 , déc. 1994. KIRA, T., 1962 - Shells of the Western Pacific in Colour. Hoïkusha, Osaka. KRAUSS, F., 1848 - Die Südafricanischen Mollusken. von Ebner & Seuberg, Stuttgart. LAMARCK, J. B.,, 1809 - zoologique...Dentu, Paris. LAMARCK, J. B., 1819 - Histoire Naturelle des Animaux sans Vertèbres, 6(1). Paris. LAMPRELL, K & T. WHITEHEAD, 1992. Bivalves of Australia 1. Crawford House Press, Bathurst. LUDBROOK, N. H., 1984 - Quaternary Molluscs of South Australia. Dept. of mines and Energy, South Australia, Handbook No 9. LYNGE, H., 1909 - The Danish Expedition to Siam 1899-1900. IV. Marine Lamellibranchiata. Mem. Acad. Roy. Denmark, Ser. 7, 3, (3): NIEBUHR, C. 1775 - Descriptiones animalium..quae observavit Petrus Forsskal. Molleri, Copenhagen. NOMURA, $S. & H. NIINO, 1940 - Mollusca dredged from off the coast of Hukui Prefecture in the Japan Sea. Rec. Oceanogr. Works Japan, 12, 1: 23-81. PILSBRY, H. AÀ., 1904 - New Japanese Marine Mollusca. Pelecypoda. Proc. Acad. Nat. Sci. Philadelphia, 56: 550-561. PRASHAD, B., 1932 - The Lamellibranchia of the Siboga expedition. Systematic, Part 2, Pelecypoda. Brill, Leiden. REEVE, L. 1844 - Conchologia Iconica, Cardium, Sp. 44-64 (Sp 47 excluded). REEVE, L. 1845 - Conchologia Cardium, Sp.47 & Sp. 65-82. REID, R. G. B. & P. K. S. SHIN, 1983 - Notes on the biology of the cockle Fulvia hungerfordi (Sowerby). Proc. 2nd Intern. Workshop on the Malacofauna of Hong-Kong and Southern China, Part 2, 1: 274-282. SAVIGNY, J. C.,1805-1812 - Planches de l'Expédition d'Egypte. Histoire Naturelle. Mollusques. Panckoucke, Paris. Philosophie Iconica, SCHNEIDER, J. A. 1992 - Preliminary cladistic analysis of the bivalve family Cardiidae. Amer. Malac. Bull.. 9 (2):145-151. SCHRÔTER, J. S. 1788 - Vollständiges Alphabetisches Namen Register von. conchylien Cabinets. Raspe, Nürnberg. SOWERBY, G. B., 1834 - The Conchological Illustrations, 48th-51st Parts. SOWERBY, G. B.. 1840 - An extensive series of new species of the Genus Cardium exhibited by 114 The genus Fulvia Mr Cuming. Proc. Zool. Soc. London (8): 105- 1 ISS SOWERBY, G. B., 1901 - Descriptions of five new Species of Marine Shells. J. Malac., 8: 100-103. SPENGLER, K., 1799 - Over den toskallede Slaegt Cardium Linnei. Skrivt. Naturhist. Selsk. Kôbenhagen 5(1). SWAINSON, W. 1840 - Malacology, or Shells Longman & al, London. TURTON, W. H., 1932 - The marine shells of Port Alfred, S. Africa. Oxford University Press, London. VOSKUIL., R. P. À. & W. J. H. ONVERWAGT, 1992 - Studies on Cardiidae. 6. Gloria Maris 31(3): 33-44. WILSON, B. R. & S. E. STEVENSON, 1977 - Cardiidae of Western Australia Western Australian Museum Spec. Publ. 9. A Treatise on and Shell- Fish. YARON, L., T. SCHIOTTE & G. WIUM-ANDERSEN, 1986 - A review of molluscan taxa described by P. Forsskal and C. Niebuhr..., Sfeenstrupia 12 (10):157-203. PLATE 1 (opposite page): Fig.1 - Fulvia undatopicta, NC SE lagoon sta.703. MNAN. Real L=9.4mm. Fig.2 - Fulvia undatopicta, lunular heart, NC NW lagoon sta.992. MNHN. Scale: x14. Fig.3 - Fulvia scalata, lunular heart, NC N lagoon sta.1063. MNHN. Scale: x14. Fig.4 - Fulvia scalata, N Qld 15°45-145°385, AMS C44644. Real L=3.0mm. Fig.5 - Fulvia mutica, periostracal insertions in the PT. Japan. MNAN. Scale: x4s. Fig.6 - Fulvia fragilis, periostracal insertions in the PT. Aden. MNHN. Scale: x55. Fig.7 - Fulvia papyracea, periostracal insertions in the MT. Subic Bay Philippines. MNAN. Scale: x150. Fig.8 - Fulvia dulcis, posterior part. NC SE lagoon sta.688. MNHN. Scale: x15. Fig.9a-b - Selected lectotype of Cardium apertum Brug. and C. hians Spengler. Real L=48.1mm. Fig.10a-b - Holotype of Fulvia voskuili. Real L=33.8mm. Fig.11a-b - Holotype of Cardium papyraceum. Real L=37.4mm. Fig.12a-b - Syntype of Cardium annae figured by Pilsbry. Real L=27.2mm. VIDAL VIDAL The genus Fulvia APEX 9(4): 93-118, déc. 1994. 115 APEX 9(4): 93-118 , déc. 1994. The genus Fu/via VIDAL PLATE 2 (opposite page): Fig.1a-b - Selected lectotype of Cardium tenuicostatum. Real L=54.0mm. Fig.2a-b - Fulvia natalensis, Port Elizabeth SA, NM 5776. Real L=39.0mm. Fig.2c - Fulvia natalensis, Port Elizabeth SA, NM 5776. Real L=37.4mm. Fig.3a-b - Fulvia australis, NC S$ lagoon sta.557, depth 44m. MNHN. Real L=31.9mm. Fig.4 - Fulvia australis, N Qld Fantome Is, QM mo 45527. Real L=34.6mm. Fig.5 - Fulvia australis, NC Nouméa, MNHN. Real L=27.1mm. Fig.6a-b - Fulvia australis, N Qld Shelburne Bay, QM mo 45517. Real L=24.8 mm. Fig.7 - Selected lectotype of Cardium apertum. (See pl.1, fig.9a-b). Fig.8a-b - Selected lectotype of Cardium dulce. Real L=11.5mm. Fig.9a-b - Cardium hungerfordi undatopictum, syntype figured by Pilsbry. Real L=17.5mm. Fig.10 - Fulvia undatopicta, detail of the AT, NC SE lagoon sta. 703, MNHN. Scale: x22. Fig.11 - Cardium hungerfordi stigmaticum, syntype figured by Pilsbry. Real L=11.03 mm. Fig,.12a-b - Two syntypes of Cardium hungerfordi. Real L respectively 8.0 and 8.4mm. Fig.13 - Fulvia hungerfordi, Philippines N of Panay Is., sta. DR 140, MNAN. Real L=13.6mm. Fig.14 - Fulvia hungerfordi, Indonesia N Sumba Is. ZMA. Real L=12.4mm. Fig.15 - Fulvia hungertordi, NC Gatope Beach, MNHN. Real L=9.0mm. 116 VIDAL The genus Fulvia APEX 9(4): 93-118, déc. 1994. 117 APEX 9(4) 93-118 , déc. 1994, The genus Fu/via VIDAL ee PLATE 3: Fig.1a-b - Fulvia fragilis, Zanzibar, MNHN. Real L=45.3. Fig.2a-b - Holotype of Fulvia fragiformis. Real L=11.0mm. Fig.3a-b - Holotype of Fulvia ballieni. Real L=49.3mm. Fig.4a-b - Holotype of Fulvia lineonotata. Real L=10.2mm. Fig.5a-b - Holotype of Fulvia scalata. Real L=19.3mm. Fig.6a-b - Holotype of Fulvia boholensis. Real L=35.2mm. Fig.7 - Fulvia scalata, detail of the AT, NC NE lagoon sta. 846, MNHN. Scale: x25. Fig.8a-b - Holotype of Fulvia prashadi. Real L=10.2mm. 118 HOUART & ABREU The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. The Muricidae (Gastropoda) from Madeira with the description of a new species of Ocenebra (Ocinebrina) (Muricidae: Ocenebrinae). Roland HOUART 3400 Landen (Ezemaal) Belgium Research Associate at the Institut Royal des Sciences Naturelles de Belgique and Anténio Domingos ABREU Museu Municipal do Funchal, Madeira KEY WORDS: Gastropoda, Muricidae, review, new species, Madeira. ABSTRACT. Nine species of Muricidae are reported here from the Archipelago of Madeira (Hexaplex trunculus, Ocenebra erinaceus, ©. edwardsi, ©. aciculata, ©. inordinata n.sp., Bedeva paivae, Muricopsis aradasii, Cytharomorula grayi, and Stramonita haemastoma). Two of these species, Hexaplex trunculus and Bedeva paivae, are new records for the Archipelago. Three species listed by previous authors remain doubtful records for the region and have not been confirmed [Muricopsis cristatus, Orania fusulus, and Trophonopsis richardi (= T. droueti or T. muricatus)]. The presence of Typhis fistulatus (= Typhis sowerbyi), once recorded from the Archipelago, is probably based on a misidentification, and is not accepted here. À new species, Ocenebra (Ocinebrina) inordinata, is described from the Island of Madeira. RESUME. Neuf espèces de Muricidae sont signalées dans l'Archipel de Madère (Hexaplex trunculus, Ocenebra erinaceus, O. edwardsi, O. aciculata, O. inordinata n.sp., Bedeva paivae, Muricopsis aradasii, Cytharomorula grayi, et Stramonita haemastoma). Deux de ces espèces, 4. trunculus et B. paivae sont signalées pour la première fois dans la région. Trois espèces listées par des auteurs précédents n'ont pas été retrouvées, leur présence dans l'Archipel reste douteuse [Muricopsis cristatus, Orania fusulus, et Trophonopsis richardi (= T. droueti ou T. muricatus)]. La présence de Typhis fistulatus (= T. sowerbyi), signalée précédemment dans l'Archipel est probablement basée sur mauvaise identification de l'espèce et n'est pas acceptée ici. Une nouvelle espèce, Ocenebra (Ocinebrina) inordinata est décrite de l'Ile de Madère. INTRODUCTION The Archipelago of Madeira is situated between 35°05' - 32°25' N and 17°15' - 16°15" W. It includes Madeira, Porto Santo, the Desertas and a group of islets around these islands. Administratively, the Selvagens Islands, situated 160 miles south of Madeira, between 30°12' - 30°02' N and 16°05' - 15°50" W, also belong to the madeiran archipelago. The largest island of the archipelago is Madeira, which comprises an area of 737 km and is situated 700 km off the coast of Morocco. Porto Santo is situated 20 miles NE of Madeira and its area is approximately 41 km” including the islets. The Desertas are composed of the Ilhéu Chäo, Deserta Grande and the Bugio, which are situated 10 miles SE off Madeira and have an area of approximately 14 km. The Selvagens have an area of 4 km? (Text Fig. 1). Despite small local and seasonal variations, the Archipelago of Madeira is strongly influenced by a current which flows from NE and is formed by a terminal branch of the Gulf Stream. The mean temperature at the surface of the water in Madeira is 19.5° C. The lowest 1s 17.0° C (February and March) and the highest is 22.5° C in September. In the Selvagens the mean values are 0.3° C higher. Faunal elements of several provinces are represented in the composition of the marine Mollusca of Madeira. There are species from North Africa, Western Mediterranean, NW and NE Atlantic, but also several dozen endemic species. From the biogeographic point of view, the Archipelago of Madeira is a part of Macaronesia which also includes the Azores, the Canary Islands and the Cape Verde Archipelago. 119 APEX 9(4)} 119-130, déc. 1994 The Muricidae from Madeira HOUART & ABREU Se ee x 4 \ 1 , 0 ‘ 1 4 4 De + e ” [42 4 RC, Lt Archipelago of Madeira and Selvagens Islands (Adapted from Michel-Thomé, 1976 - Geology of the Middle Atlantic Island) 120 HOUART & ABREU The actual known total of the marine molluscan fauna of Madeira is about 500 species. The first author giving a general view of this matter was WATSON (1897) with a list containing 382 species. Later, NOBRE (1937) added 13 species to Watson's list. Apart from these two authors, NORDSIECK & TALAVERA (1979) have made a general study which, however, was only devoted to the Gastropoda. More recently, MOOLENBEEK & FABER (1987 a- c) and MOOLENBEEK & HOENSELAAR (1989) have dedicated special attention to the Rissoidae, describing some new species. The family Muricidae in Madeira is represented by several species, but not all the species reported by previous authors were rediscovered by us. Of course, it does not mean ipso facto that these do not occur in Madeira at all, and later reports perhaps will confirm the presence of these in the Archipelago. Abbreviatons BMNH: The Natural History Museum, London. MMF: Museu Municipal do Funchal, Madeira. RH: Roland Houart collection. SYSTEMATICS Family MURICIDAE Rafinesque, 1815 Subfamily Muricinae Rafinesque, 1815 Genus Hexaplex Perry, 1811 Hexaplex trunculus (Linné, 1758) Figs 1-2 Murex trunculus Linné, 1758: 747. Records: Madeira (Garajau), Porto Santo; Desertas Is, many live and dead taken specimens at a depth of approximately 70 m. (MMF). Remarks: Æexaplex trunculus is a very common species occuring in the Mediterranean Sea. The species was not reported before from the Madeira Archipelago, despite the fact that it is very common in Madeira and the other islands. There are at least 40 synonyms known for this variable species, most of them named to designate minor morphological differences in shell structure such as the length of spines, the dimensions, the colour, etc. The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. Subfamily MURICOPSINAE Radwin & D'Attilio, 1971 Genus Muricopsis Bucquoy & Dautzenberg, 1882 Subgenus Muricopsis Muricopsis (Muricopsis) cristatus (Brocchi, 1814) Murex cristatus Brocchi, 1814: 394, pl.7, fig.15. Records: NOBRE, 1937: Funchal. Remarks: As noted above, Muricopsis cristatus was listed in NOBRE (1937), but it 1s not yet recorded by us from the Archipelago. Its presence in Madeira, even if doubtful, is possible. The species is common in the Mediterranean Sea. Subgenus Murexsul Iredale, 1915 Muricopsis (Murexsul) aradasii (Monterosato, 1883) Figs 3-4 Murex aradasii Monterosato in Poirier, 1883: 1287 Murex (Ocinebra) medicago Watson, 1897: 242, pl. 19, fig. 11. Murex medicago -NOBRE, 1937: 29 Muricopsis medicago -NORDSIECK & GARCIA- TALAVERA: 131. Records: Only a few specimens from Madeira (Caniçal) and Porto Santo (MMF); WATSON, 1897 (as M. medicago nsp.): Madeira, Punta de Lourenço to 50 fms. (91 m); Magdalena (dredged); Selvagem Grande, shore; NOBRE, 1937 (as M. medicago): same as above, and Funchal; NORDSIECK & GARCIA- TALAVERA, 1979 (as M. medicago): Madeira. Remarks: No live specimens but only fresh dead specimens have been recorded until now by us. The species is apparently rare in the Archipelago (as it generally is elsewhere). There are only a few synonyms, amongst them Murex medicago Watson, 1897, described from Madeira. 121 APEX 9(4): 119-130, déc. 1994. Subfamily ERGALATAXINAE Genus Cytharomorula Kuroda, 1953 Cytharomorula grayi (Dall, 1889) Figs 5-6 Nassarina grayi Dall, 1889: 183, pl. 32, fig. 12a. Trophon lowei Watson, 1897: 244, pl. 19, fig. 12 Trophon lowei -NOBRE, 1937: 28. Urosalpinx lowei -NORDSIECK & GARCIA- TALAVERA, 1979: 134. Records: Some live and dead taken specimens from Madeira (with no precise locality data, and Funchal Bay) and from Porto Santo, up to 100 m. depth (MMF), WATSON, 1897 (as 7rophon lowei n.sp.): Madeira, 50 fms (91 m) (Labra and Punta de Säo Lourenço): NOBRE, 1937 (as TZ. lowei): same as above: NORDSIECK & GARCIA-TALAVERA (as Urosalpinx lowei): Madeira. Remarks: Nassarina grayi was described from Barbados (West Atlantic), Trophon lowei from Madeira and a third synonym, Cantharus laevis Smith, 1891, was described from St. Helena. The species is also known from the Canary Islands. It is now classified in the Muricidae, subfamily Ergalataxinae, due to morphological affinities of shell and radular characters with other species of the subfamily. The geographical distribution is probably world- wide because closely related specimens, probably belonging to the same species, have been recorded from the Indian and Pacific Oceans. Genus Orania Pallary, 1900 Orania fusulus (Brocchi, 1814) Murex fusulus Brocchi, 1814: 409, pl.8, fig.9. Records: NOBRE, 1937: Funchal; Porto Santo; NORDSIECK & GARCIA-TALAVERA, 1979: Madeira. Remarks: The species 1s not yet recorded by us, but its presence in Madeira is not doubted due to its presence from the Mediterranean Sea to Angola (West Africa). 122 The Muricidae from Madeira Subfamily OCENEBRINAE Cossmann, 1903 Genus Ocenebra Gray, 1847 Ocenebra erinaceus (Linné, 1758) Fig. 7 Murex erinaceus Linné, 1758: 748. Murex (Ocinebra) erinaceus - WATSON, 1897: 294. Murex erinaceus -NOBRE, 1937: 28. Ocenebra erinaceum -NORDSIECK & GARCIA- TALAVERA, 1979: 132. Records: Madeira (many localities); Porto Santo; Desertas Is, many live and dead taken specimens (MMF); WATSON, 1897: from Funchal to East point and Porto Santo (abundant); NOBRE. 1937: same remarks as above; NORDSIECK & GARCIA-TALAVERA, 1979: Porto Santo. Remarks: The species is common in the Archipelago. It is also common in the Eastern Atlantic and the Mediterranean sea, with a lot of synonyms, because of the many shell variations. Subgenus Ocinebrina Jousseaume, 1880 Ocenebra (Ocinebrina) edwardsi (Payraudeau, 1826) Figs 9-10 Purpura edwardsi Payraudeau, 1826: 155, pl. 7, fig. 19, 20. Murex (Ocinebra) edwardsii -WATSON, 1897: 294. Murex edwardsi -NOBRE, 1937: 28. Ocinebrina edwardsi -NORDSIECK & GARCIA- TALAVERA: 133. Records: Madeira (locinc.) (MMF): WATSON, 1897: from Funchal westwards. Very abundant, NOBRE, 1937: Funchal, Porto Santo; NORDSIECK & TALAVERA., 1979: Madeira. Remarks: Numerous specimens are present in the collections of the Museum of Funchal, all from Madeira, but with no precise locality data. The species is widely dispersed in the Mediterranean and the East Atlantic Ocean, and many forms have been named. All specimens examined from Madeira are strongly spirally sculptured., with low, broad axial varices. HOUART & ABREU HOUART & ABREU Ocenebra (Ocinebrina) aciculata (Lamarck, 1822) Fig. 8 Murex aciculatus Lamarck, 1822: 176 Murex (Ocinebra) aciculatus -WATSON, 1897: 294. Murex aciculatus -NOBRE, 1937: 29. Ocinebrina aciculata -NORDSIECK & GARCIA- TALAVERA, 1979: 133. Records: Madeira (Ponta Gorda), Porto Santo and the Desertas Is (common) (MMF); WATSON, 1897: from Madalena to island's East point and Porto Santo; NOBRE, 1937: from Madalena to East point; Porto Santo, Funchal, Pontinha; Porto Santo; NORDSIECK & TALAVERA, 1979: Madeira. Remarks: A small species, reaching a maximum length of 15 mm. It is common in the Eastern Atlantic and the Mediterranean Sea. It is not variable morphologically, however some form names have been proposed for smaller, larger, or more colourful specimens. Ocenebra (Ocinebrina) inordinata n.sp. Figs 11-13 Type material: Madeira Is (no other locality data), holotype MMF 25429, 19.2 mm; 1 paratype coll. R. Houart, 14.2 mm, 1 paratype coll. J. Verstraeten, 21 mm. Description: Shell medium sized for the subgenus, up to 21 mm in length at maturity, heavy, tuberculate. Spire high with 1.25-1.50 protoconch whorls and up to 6 shouldered, strongly nodose teleoconch whorls. Suture appressed. Protoconch whorls rounded, weakly elongate, smooth, terminal varix very shallow, nearly straight. Axial sculpture consisting of ridges and varices: 12 low axial ridges on first teleoconch whorl; 11 on second whorl; 10 or 11 low to high ridges on third whorl;, 9 high, strong ridges on 4th whorl; 7 high ridges, and varices on Sth whorl; last teleoconch whorl with 4 or 5 erratically placed varices, some with low, blunt open spines, and one or two high, strong axial node. Spiral sculpture consisting of 2 nodose cords on first teleoconch whorl; 2 primary cords and 1 secondary cord on the shoulder on second whorl; third to fifth teleoconch whorls with 2 primary cords and narrow threads between them, some 2 or 3 additional threads on shoulder; last teleoconch whorl with 5 or 6 low. The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. obsolete cords, forming short, rounded, broadly open spines on the varices, chiefly on apertural varix. Occasionally with 2 or 3 low, shallow threads between cords. Aperture ovate, moderately large. Columellar lip smooth, margin partially weakly erect, adherent at adapical extremity. Anal notch broad. Outer lip erect, smooth, with 5 or 6 strong nodes within, adapical node strongest. Siphonal canal short, narrow, straight, closed, smooth. Shell entirely light brown; aperture glossy white. Remarks: We are aware of the great diversity of forms existing in the Ocenebra (Ocinebrina) edwardsi group of shells, but ©. inordinata does not fit any of these forms. A great number (more than 500 specimens) of ©. edwardsi were observed from different localities, representing many varieties (both colour and morphological forms). Moreover, the varieties of ©. edwardsi are generally mixed, and many forms live together in the same region. We also examined more than 200 specimens of ©. edwardsi from Madeira, all are very similar morphologically. Ocenebra inordinata constantly differs in its completely white aperture, in the few, strong nodes on its last whorl, and in the erratically placed varices with blunt, broad, open spines. One form of ©. edwardsi (valid taxon?) from Vigo (Spain) has a completely white aperture, with a white shell, but it differs strongly morphologically from ©. inordinata. That shell was 1illustrated by ROLAN (1983: 231) as Ocinebrina cf. nicolai (Monterosato, 1884). From ©. miscowichi (Pallary, 1920), a species occuring off the North-West African coast, ©. inordinata differs in its white aperture, stronger and higher axial sculpture, in its erratically placed varices, and in its fewer, broader spiral cords. Other species of European or West African Ocenebra or Ocinebrina are very different and need not to be compared. Etymology: inordinata (Latin): not arranged, disorderly. Named for the erratically placed varices and axial ridges. 123 APEX 9(4): 119-130, déc. 1994 Genus Bedeva Iredale, 1924 Bedeva paivae (Crosse, 1864) Figs 15-17 Trophon paivae Crosse, 1864: 278. Records: Madeira, Funchal Harbour, 0-10 m, many spécimens. Remarks: Bedeva paivae was originally restricted to Australia, from South Queensland to Shark Bay, West Australia, and in Tasmania (WELLS & BRYCE. 1986). KILBURN & RIPPEY (1982: 91) recorded the species from the eastern Cape Province (South Africa) where it lives in colonies of up to 72 individuals per square meter. Bedeva paivae was also collected alive in the Canary Islands (GOMEZ, 1983) It is obvious that the species was introduced to South Africa, to the Canary Islands, and now to the Archipelago of Madeira in the hull of ships (oil tankers, merchant ships...), as already noted in KILBURN & RiPPEY (1982) and in GOMEZ (1983). The presence of B. paivae in the Archipelago of Madeira was never reported before. It is apparently very common but not outside of the harbour. The first known specimen was collected by Nicolas Vassart. The classification of Bedeva in the Ocenebrinae is tentative and based on the observation of radular morphology (Vokes, pers. comm.). Subfamily TROPHONINAE Genus Trophonopsis Bucquoy & Dautzenberg, 1882 Trophonopsis richardi (Dautzenberg & Fischer, 1896) Trophon richardi Dautzenberg & Fischer, 1896: 438, pl. 18, fig. 6. Records: NORDSIECK & GARCIA- TALAVERA, 1979: Madeira. Remarks: No recent record is known for this species. Moreover, the real identity of the recorded species in NORDSIECK & GARCIA- TALAVERA (1979: 132) is very doubtful because of the confusion existing between 7rophonopsis muricatus (Montagu, 1803) and 7rophonopsis richardi (Dautzenberg & Fischer, 1896) (= Trophon droueti Dautzenberg, 1889) (HOUART, 1981: 33). The presence of 7. droueti or T. muricatus in the Archipelago thus remains doubtful. 124 The Municidae from Madeira Subfamily TYPHINAE Genus Typhis Montfort, 1810 Subgenus Typhinellus Jousseaume, 1880 Typhis (Typhinellus) fistulatus (Risso, 1826) Murex fistulatus Risso, 1826: 191 (not Muricites fistulatus Schlotheim, 1820 = Lyrotyphis). Typhis sowerbii Broderip in Broderip & Sowerby 1833: 178. Records: NORDSIECK & GARCIA- TALAVERA, 1979: Madeira (as Typhis sowerbyi Broderip, 1833). Remarks: The record of this species in Madeira is probably based on a misidentification. The specimen in the possession of Garcia-Talavera (from the Canary Islands) is an example of 7yphis (Typhina) belcheri Broderip, 1833. Moreover, Garcia- Talavera (pers. comm.) has no material from Madeira. The presence of Zyphis fistulatus in the Archipelago of Madeira is rejected here. Subfamily RAPANINAE (ex Thaidinae) Genus Stramonita Schumacher, 1817 Stramonita haemastoma (Linné, 1767) Fig. 14 Buccinum haemastoma Linné, 1767: 1202. Purpura haemastoma -WATSON, 1897: 306. Purpura haemastoma -NOBRE, 1937: 30. Thais haemastoma -NORDSIECK & GARCIA- TALAVERA, 1979: 132. Records: Many live and dead taken specimens from the Desertas Is and the Selvagens Is (MMF),;, WATSON, 1897: Everywhere, very common; NOBRE, 1937: Porto da Cruz, Funchal, Porto Santo, Zimbral; NORDSIECK & GARCIA-TALAVERA, 1979: Madeira. Remarks: Sramonita haemastoma 1 variable and is recorded from both the eastern and western Atlantic. It also occurs all along the West African coast, as well as in the Mediterranean Sea. Many synonymous names and a few subspecies have been proposed for this species. HOUART & ABREU HOUART & ABREU Acknowledgements. We wish to express our sincere thanks to F. Garcia-Talavera (Santa Cruz De Tenerife, Canary Is) and Johan Verstraeten (Oostende, Belgium), for lending material from their collections, and to EH. Vokes (Tulane University, New Orleans, U.S.A.) for critical reading of the manuscript. REFERENCES BROCCHI, G.B., 1814. Conchiologia fossile subappenina, con osservazioni geologiche sugli Appenini e sul suolo adjacente, Milano, 2 vols: 1-712. CROSSE, H, 1864. Description nouvelles de l'Australie Conchyl. 12: 275-279. BRODERIP, W.J. & G.B. SOWERBY, 1833 (1832). Characters of new species of Mollusca and Conchifera collected by Mr. Cuming. Proc. Zool. Soc. London 2: 173-179 (published 14 Jan. 1833), 194-202 (published 13 Mar. 1833). DALL, W.H., 1889. Reports on the results of dredgings, under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U.S. Coast Survey Steamer "Blake". 29. Report on the Mollusca. 2, Gastropoda & Scaphopoda. Bull. Mus. Comp. Zool. 18: 1-492. d'espèces méridionale. J. DAUTZENBERG, P., & H. FISCHER, 1896. Dragages effectués par l'Hirondelle et par la Princesse-Alice: 1. Mollusques Gastéropodes. Mémoires de la Société Zoologique de France, 9: 395-498. GOMEZ, R., 1983. Primera cita para el Atlantico (Islas Canarias) de Bedeva paivae (Crosse, 1864). Boll. Malac. 19 (9-12): 249-252. HOUART, R., 1981. Révision des Trophoninae d'Europe (Gastropoda: Muricidae). /nf. Soc. Belge de Malac. 9 (1-2): 1-70. KILBURN, R. & E. RiPPEY, 1982. Seashells of southern Africa, Johannesburg, Macmillan: 1- 249. LAMARCK, J.B.P.A.. de M. de, 1822. Histoire naturelle des animaux sans vertèbres, vol. 7, Paris: 1-232. LINNE, C. von, 1758. Systema naturae per regna trià natura. editio decima, reformata. Stockholm, vol. 1, Regnum animale: 1-824. The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. LINNE, C. von, 1767. Systema naturae per regna tria natura. 12th ed. 1(2): 533-1327 + index. MOOLENBEEK, R.G. & M.J. FABER, 1987a. The Macaronesian species of the genus Manzonia (Gastropoda: Rissoidae) Part I. De Kreukel 23 (1): 1-16. MOOLENBEEK, R.G. & M.J. FABER, 1987b. The Macaronesian species of the genus Manzonia (Gastropoda: Rissoidae) Part II. De Kreukel 23 (2-3): 23-31. MOOLENBEEK, R.G. & M.J. FABER, 1987b. The Macaronesian species of the genus Manzonia (Gastropoda: Rissoidae) Part III. De Kreukel 23 (10): 166-179. MOOLENBEEK, R.G. & H.J. HOENSELAAR, 1989. The genus A/vania in the Canary Islands and Madeira (Mollusca: Gastropoda) Part I. Bull. Zool. Mus. 11 (27): 215-227. NOBRE, À. 1937. Moluscos testaceos marinhos do arquipelago da Madeira. NORDSIECK, F. & F. GARCIA-TALAVERA, 1979. Moluscos marinos de Canarias y Madera. Aula de Cultura de Tenerife: 1-208. PAYRAUDEAU, B.C., 1826. Catalogue descriptif et méthodique des Annélides et des Mollusques de l'île de Corse. Paris: 1-218. POIRIER, J.. 1883. Révision des Murex du Muséum. MVouvelles Archives du Muséum d'Histoire Naturelle, Paris. Ser. 2, 5: 13-128. RIsso, A, 1826. Histoire naturelle des principales productions de l'Europe Méridionale et particulièrement de celles des environs de Nice et des Alpes Maritimes. Paris. Vol. 4: 1-vi, 1-439. ROLAN, E., 1983. Moluscos de la ria de Vigo, 1. Gasteropodos. Privately published, Vigo: 1- 383. WATSON, R.B., 1897. On the marine Mollusca of Madeira; with descriptions of thirty-five new species, and an index-list of all the known sea- dwelling species of that island. J. Linn. Soc. Lond., Zool. 26: 233-329. WELLS. F.E. & C.W. BRYCE, 1985. Seashells of Western Australia. Perth. Western Australian Museum: 1-207. 125 APEX 9(4): 119-130, déc. 1994. The Muricidae from Madeira HOUART & ABREU Figures 1-8 (opposite page). 1-2. Hexaplex trunculus (Linné, 1758), Madeira, Garajau, 70, MMF. 1. 83.3 mm. 2. 41.2 mm. 3-4. Muricopsis (Murexsul) aradasii (Monterosato, 1883). 3. Murex medicago Watson, 1897. Holotype BMNH 1911.7.17.3, 13.5 mm. 4. Madeira, Caniçal, MMF JSL-3004, 13 mm. 5-6. Cytharomorula grayi (Dall, 1889). 5. Trophon lowei Watson, 1897. Holotype BMNH 1911.7.17.2, 19 mm. 6. Madeira (no other locality data), MMF, 19.5 mm. 7. Ocenebra erinaceus (Linné, 1758), Madeira (no other locality data), MMF 24702, 44.1 mm. 8. Ocenebra (Ocinebrina) aciculata (Lamarck, 1822), Madeira Arch., Desertas, MMF 25160, 10.8 mm. 126 HOUART & ABREU The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. 127 APEX 9(4) 119-130, déc. 1994. The Muricidae from Madeira HOUART & ABREU Figures 9-14 (opposite page). 9-10. Ocenebra (Ocinebrina) edwardsi (Payraudeau, 1826), Madeira (No other locality data), MMF 24631. 9: 17 mm. 10: 14 mm. 11-13. Ocenebra (Ocinebrina) inordinata n.sp. 11. Holotype MMF 25429, 19.2 mm. 12-13. Paratype coll. J. Verstraeten, 21 mm. 14. Stramonita haemastoma (Linné, 1767), Madeira Arch., Selvagens, MMF 14332, 60.3 mm. 128 HOUART & ABREU The Muricidae from Madeira APEX 9(4): 119-130, déc. 1994. # "4 NT À 129 APEX 9(4): 119-130, déc. 1994. The Muricidae from Madeira HOUART & ABREU Figures 15-17. 15-17. Bedeva paivae (Crosse, 1864). 15. Syntype BMNH 1870.10.26.70, 24 mm. 16-17. Funchal Harbour, Funchal, Madeira Arch., RH, 16: 19.2 mm; 17: 16.8 mm. 130 TURSCH The scale of sympatry APEX 9(4): 131-142, déc. 1994. The scale of sympatry in the genus Oliva (Gastropoda, Olividae) () Bernard TURSCH Laboratoire de Bio-Ecologie, Faculté des Sciences, Université Libre de Bruxelles, 50 av. F.D.Roosevelt, B-1050 Brussels, Belgium. KEY WORDS. Sympatry, syntopy, variation, taxonomy, Mollusca, Gastropoda, Oliva. ABSTRACT. Sharp morphological discontinuities have been evidenced between conspecific, close neighbouring Oliva populations in Hansa Bay (Papua New Guinea). Taxonomic implications are discussed. The broad notion of sympatry usually applied in the genus Oliva should be replaced by that of syntopy. RESUME. De nettes discontinuités morphologiques ont été mises en évidence entre des populations conspécifiques très voisines d'Oliva à Hansa Bay (Papouasie- Nouvelle- Guinée). Les implications taxonomiques sont discutées. La notion large de sympatrie habituellement appliquée au genre Oliva devrait être remplacée par celle de syntopie. 1. INTRODUCTION Much more effort has been invested on the nomenclature of Oliva species than on their natural history. The taxonomy of this genus (like that of the immense majority of marine molluscs) is therefore still at the stage of the morphospecies approach, based upon the dem- onstration of gaps in the distribution of shell characters. This method is obviously open to er- rors. Some are inherent to the method itself (amongst others, it is likely to overlook sibling species). Other errors can result from the quality of data (these vary from the statement of per- sonal impressions to multivariate analysis, de- pending on the requirements of authors). I wish to report and discuss here some data which un- derline another, frequently overlooked, possible source of error: unwarranted assumptions on sympatry. Data on sympatry are indeed crucial be- cause they largely determine the reasoning of the morphospecies taxonomist. On the one hand, if two sympatric populations are separated by a constant morphological gap, one can infer that interbreeding does not take place and that one deals with two distinct species. In the ab- sence of potentially misleading factors (such as sexual dimorphism, for instance) this procedure is quite straightforward. On the other hand, one can reasonably combine allopatric, morphologi- çally distinct populations into one species 1f one establishes that their discriminating characters are bridged by other populations, forming one morphological continuum. Yn this case, one cannot exclude potential interbreeding. Decisions on allopatric populations are of course more delicate (see MAYR & ASHLOCK, 1991) and their reliability will depend very much on the quality of the data. Suspicion of sympatry-related errors arose during a study on the "Oliva oliva complex" (TURSCH ef al., 1992). The existence of separate species was detected at the local level by the observation of total morphological gaps between sympatric populations. On a global scale, the populations of each species form a morphologi- cal continuum. As one could expect, extreme forms of the same species (sometimes separated by hundreds or thousands of miles) often differ to the point of complete morphological separa- tion. But these differences were always bridged in intermediate, allopatric populations. In con- trast, each conspecific continuum is separated from the others by demonstrable gaps reflecting the boundaries of intraspecific variability. Although the overall picture was quite satis- factory, unexpected facts were observed in two localities (MIsSA, 1991; TURSCH et al., 1992). In Phuket (Thailand) one finds two morphs that are locally entirely separable, while being obvi- ously bridged by series of allopatric intergrades. An entirely similar situation occurs in Carita Beach (Java). These observations were rather disquieting, all the more so because similar situations were soon detected in several other (°) This is paper 21 in the series Studies on Olividae and Laing Island Biological Station contribution n° 286. 131 APEX 9(4): 131-142, déc. 1994. Oliva species, as will be seen here below. On the one hand, membership of the same morpho- logical continuum is a very strong argument for conspecificity. On the other hand, sympatric forms separated by an absolute morphological gap just have to be distinct species, at least in groups such as Oliva where sexual dimorphism and allometry are absent or negligible. This apparent contradiction could be ex- plained by invoking circular overlap, a situation in which the extremities of a chain of inter- breeding local races come into contact and where the extreme forms, now sympatric, can- not anymore interbreed. À much simpler expla- nation would be that there is something wrong in the way we usually apply the concept of sym- patry to Oliva (and possibly many other marine molluscs). Sympatry has been conceived broadly to include "populations the individuals of which are within cruising range of each other during the breeding season, even though the habitats in which they occur do not overlap in space" (CAIN, 1953; MAYR, 1963; MAYR & ASHLOCK, 1991). We know very little on cruising ranges, yet the common practice of marine mollusc tax- onomy 1s to consider as sympatric two popula- tions separated by a few miles, living in the same depth range and on the same type of sediment. The hypothesis of a possible error in our application of the concept of sympatry to the genus Oliva could be tested by studying intras- pecific variation over very short distances. A suitable location for such a study was provided by Hansa Bay, on the North coast of Papua New Guinea. It is a small semicircular bay (roughly 10 km in diameter) located in Madang Prov- ince, near the mouth of the Ramu River, about 110 nautical miles West of Madang. A rough sketch of Hansa Bay is given in Fig. 1. Laing Island (4°10'30"S-144°52'"20"E) lying roughly at the middle of the bay is a raised coral reef, covered with vegetation and separated from the mainland by depths of 45-50 m, with a muddy bottom. Climatic and hydrological data are given by BOUILLON et al. (1986). Detailed envi- ronmental data can be found in CLAEREBOUDT (1989). The sediments in which Oliva are found in Hansa Bay have been analysed by VAN OSSELAER (1992) (see VAN OSSELAER et al.,1994). All the coast is lined with a long, black sand beach, with the exception of a small stretch at Boro Beach, formed of white, coarse coral sand. 132 The scale of sympatry The Oliva species of Hansa Bay have been under survey for nearly 20 years, since the establishment of King Leopold III Biological Station on Laing Island in 1974. 29 species have been found so far and nearly all of them have been collected in adequate numbers, with accurate locality data. Many of these species have distinct micropopulations, separated by short distances. The differences observed between these populations concern not only the variations of colour pattern observed in cryptic, polytopic species (VAN OSSELAER et al., 1994) but also, as will be shown here, the morphology of the shell. In order to avoid repetition, only three cases (ranging from a rather subtle discontinuity to a large, obvious morphological gap) will be described here, although many additional examples could be given. 2. MATERIAL AND METHODS 2.1. Collection Oliva specimens have been obtained mostly by SCUBA diving (day and night dives) but a variety of other methods have also been used. These include dredging (using a small rectangu- lar steel mesh dredge with an opening of 60 x 22 cm), trawling (with a small mesh 3 m rigid frame trawl), snorkelling in shallow waters, or beach collecting at the turn of the low tide. Baiting and trapping have often been used. Quantitative quadrats and transects have also been effected (VAN OSSELAER, 1992). When diving, small rigid steel mesh "hand dredges" (in the shape of a dustpan, about 20 x 30 cm) have been especially productive. Special care has always been taken to avoid overcollecting. Specimens of all 29 Oliva species present in Hansa Bay have been kept and observed in aquaria, sometimes for several months. 2.2. Nomenclature Oliva amethystina (Rôding, 1798) was formerly part of O. annulata (Gmelin, 1791), a nomen dubium encompassing at least two distinct species (TURSCH ef al, 1986). The nomenclatural puzzle of the "©. oliva complex" is not yet completely solved (see TURSCH ef al., 1992), so for the time being, the name ©. smithi Bridgmann, 1906 will be used for one of the species discussed here. This name is probably a junior synonym but has been selected because it is based upon indisputable type material. TURSCH TURSCH The scale of sympatry APEX 9(4): 131-142, déc. 1994. CONDOR POINT SISIMANGUM village NUBIA station 0 1 2 + ——— km Fig. 1. Hansa Bay. 2.3. Material measured BT-numbers refer to specimens in the author's collection, DG- numbers to the collection of Dr. Dietmar Greifeneder (Schwenningen) and JS- numbers to that of Dr. Jacques Senders (Brussels). Oliva amethystina (Rôding, 1798). PAPUA NEW GUINEA, HANSA BAY: - DURANGIT REEF, 6m: specimens BT-7002, BT- 7003, BT-7004, BT-7006 to BT-7001; - "DaAviT's WRECK" (near Awar river), 6 m: specimens BT-6779 to BT-6786, - "MaAST WRECK" (near Awar river), 5 m: specimens BT- 7178 to BT-7182. OTHER LOCALITIES: SOLOMON Is. specimens BT-2454, BT-3512, BT-3516, DG- 2521/5, DG-2521/8, DG-2523/6, DG-2524/3. Oliva smithi Bridgmann, 1906. PAPUA NEW GUINEA, HANSA BAY: - LAING ISLAND West coast (lagoon), 0.5-1 m: specimens BT-2174 to BT-2183. - OFF BORO BEACH, 6-8 m: specimens BT-2164 to BT-2173. OTHER LOCALITIES: AUSTRALIA (North Queensland): specimens BT-5767, BT-5807, BT-5808, BT-6122 to BT-6126, BT-6128, BT- 6130; INDONESIA (Ceram IL): specimens BT- 167, BT-169, BT-296, BT-298; PHILIPPINES (Cebu): specimens BT-1312, BT-4999 to BT- SUOS NN BT-5789 MBT-5791 "10 WBT-5793; PHILIPPINES (Pamilacan): specimens BT-6277 to BT-6279, BT-6281, BT-6284. Oliva oliva (Linnaeus, 1758). PAPUA NEW GUINEA, HANSA BAY: - BORO BEACH, low tide: specimens BT-2154 to BT-2163 (all of very light ground colour, here- under designated as "white"). - SISIMANGUM BEACH, low tide: specimens BT-2154 to 2158 (all of very light ground colour, hereunder des- ignated as "white") and specimens BT-2149 to BT-2153 (all of very dark ground colour, here- under designated as "black"). 133 APEX 9(4) 131-142, déc. 1994. The scale of sympatry TURSCH OTHER LOCALITIES: PAPUA NEW GUINEA (Samarai, Milne Bay): specimens BT-5245 to BT-5247, BT-5251, BT-5254, BT-5259, BT- 5263, BT-5269, BT-5272, BT-5273; PHiL- IPPINES (Zamboanga): specimens BT-4589 to BT-4593, BT-5700, BT-5779 to BT-5784; THAILAND (Patong Beach, Phuket): specimens BT-4768, BT-6142, BT-6149, BT-6154, JS-035, JS-037, JS-040, JS-176, JS-186. In addition to the measured material, sev- eral hundred Hansa Bay specimens of the most common populations of these species have been visually checked, in order to verify the constancy of the discriminating characters described below. 2.4. Tests of conspecificity. Special care was taken to verify that the populations presented here as conspecific do really belong to the same species. All are common, widely distributed Oliva species for which abundant comparison material is available. In each case it was established (by principal factor analysis and/or discriminant factor analysis) that these populations are parts of a much larger morphological continuum, covering the whole distribution area of the species. Within a species continuum, no population (or group of populations) could be distinguished from all the others by any of the many metric shell variables that were tested (alone or in combination). Examples of the applications of these methods to O/iva have already been reported (for instance in TURSCH ef al., 1992) and, for the sake of brevity, details of these lengthy procedures will not be reported here. For every case, it was demonstrated (by scatter diagrams) that the morphological gaps distinguishing the Hansa Bay populations are bridged in other, allopatric populations. For each species, only one scatter diagram will be illustrated here. One should note that the "bridging populations" do not necessarily occur in Hansa Bay. 2.5. Measurements The following measurements have been used in this paper: H, L, D (Fig. 2) and R (Fig. 3) are teleoconch measurements defined in TURSCH & GERMAIN (1985). Pat15, pat16, pat17 and pat18 (Fig. 4) are protoconch measurements defined in TURSCH & GERMAIN (1987). 134 y *_ Fig. 2. Sketch of the measurements H, D and L. Fig. 4. Sketch of the protoconch measurements pat15, pat16, pati7 and pat18. TURSCH The scale of sympatry APEX 9(4): 131-142, déc. 1994. Plate 1. Figs. 1-2: Oliva amethystina (Rôding, 1798). Fig. 1. Typical specimen (BT-6781) from "Davit's wrecK", 6 m. Fig. 2. Typical specimen (BT-7009) from Durangit Reef, 6 m. Figs. 3-4: Oliva smithi Bridgman, 1906. Fig. 3. Typical specimen (BT-7107) from Laing |. lagoon, 0.5-1 m. Fig. 4. Typical specimen (BT-6543) from Boro Beach, 6-8 m. Figs. 5-6: Protoconchs of Oliva smithi. Note the difference in convexity of the penultimate protoconch whorl (arrows) and the fusion of the two last protoconch whorls in specimen BT-7107 (fig. 5). Fig. 5. Protoconch of specimen BT-6228 from Laing Il. lagoon, 0.5-1 m. - Fig. 6. Protoconch of specimen BT-6543 from Boro Beach, 6-8 m. Figs. 7-8: Oliva oliva (Linnaeus, 1758). Fig. 7. Typical specimen (BT-7022) from Sisimangum Beach, low tide. Fig. 8. Typical specimen (BT-6573) from Boro Beach, low tide. 195 APEX 9(4): 131-142, déc. 1994. 3. OBSERVATIONS AND RESULTS 3.1. Oliva amethystina (Rôding, 1798). In Hansa Bay, this common species lives mostly between -1 and -10 m, exclusively in coral sand, in proximity to live coral (where it is easily mistaken for a dead Acropora coral fragment). Two populations (Plate 1, figs. 1-2) have been compared: one living on the top of Durangit Reef and another living around "Davit's wreck", two localities distant of only 2.8 km and separated by a bottom of very fine dark mud, in 40-48 m, extending over about 1 kmMany specimens have been collected at -6 m, in coarse, white coral sand, on the top of the large Durangit Reef. This biotope is often exposed to heavy water motion (high waves and swift current), so Oliva tracks are immediately erased and specimens have to be collected with a small hand dredge. Adult specimens are mostly found around the rocky ledges of the sand pockets, while juveniles are mostly found in the sand ripples. The population of ©. amethystina at the top of Durangit Reef is very similar to (and could not be separated from) that of the Laing Island reef (excepted that the percentage of large, adult shells is much higher at Durangit). A much smaller number of specimens were collected at -6 m, around a Japanese wreck called "Davit's wreck" because its davits could be seen above water until a few years ago. The specimens lived in a mixture of fine and coarse coral sand( including dark terrigeneous material). In this biotope where the water 1s mostly quiet, all our juvenile amethystina were found (by hand dredging) only in the white coral sand recently deposited just along the hull, whereas some adults were found by their tracks in the darker sand further away from the wreck. The wreck is quite small and we have been careful not to overcollect this little biotope. The shells are quite characteristic and no matching specimens have been found in or around Hansa Bay despite extensive exploration over 20 years, with the exception of a few rather similar specimens found around "Mast wreck", another sunken ship lying close to "Davit's wreck". The two wrecks are separated by about 200 m of hard, fine dark sediment, in 5-7 m depth. AII available specimens (8) from "Davit's wreck" were compared to an equivalent number of randomly selected specimens from Durangit. The two samples differ by protoconch as well as by teleoconch characters and are completely separable, as demonstrated (Fig. 5) by a scatter diagram of pat17/pat18 vs. D/L (pat17/pat18 136 The scale of sympatry patl/7/pat18 0.72 0.68 0.64 0.60 0.56 L] LJ Ô 0.58 0.62 0.66 Fig. 5. Comparison of populations of Oliva amethystina. Scatter diagram of D/L vs. pat1/7/pat18. Minimum convex polygons. The populations from Hansa Bay {thick contour lines - black circles: shells from Durangit Reef - black squares: shells from "Davit's wreck") are separated by an obvious moprphological gap. This gap is bridged by other conspecific populations (thin contour lines - white circles: shells from "Mast wrecK" - white squares: shells from Solomon Is.). roughly reflects the conical angle of the penultimate protoconch whorl). All specimens collected at Durangit are squat, heavy, with a tendency to bulge on the body whorl (Plate 1, fig. 2), all have a light coloured background. AIl specimens from "Davit's wreck" are more slender, have a darker background (Plate 1, fig. 1) and the surface of their protoconch is mostly corroded. This is also the case for the "Mast wreck" specimens. Fig.5 also shows that the morphological gap separating the Durangit Reef and the "Davit's wreck" populations 1s bridged in other, allopatric populations of the same species. It should be noted that "Davit's wreck" was sunk during World War Il Like the other wrecks in the bay, it is now overgrown with a luxuriant reef, the decay of which surrounds the hull with coral sand, the exclusive habitat of ©. amethystina. This small pocket of white coral sand is isolated in a plain of fine black sedi- ment. If one estimates the time required for the TURSCH TURSCH growth and the subsequent decay of the coral it is unlikely that the "Davit's wreck" population of O. amethystina is more than 30 years old. 3.2. Oliva smithi Bridgman, 1906. Two distinct populations of this common species are found in Hansa Bay (Plate 1, figs. 3- 4). One is widely spread along the coast, in depths of 2-14 m (generally 5-9 m), on sediments ranging from fine, dark terrigeneous material (off Awar, in usually quiet waters) to fine, white coral sand (off Boro Beach, where the sediment is in nearly constant motion, due to heavy swell). The colour pattern of the shells is very variable, mostly matching the colour of the sediment. The other population has a very restricted distribution in the lagoon of Laing Island, in shallow water (0.5-1 m), coarse coral sand, occasional moderate wave action, and is quite constant in colour pattern. It is a recent (maybe accidental) introduction, not found before 1992 although the lagoon has been the object of regular, intensive search for 20 years. Furthermore, the lagoon population happens to live in the place where our native co-workers (all experienced Oliva watchers) daily wash their dishes, and it is unlikely that the shells would have escaped their trained eye for very long. The lagoon beach appears quite uniform, but during a thorough search in early 1993, ©. smithi was found only on a stretch of about 50 meters. Only a few shells were gathered, in order not to upset this small population. A sample of ©. smithi from Laing Island lagoon was compared to a sample from off Boro Beach. These localities are roughly 3.8 km apart and are separated by a bottom of very fine dark mud, at 40-48 m, extending over about 1.5 km. The two samples are completely separated (Fig. 6) on a scatter diagram of D/L vs. pat15/pat16 (pat15/pat16 is an expression of the convexity of the penultimate nuclear whorl). Specimens from Laing Island reach a larger size than the Boro Beach shells (H,n2% 22.8 mm vs. 15.8 mm) and their body whorl is less cylindrical (see Plate 1, figs. 3-4). The protoconchs of the two samples, albeit similar, show constant differences (see Plate 1, figs. 5-6). In the Laing Island specimens, the two last whorls of the protoconch are fused (the suture 1s covered by a thin, transparent layer of enamel) and the profile of the penultimate nuclear whorl is quite flat. In the Boro Beach specimens, the two last whorls of the protoconch are distinct and the profile of the penultimate nuclear whorl is The scale of sympatry APEX 9(4): 131-142, déc. 1994. convex. Fig.6 also shows that the morphological gap separating the Laing Island and the Boro populations is bridged in other, allopatric populations of the same species. 3.3. Oliva oliva (Linnaeus, 1758). In Hansa Bay, this abundant species is restricted to the low water level on sandy beaches exposed to frequent surf. It is polytopic and cryptic (VAN OSSELAER ef al., 1994), the background colour of the shells ranging from very pale to very dark (hereafter called "white" and "black" shells). One small population lives on white, coarse coral sand at Boro Beach, where heavy swell is generally prevalent. Specimens from Boro (see Plate 1, fig. 8) reach a moderate size (H,nax 33.15 mm). All 42 specimens collected are "white" and have an elongated spire. The aper- ture is short and consistently reddish brown. The protoconch of all specimens is severely eroded. Another population (this one very large) extends all the way from the northern tip of the bay to the mouth of the Sakula river, on fine, black volcanic sand, with occasional, moderate swell. Specimens from Sisimangum (see Plate 1, fig. 7) reach a larger size (H,n,% 42.31 mm). On nearly one thousand specimens observed, 76% are "black", 15 % are "white" and 9 % do not fit into these categories. The spire is short, the aperture long and mostly dark purple. Most spécimens have an intact protoconch. A sample of 10 specimens (all "white") from Boro Beach was compared to a sample of 10 specimens (5 "white", 5 "black", all protoconchs intact) from Sisimangum. These localities are distant of roughly 12 km along the coast but the Sisimangum population reaches the Sakula river and the two populations actually come within 1.5 km of each other. The only physical obstacles separating these populations are a small rocky point between Boro and Kalagima and the mouth of the Sakula River. The two samples are entirely distinct, separated by a large morphological gap, as shown (Fig. 7) on a scatter diagram of H/L vs. R/D. Within the Sisimangum sample, "white" and "black" specimens are not separated; shell morphology seems unrelated to colour. Fig. 7 also shows that the morphological gap separating the Sisimangum Beach and the Boro Beach populations is bridged in other, allopatric populations of the same species. 137 APEX 9(4): 131-142, déc. 1994. pat15/patlé R/D 138 0.52 0.56 The scale of sympatry TURSCH 0.60 Fig. 6. Comparison of populations of Oliva smithi. Scatter diagram of D/L vs. pat15/pat16. Minimum convex polygons. The populations from Hansa Bay (thick contour lines - black circles: shells from Boro - black squares: shells from Laing Island lagoon) are separated by an obvious moprphological gap. This gap is bridged by other conspecific populations (thin contour lines - white circles: shells from Australia, North Queensland - white squares: shells from Philippines, Cebu - white circles, centered: shells from Indonesia, Ceram - white lozenges: shells from Philippines, Pamilacan). Fig. 7. Comparison of populations of Oliva oliva. Scatter diagram of D/L vs. pat15/pat16. Minimum convex polygons. The populations from Hansa Bay (thick contour lines - black circles: shells from Boro Beach - black squares: black shells from Sisimangum Beach - thick white squares: white shells from Sisimangum Beach) are separated by an obvious moprphological gap. This gap is bridged by other conspecific populations (thin contour lines - white circles: shells from Philippines, Zamboanga - white squares: shells from Papua New Guinea, Samarai - white lozenges: shells from Thailand, Phuket). TURSCH It should be noted that the characters of these two populations have been stable for 20 years (judging from specimens in our collection) and probably for much longer a time, judging from specimens in ancient leg ornaments, still used in traditional sing-sing ceremonies by the local Dawar people. 4. INTERPRETATION In short, it has been shown that: a. Close neighbouring populations of the same Oliva species often present notable differences. These differences reach complete morphological separation in the examples presented here, but it should be stressed that this point is not essential. Some overlap in the distribution of population characters would not affect the conclusions here under. Many Oliva variants exhibit a remarkable fidelity to their locality, to the point that an experienced collector can often guess the precise origin of a given specimen. b. The observed divergence of micropopulation characters can occur fast. It can be estimated to less than 30 years for ©. amethystina (see section 3.1, last $) and to probably a few years for ©. smithi (see section 3.2, $ 2). In contrast, all the established populations regularly sampled for a long time have been stable in their characters at least over a period of 20 years and probably for much longer (see section 3.3, last $). The observed differences in shell morphol- ogy could stem both from phenotypic plasticity and genetic isolation. On the one hand, envi- ronmental effects are probably an important contributor to colour variation, as 1llustrated by the fact that 24 of the 29 Oliva species encoun- tered in Hansa Bay are cryptic to some degree, a phenomenon suggesting intense pressure from visual predator(s) (this 1s discussed in VAN OSSELAER ef al., 1994). Considerable predation has indeed been demonstrated in the case of ©. oliva, where nearly every specimen of the Sisimangum population bears the scars of at least one unsuccessful attack (study in pro- gress). No data for linking Oliva shell morphol- ogy to environmental effects are yet available. On the other hand, genetic isolation 1s strongly suggested by the frequent observation of differences in the protoconch (e.g. in the case of Q. smithi). The protoconch of Oliva species is produced inside the egg capsule (OLSSON & CROVO, 1968; TURSCH, 1991), where it 1s shielded against direct environmental The scale of sympatry APEX 9(4): 131-142, déc. 1994. influences. Morphological gaps at the proto- conch level are thus likely to be of genetic ori- gin. Another, indirect argument stems from the very rapid changes observed for the small population of ©. smithi in the lagoon of Laing Island. Such a situation immediately evokes genetic drift, a phenomenon requiring genetic isolation. One more indirect argument can be found in the morphological stability observed in es- tablished Oliva populations. This stability sharply contrasts with the rapid changes occur- ring in the topography of the shallow water biotopes of Hansa Bay. During the last 20 years, the beach at Sisimangum has diminished by some 100 m, the East coast and the northern tip of Laing Island have also considerably re- gressed, while the beach in the southern part of the lagoon has extended. According to the oral tradition of the local Dawar tribesmen, Laing Island was actually linked to the mainland at the time of their ancestors. Elderly people all say that in their youth Laing island had at least the double of its present size. In addition, spo- radic eruptions of the nearby Manam volcano frequently pepper the whole area with large quantities of igneous material. The modifica- tions of the shoreline produce important changes in physical conditions (slope, exposure to waves, etc.), to the point that some of our boat moorings had to be moved over the years. So the morphological stability of most popula- tions shows that many of the discriminant char- acters used by the O/iva taxonomist do not reflect subtle environmental differences but most probably partial genetic isolation. AIl the available data thus indicate that there appears to be severe restrictions on gene flow between many of the Hansa Bay conspeci- fic Oliva populations. Each of these populations could be considered as a topogamodeme, this is a relatively isolated, naturally interbreeding, population (gamodeme) occupying a particular area (LINCOLN ef al., 1982). 5. COMMENTS AND DISCUSSION Although I have no simple solution to offer, it might be worth discussing the mechanism(s) that could explain the partial genetic isolation of the close neighbouring Hansa Bay Oliva populations. Interpopulation gene flow depends upon dispersal and this could take place either at the larval or at the adult stage. Nothing is known about the vagility of Oliva larvae, but the present observations 139 APEX 9(4): 131-142, déc. 1994. suggest that it must generally be low. Indeed, the larval stage lasts several days (OLSSON & CROVO, 1968; TURSCH, 1991) and the season- ally variable currents could easily transport pelagic veligers anywhere in Hansa Bay within hours. It has been shown that the veliger of Olivella verrauxii is not pelagic but swims on the bottom (MARCUS & MARCUS, 1959). I could find no report of Oliva larvae being caught in plankton hauls. Even if the larvae were effectively dispersed, successful immigration could still be severely restricted by the necessity of settling on a suitable substrate. Effective larval dispersal over large distances certainly does take place, as evidenced by the very large distribution of several Oliva species, but it must be quite infrequent. The Oliva populations of Hansa Bay are so closely spaced that immigration could possibly also occur at the adult stage. Very little is known about the actual cruising range of adult Oliva species. They can live several years and do easily reach speeds of 25 cm/min (TURSCH, 1991). Tracks several meters long are frequently observed and an Oliva could conceivably travel many kilometers in its lifetime. One could even imagine that adult Oliva could cross small stretches of moderate depths: they have no pressure-sensitive organ and they could certainly tolerate reduced light (being mainly nocturnal, see VAN OSSELAER & TURSCH, 1993). In addition, they can easily subsist on a wide variety of foods that are completely foreign to them. In theory, ©. oliva could thus travel from Boro to the black Sisimangum beach in a matter of days or weeks. But no specimen of the “Boro type" has ever been collected from Kalagima to Awar. (Conversely, no specimen of the "Sisimangum type" has ever been caught at Boro Beach. It is doubtful that the mouth of the Sakula river could constitute a serious barrier: it is only about 10-20 m wide and is completely closed by a sand bar at low tide during the dry season. The similar (but smaller) Awar river does not divide the Sisimangum population. It is more likely that the obstacle is constituted by the small rocky point separating Boro from Kalagima, where the soft substrate that all olives require is found at 5-6 m, an environment possibly unsuitable for a species restricted to surf beaches. This same rocky point is no obstacle for ©. smithi that often lives in deeper water. In practice, adult Olives seem rather sedentary, as shown by the lagoon population of O. smithi (see section 3.2, $2). One could conjecture they are prevented from moving by 140 The scale of sympatry being very sensitive to minute changes in the nature of the substrate, but this is unlikely because in sediment choice experiments, several species of Oliva have shown only very slight substrate preferences (VAN OSSELAER & TURSCH, 1993). One could also imagine that the micropopulations are kept together by some cohesive force such as chemical attractants (TURSCH, 1991) but no firm data yet support this hypothesis. 6. CONCLUSIONS The Oliva species of Hansa Bay consist in a mosaic of populations, the map of which fairly reflects the discontinuities of the habitat. These populations are not only ecological races; in many cases they also appear to be temporarily isolated by restricted gene flow, even over very short distances. Neighbouring, conspecific populations frequently differ to the point of complete morphological separation. That different populations could be distinguished by some combination of characters is no great discovery and was indeed fully expected (see MAYR, 1969 and FUTUYMA, 1986), even for shells that are collector's favourites. The observations at Hansa Bay just give a somewhat sharper image of the spatial scale, the extent and the tempo of intraspecific variation in the genus Oliva. They also point to the necessity of moderating the current taxonomic approach to this genus. First, the scale of sympatry in Oliva can be much smaller than it is generally considered. The pattern observed in tiny Hansa Bay is certainly not unique and can be expected to occur over the very large distribution areas of many species. In addition to the examples given in the introduction (see TURSCH ef al., 1992), the populations of Oliva oliva I have sampled in two very similar small bays separated by only 5.8 km on the North coast of Hon Lon Island, off Nha Trang (Vietnam) can be separated at a glance. Many other similar examples could be given and it is a safe bet that morphological discontinuities between close-set populations will be commonly observed in any careful field study of Oliva. Special caution is thus necessary in the application of our most reliable tool for taxo- nomic decisions at the morphospecies level: the demonstration of morphological gaps between sympatric populations (see Introduction). When comparing populations from a same, broad lo- cality, data such as "New Guinea, shallow TURSCH TURSCH water" or even "Hansa Bay, sand, 8 m" are now clearly insufficient to demonstrate sympatry. We can be sure that Oliva populations are sympatric only when they actually overlap in space. This can be known only if specimens of both taxa have been observed together (or within a short distance) in the same, continuous microhabitat. Such a relationship could be described by the word "syntopic" in a slightly restricted sense. "Syntopic" has been defined as: "pertaining to populations or species that occupy the same macrohabitat, are observable in close proximity (italics are mine) and could thus interbreed" (LINCOLN et al, 1982). Conversely, in the absence of actual spatial overlap, one could use the word ‘“allotopic" in the somewhat restricted sense of "not observed in close proximity". In the case of Oliva (at least) it is obviously safer to replace our broad criterion of sympatry by that of syntopy. This does not upset any fundamental concept, as it is only a reduction of spatial scale. It will actually facilitate the task of the taxonomist because differences between similar species will generally be more pronounced wherever these species come into contact (the well-known phenomenon of character displacement). Without this reduction of the commonly accepted scale of sympatry, one could be led to create as many Oliva "species" as there are local populations within an arbitrary range of proximity. Many "species" of Oliva have indeed been created upon differences smaller than those observed between some of the Hansa Bay conspecific populations. This solution is unfortunately attractive to some (see the criticism of MAYR, 1969) but is demonstrably wrong. In all the cases studied so far in this laboratory and for which enough data were available (for example the protean "Oliva oliva complex", see TURSCH ef al., 1992), the set of all conspecific populations (considered over the whole distribution range) always form a morphological continuum. The characters discriminating any two given conspecific populations are invariably bridged by at least another population (or by a chain of intergrading populations). This indicates that gene flow, even if restricted, does nevertheless take place. Some could then be tempted to consider each of the morphologically distinct local populations of Oliva as a subspecies in the sense of "local variety". This would lead to obvious nomenclatural excess and the notion of The scale of sympatry APEX 9(4): 131-142, déc. 1994. subspecies (largely a category of convenience) would then lose any meaning. The sensible definition of MAYR (1963) restricts subspecies to "aggregates of populations", on much larger spatial scale. Acknowledgments. I am grateful to the Belgian National Fund for Scientific Research (F.N.R:S), the King Léopold III Fund for Nature Exploration and Conservation, and BIOTEC, S.A. for material support. I am much indebted to my team-mates Sid Johnson, Jean-Marc Ouin, Jean Pierret and Christian Van Osselaer, who have been superlative Oliva watchers as well as fine diving and dredging companions for many years. I thank Dr. Jacques Senders for the loan of comparison specimens and Dr. Dietmar Greifeneder for his usual, constructive criticism. REFERENCES. BOUILLON, J., M. CLAEREBOUDT & G. SEGHERS, 1986. Hydroméduses de la baie de Hansa (Papouasie Nouvelle Guinée): conditions climatiques et hydrologie. /ndo-Malayan Zoology 3: 105-152. CAIN, À. J, 1953. Geography, ecology and coexistence in relation to the biological definition of the species. Evolution 7: 76-83. CLAEREBOUDT, M., 1989. Répartition spatiale et diversité des Scléractiniaires sur un récif corallien de Papouasie Nouvelle Guinée. Thèse de doctorat, Université Libre de Bruxelles. FUTUYMA, D.J., 1986. Evolutionary Biology. Sinance Associates: Sunderland, Mass. 600 pp. LINCOIN, R.J., G.A. BOXSHALL & P.F. CLARK, 1982. A Dictionary of Ecology, Evolution and Systematics. Cambridge University Press: Cambridge, U.K.. 298 pp. MARCUS, E. & E. MARCUS, 1959. On the reproduction of Olivella. Bol. Fac. Filosofia, Ciéncias e Letras, Univ. Sâäo Paulo 232 (Zool. 23): 189-196. MAYR, E., 1963. Animal Species and Evolution. Harvard University Press: Cambridge, Mass. 797 pp. MAYR, E. 1969. Principles of Systematic Zoology. McGraw Hill: New York. 428 pp. MAR, E. & P.D. ASHLOCK, 1991. Principles of Systematic Zoology. McGraw Hill: New York. 475 pp. 141 APEX 9(4): 131-142, déc. 1994. The scale of sympatry MissA, O., 1991. La structure taxonomique de Oliva oliva (Mollusca, Gastropoda). Mémoire de Licence, Université Libre de Bruxelles. OLSSON, À: À & LE'/CROVO 1965: Observations on aquarium specimens of Oliva sayana Ravenel. The Veliger 11(1): 31-32. TURSCH, B., 1991. Studies on Olividae. XIIL. Behaviour of Oliva vidua in aquarium: preliminary observations. Apex 6(1): 1-10. TURSCH, B. & L. GERMAIN, 1985. Studies on Olividae. I. À morphometric approach to the Oliva problem. /ndo-Malayan Zoology 331-352. TURSCH, B.& L. GERMAIN, 1987. Studies on Olividae. V. Five additional protoconch characters for Oliva taxonomy. Apex 2: 59-68. TURSCH, B., L. GERMAIN & D. GREIFENEDER, 1986. Studies on Olividae. IV. Oliva annulata Gmelin, 1791 (of authors) : a confusion of species. /ndo-Malayan Zoology 3: 189-216. TURSCH, B., O. MissA & J. BOUILLON, 1992. Studies on Olividae XIV. The taxonomic structure of Oliva oliva (auct.). Apex 7(1): 3-22. VAN OSSELAER, C., 1992. Contribution à l'étude écologique du genre Oliva (Mollusca, Gastropoda) à Hansa Bay (Papouasie-Nouvelle Guinée). Travail de fin d' Etudes, Université Libre de Bruxelles. 113 pp. VAN OSSELAER, C., J. BOUILLON, J.M. OUN & B. TURSCH, 1994. The distribution of Oliva species and the variations of their colour patterns in Hansa Bay (Papua New Guinea). Apex 9(2/3): 27-46. VAN OSSELAER, C. & B. TURSCH, 1993. Studies on Olividae. XVII. Data on depth of burrowing, motion and substrate choice of some Oliva species. Apex 8(4): 151-158. 142 TURSCH PANIGRAHI & RAUT Cerebral ganglia of Achatina fulica APEX 9(4): 143-147, dec 1994. Noradrenaline and adrenaline in the cerebral ganglia of the giant African land snail Achatina fulica Bowdich. A. PANIGRABHI and S.K. RAUT Department of Zoology, University of Calcutta, 35, Ballygunge Circular Road, Calcutta - 700 019, India KEY WORDS : Achatina fulica, cerebral ganglia, noradrenaline, adrenaline. ABSTRACT. Both histochemical and biochemical studies were conducted to note the state of noradrenaline and adrenaline in the cerebral ganglia of the giant African land snail Achatina fulica Bowdich. The occurrence and distribution of noradrenaline and adrenaline containing granules in the cerebral ganglia have been noted following histochemical studies. It is revealed that the noradrenaline and adrenaline containing granules are distributed to central and peripheral regions of the ganglia respectively. The amount of fluorometrically estimated noradrenaline in the cerebral ganglionic tissues was almost equal to the estimated amount of adrenaline. INTRODUCTION. The occurrence and distribution of monoamines in the cerebral ganglia of molluscs have been the subject of study by many workers. Thus, catecholamine containing cell bodies have been demonstrated in Zimax maximus Linnaeus (OSBRONE and COTTRELL, 1971), Aplysia (GOLDSTEIN, 1984), Helisoma (TRIMBLE et al, 1984), Helicella virgata Da Costa (FRANCHINI et al., 1985) and Zymnaea stagnalis Linnaeus (AUDESIRK, 1985). The distribution of monoamines in the central nervous system of the gastropods, Hermissenda crassicornis Eschscholtz and Achatina fulica Bowdich has also been described by CROLL (1987a, b, 1988). VON EULER (1953) reported the presence of noradrenaline in Octopus. MCCAMAN et al. (1979, 1984) were successful in isolating dopamine, 5-Hydroxytryptamine and N-acetyldopamine from molluscan ganglia. HERNADI et al. (1989) demonstrated the distribution of serotonin (5-HT)-containing neurones in the central nervous system of the land snail Helix pomatia Linnaeus. Distribution of dopamine in the central nervous system of the pond snail Z. stagnalis has also been studied by ELEKES et al. (1991) and WERKMAN et al. (1991). Even. studies on chemical transmission in invertebrate central nervous system have been made by GERSCHENFELD (1973). The present study provides qualitative and quantitative accounts of noradrenaline and adrenaline in the cerebral ganglia of the giant African land snail, À. fulica. MATERIALS AND METHODS. Adult Achatina fulica Bowdich were collected from local vegetable gardens in October, 1987. They were 68.0-71.0 mm in shell length, 32.0-34.5 mm in shell breadth and 36.74-40.65 g in body weight. The snails were anaesthetized by applying chloroform and the cerebral ganglia were quickly dissected out for both the histochemical and biochemical determinations of noradrenaline and adrenaline. The snails were anaesthetized under field conditions and the samples were collected immediately after the same. The following methods were used in the study. Histochemical method. Control. Cerebral ganglia dissected out from five À. fulica were immersed in Carnoy's fixative, embedded in mixture of Beeswax, cerecin and paraffin and 30 UM sections were cut. Chromate-dichromate reaction. For total catecholamines, cerebral ganglia dessected out from five À. fulica were treated with a mixture of 5% potassium dichromate and 5% potassium chromate (10:1) solutions for 48 hours and subsequently transferred to 10 % formalin for 24 hours (HILLARP and HÔKFELT, 1955). Then, the cerebral ganglia were 143 APEX 9(4): 143-147, dec 1994. embedded in a mixture of Beeswax, ceresin and paraffin for a period of 30 minutes and required 30 UM sections were prepared. lodate reaction. To detect noradrenaline, cerebral ganglia dessected out form five À. fulica were fixed in saturated solution of potassium iodate for 24 hours and in turn these were immersed in 10 % formalin for 48 hours (HILLARP and HÔKFELT, 1955). After a 30 minute embedding in a mixture of Beeswax, ceresin and paraffin 30 lL M sections were prepared. Biochemical method. Cerebral ganglia of twelve other snails (4. fulica) were processed for spectrophotofluorometric determinations of noradrenaline and adrenaline. Noradrenaline and adrenaline were extracted and purified following the method of COX and PERHACH (1973) and estimated according to the method of LAVERTY and TAYLOR (1968). The cerebral ganglia of four snails were sorted for each measurement. Three such samples were considered for reading by a Hitachi (Model 650- 10 M) fluorescence spectrophotometer (for details see MAHATA and GHOSH, 1989). The mean of three readings was taken as the final data. Student's 't' test (SNEDECOR and COCHRAN, 1967) was applied for statistical analysis of the data. RESULTS. Histochemistry. Control. No colouration was observed either in the central or peripheral region in the cerebral ganglia of À. fulica Bowdich (Figure 1). Treatment. Following treatment with potassium chromate-dichromate solution, numerous brown and yellow coloured granules were noted in cerebral ganglionic tissues of the snail, À. Jfulica. The yellow coloured granules (indication of presence of noradrenaline) were exclusively confined to central portion of the cerebral ganglia (Figures 2, 4) while the brown coloured granules (indication of presence of adrenaline) were distributed to the peripheral region of the cerebral ganglia (Figures 3,5). The occurrence of yellow granules (indication of presence of noradrenaline), in an irregular fashion in the central portion of the tissues was also noted following potassium iodate treatment (Figure 6). 144 Cerebral ganglia of Achatina fulica Biochemistry. The spectrophotofluorometric determina- tions revealed that the cerebral ganglia of A. Julica contain both noradrenaline 14.09 + SE. 0.80 u1g/g tissue weight) and adrenaline (12.24 + S.E. 0.61ug/g tissue weight). However, such difference in amount is not statistically significant (t.