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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

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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

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

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

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

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VOL. 9 (2-3)

C. Van Osselaer J. Bouillon J.M. Ouin B. Tursch

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SOMMAIRE

Studies on Olividae. 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)

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Dr. Y. Finet

L. Germain

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VAN OSSELAER, BOUILLON, OUIN, TURSCH Studies on Ohvidae. 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

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

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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-

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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

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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.

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SNEATH, P.H. & R.R. SOKAL. 1973. Numerical taxonomy. WH. Freeman and Co, San Francisco, 573 pp.

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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.

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

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

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

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

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

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 ©.

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

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.

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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

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

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.

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.