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ez Uo| AN ISSN 0042-3211



A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California

R. Stohler, Founding Editor

TAA aes} pe



Volume 39 January 2, 1996 Number 1


The taxonomic status of Xerotropis Monterosato, 1892, and redescription of its type species (Pulmonata: Hygromiidae) GIUSEPPE MANGANELLI, LEONARDO FAVILLI, AND FOLCO GIUSTI ........ 1

Homoplastic loss of dart apparatus, phylogeny of the genera, and a phylogenetic taxonomy of the Helminthoglyptidae (Gastropoda: Pulmonata) BAR RM NO DEMIR een se auth A es crete a My cunpa as ubiaed ana & pst sely 18

Lecithotrophic development in Doto amyra (Nudibranchia: Dendronotacea), with a review of developmental mode in the genus PEDREVehn We GODDARD Maan iis Sein dilis ued: alclet ks tee ened Die EY 43

Redescription of Nembrotha megalocera Yonow, 1990 (Gastropoda: Nudibran- chia: Polyceratidae) from the Red Sea J. LL: CERVERA, JC. GARCIA-GOMEZ, AND CG. MEGINA ................ 55

Embryonic and larval development of Spisula solidissima similis (Say, 1822) (Bi- valvia: Mactridae) INANDAT IG. WALKERVAND FRANCIS X. ©>BEIRN ./...........0 00 0re 60

A new species of Abyssochrysos (Gastropoda: Loxonematoidea) from a middle Eocene cold-seep carbonate in the Humptulips Formation, western Wash- ington


CONTENTS Continued

The Veliger (ISSN 0042-3211) is published quarterly in January, April, July, and October by the California Malacozoological Society, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. Second Class postage paid at Berkeley, CA and additional mailing offices. POSTMASTER: Send address changes to The Veliger, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105.


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The Veliger 39(1):1-17 (January 2, 1996)

THE VELIGER © CMS, Inc., 1996

The Taxonomic Status of Xerotropis Monterosato, 1892,

and Redescription of Its ‘Type Species (Pulmonata: Hygromiidae) by


Dipartimento di Biologia Evolutiva, Universita di Siena, Via Mattioli 4, I-53100 Siena Italy

Abstract. The taxonomic status of the nominal genus Xerotropis Monterosato, 1892, was revised by studying its type species Carocolla gargottae Philippi, 1836. Anatomical research showed that the species belongs to the genus Cernuella Schliiter, 1838. Consequently, Xerotropis becomes a junior synonym of Cernuella.

A long taxonomic and nomenclatural controversy exists on the name of the type species of Xerotropis. Carocolla gargottae was considered to be a junior synonym of one of the following nominal species: “Helix rugosa” Chemnitz, 1786, Helix corrugata Gmelin, 1791, Helix scabra Salis Marschlins, 1793, “Helix groyana” Férussac, 1821, and Helix rugosa Lamarck, 1822.

Analysis of the literature showed that two of these names were not valid: “Helix rugosa” Chemnitz, 1786, being part of a polynomial name (Helix scabra et rugosa) and Helix groyana Ferussac, 1821, being a nomen nudum; and that two others were not available for the Sicilian species: H. corrugata Gmelin, 1791, being introduced for the polynomial Helix scabra et rugosa but, despite Pfeiffer’s claim, without objective evidence that the species illustrated by Chemnitz is the same as that of Philippi; and H. scabra Salis Marschlins, 1793, being introduced for material collected at Paestum in southern Italy, but with an original description insufficient for establishing its identity.

Examination of the type material of Helix rugosa Lamarck, 1822, demonstrated that it is the same species as Carocolla gargottae Philippi, 1836. Consequently, Helix rugosa Lamarck, 1822, is the correct

name for the species.


Monterosato (1892:23) established Xerotropis for a group of species of the so-called xerophilae having shells “de- presse a forti rugosita interrotte da un orlo carenale [de- pressed, with marked ribs interrupted by a peripheral carina].” He placed six species in it: Carocolla gargottae Philippi, 1836, from Sicily; Helix joly: Ancey, 1882, from Algeria; Helix prietoc Hidalgo, 1878, from the Balearic Islands; Helix lederer: Pfeiffer, 1857, from Jaffa; Helix milaschewrschi [sic] Retowski, 1886, from Crimea; and He- lix spratt: Pfeiffer, 1846, from the Maltese Islands. In the same year, ‘“‘Hel|[1x]. gargottae” was designated as the type species of Xerotropis in an anonymous comment (presum- ably written by Kobelt, editor of the Nachrichtsblatt) (Ko- belt, 1892:152) on Monterosato’s paper.

Xerotropis was confirmed as a subgenus of Xerophila Held, 1837, by Kobelt (1904), who included Helix cor-

rugata Gmelin, 1791 [= Carocolla gargottae], and H. prae- clara Cafici, 1882, in it. The other species originally in- cluded in Xerotropis by Monterosato were moved to dif- ferent subgenera of Xerophila: Helix lederert and H. spratti to Xeroamanda Monterosato, 1892; H. milachewitchi [sic] and Helix sigensis Kobelt, 1883 [= H. jolyz], to Jacosta Gray, 1821; and H. prietoi to Xeroplexa Monterosato, 1892.

On the other hand, Xerotropis was regarded as a junior synonym of Jacosta by Pilsbry (1895) and Gude & Wood- ward (1921), and as a taxon of unclear and dubious status by Hesse (1926, 1934) and Thiele (1931).

The taxonomy of Xerotropis was subsequently discussed by Zilch (1960), Alzona (1971), and Richardson (1980). Zilch (1960) regarded Xerotropis as being of uncertain status due to a lack of anatomical information. Alzona (1971) regarded Xerotropis as a subgenus of Helicella Fer- ussac, 1821, and assigned four Sicilian species to it. Finally, Richardson (1980) considered it to be a subgenus of the

Page 2

The Veliger, Vol. 39, No. 1

Miocene genus 77tthodomus Pfeffer, 1929, misinterpreting Zilch (1960), who listed two uncertain helicellid genus group taxa, Xerotropis and Xeronexa Monterosato, 1892, using the typographic character used for subgeneric names, after the last genus of the Helicellinae Ihering, 1909, 1.e., Titthodomus!

In view of persisting uncertainty about the status of Xerotropis (as in the case of many other Monterosatian genus group taxa of the xerophilae), we asked some Sicilian colleagues for living specimens of Carocolla gargottae, the nominal species designated at its type species. V. E. Or- lando and I. Sparacio succeeded in providing this material in 1987, so that we were able to perform anatomical study and publish the following revision.


Whole shells were photographed under the optical micro- scope (Wild M5A). All dimensional parameters (shell height, maximum shell diameter, aperture height, and ap- erture diameter) were measured using a calipers.

Living specimens were drowned in water, then fixed and preserved in 75% ethanol buffered with NaHCO,. The bodies were isolated after crushing the shells, and dissected under the optical microscope using very thin, pointed watchmaker’s forceps. Anatomical details were drawn using a Wild camera lucida. The dimensions of anatomical tracts were measured using a millimetric lens on the same microscope.

Radulae were manually extracted from the buccal bulbs, washed in pure 75% ethanol, mounted on copper blocks with electronconductive glue, sputter-coated with gold, and photographed using a Philips 505 SEM.

The material examined is listed as follows: locality, mu- nicipality and province names in parenthesis, UTM ref- erence, collector(s), date, number of specimens in paren- thesis. Locality names and UTM references were accord- ing to the official 1:50,000 scale map of Italy (series M 792, sheets 593 and 626).


In the past century, much controversy arose over the correct name for the species selected as the type species of Xero- tropis.

Many authors considered the name of the type species of Xerotropis, Carocolla gargottae, introduced by Philippi (1836:136-137; pl. 8, fig. 10; type locality: “prope Pan- ormum et Termini”) for a species of western Sicily, to be a junior synonym of one of the following species: “Helix rugosa” Chemnitz, 1786; Helix corrugata Gmelin, 1791; Helix scabra Salis Marschlins, 1793; “Helix groyana” Feér- ussac, 1821; and Helix rugosa Lamarck, 1822.

The matter was raised by L. Pfeiffer (1841:17) who,

on the basis of material in the Lamarck collection, claimed that Philippi’s species was a junior synonym of both Helix rugosa Lamarck, 1822 (p. 90; type locality: “. . . en Italie, sur la route d’Anc6ne a Sinigaglia”) and Helix groyana

Ferussac, 1821 (p. 44, nomen nudum!), the latter intro- duced for material collected by Menard de la Groye “Entre Fiumesino et le case Brusciota, route de Sinigaglia, a An- cone.”

Philippi (1844:110) denied this and alleged that there had been an exchange of material in the Lamarck collec- tion. In fact, the original description of Helix rugosa did not appear to fit his C. gargottae: “‘. . . sed error manifestus collectionis Lamarchianae videtur; verba Lamarckii angle de son dernier tour est un peu prononcé minime quadrant.”

Nevertheless, Pfeiffer (1845:23) persisted in his hy- pothesis. According to him, there was no doubt that Phi- lippi’s C. gargottae was Lamarck’s Helix rugosa (on the basis of his examination of the material in the Lamarck collection and confirmed by the figures of Delessert, 1841- 1842:pl. 26, fig. 4a-d), despite the fact that this evidence was in contrast with Lamarck’s description.

Pfeiffer (1845) also claimed that the authorship of the species should be assigned to Chemnitz (1786) and not to Lamarck (1822). According to him, the species described and illustrated by Chemnitz (1786:152-153; pl. 133, fig. 1208) as “Helix scabra et rugosa,” for which Gmelin (1791: 3623) introduced the binomial name Helix corrugata, cor- responded (at least in part, see below) to that later intro- duced by Lamarck under the name of H. rugosa. It was therefore not necessary to change its name, but to maintain a part of Chemnitz’s original polynomial name.

In the second edition of Martini & Chemnitz’s Conchy- lien-Cabinet, Pfeiffer again published the original descrip- tion and figures by Chemnitz (Pfeiffer, 1846a:51-52, pl. 6, figs. 12, 13). However, since he realized that incongru- ities existed between the description and figure 1208, he wrote: “Bei einer spatern bessern Abbildung werde ich die vollstandige Synonymik und Beschreibung geben” (Pfeif- fer, 1846a:52). Two years later, Pfeiffer (1848a:173-174) gave a new description of the species and referred to new figures previously published by himself (Pfeiffer, 1843, pl. 23, figs. 3, 4). He also reintroduced Lamarck as the author of the species, by. way of explanation offering only: “Ich halte diese Art fur dieselbe Schnecke, welche als H. rugosa Chemn. friher gegeben word ist.” In subsequent years, he curiously reverted to assigning the authorship of the species to Chemnitz (Pfeiffer, 1848b, 1853, 1859, 1868, 1876).

Pfeiffer created further confusion when he cited a dif- ferent nominal species: Helix groviana Ferussac, 1832 (Ex- plication: ij, pl. 46A, fig. 1), which he reported in the synonymy of the Madeiran hygromiid Leptaxis undata (Lowe, 1831) as “Helix groviana (Helicogena) Fér. pr. 276. Hist. t. 46 Af. 1.” (cf. Pfeiffer, 1848b:192; Pfeiffer, 1852: 154). “Helix (Helicogena) groviana”’ does not exist in the Prodromus: the species listed at number 276 is Helix groy- ana, not Helix groviana! Helix groviana is only reported in the “Explication des planches des livraisons XXII, XXIII, XXIV, XXV, XXVI et XXVII” of 1832.

Pfeiffer’s (1845) hypothesis that Helix corrugata Gmelin, 1791, “Helix groyana”’ Ferussac, 1821, Helix rugosa La-

G. Manganelli et al., 1996

Page 3

: My Wo y Y=

Lamac .

= js ey ¢ SL cochlea xr wacata, 72272077 Subriefe Pee

Ware circerzalo, el opertulile,

Figures 1, 2

Two illustrations involved in early taxonomic and nomenclatural history of Cernuella rugosa (Lamarck, 1822). 1. The reference to “Lister Hist. Conchyl. tab. 55. fig. 51” by Chemnitz (1786) and Gmelin (1791) corresponds to a figure in Lister’s Historiae sive Synopsis Methodicae Conchyliorum. The fig. 51 of plate 55 of this book illustrates a Jamaican cyclophorid collected by H. Sloane and described as Cochlea umbilicata minor, subrufa, ore circinnato et operculato. 2. Die rauhe runzelhafte Schnirtelschnecke of Chemnitz (1786: pl. 133, fig. 1208). This shell was assigned by Draparnaud (1805) to his Helix variabilis [= Cernuella virgata (Da Costa, 1778)].

marck, 1822, and Carocolla gargottae Philippi, 1836, were junior synonyms of ‘“‘Helix rugosa’? Chemnitz, was accepted by many authors (Albers, 1850; Reeve, 1854; Benoit, 1862, 1882; Paulucci, 1878; Reuleaux, 1889; Westerlund, 1876, 1889; De Gregorio, 1896; Alzona, 1971).

Albers (1860:111) later preferred to use the name Helix

corrugata Gmelin, 1791 (presumably because Chemnitz’s name was polynomial) and Martens in Albers (1860:115), that of Helix scabra Salis Marschlins, 1793. While some authors followed Albers (1860) in using the name H. cor- rugata (Schaufuss, 1869; Kobelt, 1871, 1904; Tryon, 1887; Pilsbry, 1895), no one followed Martens in using Salis Marschlins’s name.

The matter is obviously complex and requires detailed historical analysis from the beginning.

Chemnitz (1786:152-153; pl. 133, fig. 1208) described “Die rauhe runzelhafte Schnirtelschnecke” as follows “Helix scabra et rugosa, testa umbilicata, carinata, ciner- eoalbida, longitudinaliter obliqua striata, anfractibus sex ro- tundis, apertura lunata, labro intus reflexo et incarnato.”

He referred it to “Lister Hist. Conchyl. tab. 55. fig. 51. Cochlea umbilicata minor, subrufa, ore circinnato et opercu- lato. lamaic.”’ and, in his remarks, listed a few differential features mentioning that “Lister nennet Jamaica als das wahre Vaterland dieser Schneche” and that “Eine fast vollig gleichformige, die auf Otaheite [Haiti] gefunden worden, verwahre ich unter meinen Stdlandischen Con- chylien.”

The bibliography of M. Lister (1638-1711/12) is very complex (Keynes, 1981), and Chemnitz’s reference to “Lister Hist. Conchyl. tab. 55, fig. 51” concerns the His- toriae swe Synopsis Methodicae Conchyliorum published in 1685-1692 and reprinted in 1770. As Pfeiffer (1840:89) was first to realize, Lister’s description (cochlea umbilicata, minor subrufa, ore circinato, et operculato) and the figure (pl. 55, fig. 51) (Figure 1) clearly indicate a cyclophorid land snail. The indications “Iamac.” and “SLOANE” which follow the figure indicate that the material was collected by H. Sloane on the island of Jamaica. When Sloane went to Jamaica in 1687, Lister asked him to collect terrestrial and freshwater snails (Dance, 1986). Descrip- tions of this material were published in the Historiae sive Synopsis Methodicae Conchyliorum and in its early versions (De Cochleis and Historiae Conchyliorum) (Keynes, 1981).

Chemnitz’s figure 1208 (Figure 2) shows dorsal and umbilical views of a shell from the Chemnitz collection (“Ex Museo nostro”’), but in what remains of the Chemnitz collection at the Zoologisk Museum of the University of Copenhagen (Denmark), the figured specimen no longer exists (J. Knudsen, personal communication, June 1993).

This shell has transverse striae, which could be growth lines or ribs, and a very small umbilicus. This feature does not agree with the description in the German text which speaks of a wide umbilicus (“Der Nabel ist rund gross und weit”). Although it is impossible to determine the exact shape of the shell (globose or depressed and cari- nated) from the figure, one can suppose it to be globose, as the figure is one of many in a plate (tab. 133) in which only species with globose shells (““Helices rotundatae”’) are illustrated (e.g., Cepaea, Arianta, Cantareus, Acanthinula). This conclusion seems to be supported by Draparnaud (1805:84) who considered the species identical to that of fig. 1207 and listed it as “Helix variegata Chemnitz” among

Page 4

The Veliger, Vol. 39, No.


Figures 3-7

a Lamarck, 1822, and their old label. The type series is composed of six syntypes and is

Lamarck collection at the Muséum d’Histoire Naturelle of Geneva (Switzerland). Photos by G. Dajoz,

ire Naturelle of Geneva (Switzerland).

G. Manganelli et al., 1996

the synonyms of his Helix variabilis [= Cernuella virgata (Da Costa, 1778)]. Moreover, the periphery of the last whorl is linear and entire both in the original figure (Chemnitz, 1786:pl. 133, fig. 1208) and in the subsequent reproductions (Wood, 1825:pl. 33, fig. 38; Pfeiffer, 1846a: pl. 6, figs. 12, 13). On the contrary, the periphery of the shell of the Sicilian species in dorsal and umbilical view shows a crenulated rim, and this is evident both in our figures (Figures 3, 8, 12), and in the classic iconography of the species (Philippi, 1836:pl. 8, fig. 10; Rossmassler, 1837:pl. 26, fig. 357; Delessert, 1841-42:pl. 26, fig. 4a, b; Pfeiffer, 1843:pl. 23, fig. 4; Benoit, 1862:pl. 4, fig. 25; Tryon, 1887:pl. 61, figs. 79-81).

It is impossible to establish whether Chemnitz (1786) redescribed Lister’s cyclophorid or another species. In any case, he included two (or three?) different species under the name “Die rauhe runzelhafte Schnirtelschnecke” that described with ‘“‘der Nabel rund gross und weit” does not correspond to the one in fig. 1208 which has a very small umbilicus, but possibly to the Jamaican cyclophorid snail illustrated by Lister. The short description may be re- garded as fitting the Sicilian species which Philippi (1836) named Carocolla gargottae, but the shell in figure 1208 does not correspond at all to this species, as Pfeiffer (1846a) realized.

Gmelin (1791:3623) introduced the binomial name He- lix corrugata for a species, to be assigned to a sectio of his genus Helix including species “‘carimatae” i.e., with “‘an- fractuum margine acuto.” This species, living ‘“‘in Jamaica,” was described as “‘testa umbilicata rugosa cana oblique stria- ta: apertura lunata, labro incarnato intus reflexo” and was said to have been described and figured by ‘““Chem. Conch. Sapo memOoo men 208erande dist. Conch. t. 55)f.) 5172

The name Helix corrugata Gmelin, 1791, was assigned to the Sicilian species for the first time by Pfeiffer (1845), and it was used as the latter’s correct name by Albers (1860), Schaufuss (1869), Kobelt (1871, 1904), Tryon (1887), and Pilsbry (1895).

The Jamaican cyclophorid land snail Cyclostoma cor- rugatum Sowerby, 1843, has nothing to do with the Ja- maican cyclophorid reported by Lister and assigned by Gmelin (1791) to his Helix corrugata. Cyclostoma corru- gatum was introduced by Sowerby (1843a:30), without reference to the preceding literature (cf. also Sowerby, 1843b; Pfeiffer, 1846b, 1854; Kobelt, 1901).

Two years later, Salis Marschlins (1793:378 not seen; repeated on p. 472 of the English translation) used the name Helix scabra for an Italian land snail found at Paes- tum (southern Italy) and described as “Testa umbilicata, depressiuscula transversim sulcata, aspera, apertura sublabia- ta ovata.” Although Salis Marschlins referred it to “Mart. [i.e., Chemnitz] t. ix. tab. 133. fig. 1208,” he should be regarded as having described it as a new nominal species (P. Tubbs, personal communication, 28 April 1994).

The name Helix scabra Salis Marschlins, 1793, was

Page 5

assigned to the Sicilian species only by Martens in Albers (1860).

Ferussac (1821:44) also used the name Helix scabra for the species illustrated by Chemnitz (1786:fig. 1208), re- garded Helix corrugata Gmelin, 1791 as its junior synonym and, without a line of description, cited “Helix groyana, nobis. Habit. Entre Fiumesino et le case Brusciota, route de Sinigaglia, 4a Ancone; Comm. Ménard de la Groye,” as a different species.

Pfeiffer’s (1845:24) statement “Dass Hel. Groyana, Fér. pr. 376 [sic] mit der Chemnitz’schen Figur 1208 identisch ist, hat Férussac, wahrscheinlich in Folge fliichtiger Prii- fung des Textes, eben so gut tibersehen, als alle folgende Auctoren” is unfounded. When Feérussac realized this, he could no longer list the two species separately!

Lamarck (1822:90) introduced the name Helix rugosa for a species inhabiting “L’Italie, sur la route d’Ancéne a Sinigaglia,” previously cited as “Helix groyana” at no. 276 of Ferussac’s Prodromus. Finally, Philippi (1836) estab- lished his Carocolla gargottae on shells from “prope Pan- ormum et Termini.”

In conclusion, Chemnitz’s name is not available because itis polynomial (ICZN, 1985:Art. 11c). The names Helix rugosa and Helix scabra, derived from different parts of Chemnitz’s polynomial name, take the author (and date) of the publication in which they were first accompanied by a description, indication, or definition (ICZN, 1985:Art. 50a). Thus Helix rugosa “Chemnitz, 1786” is available from Pfeiffer, 1846a, and Helix scabra “Chemnitz, 1786” from Feérussac, 1821. Since Helix scabra Férussac, 1821, and Helix rugosa Pfeiffer, 1846a, are junior primary hom- onyms of Helix scabra Salis Marschlins, 1793, and Helix rugosa Lamarck, 1822, respectively, they must be rejected.

Four names remain: Helix corrugata Gmelin, 1971, in- troduced for Chemnitz’s “Die rauhe runzelhafte Schnir- telschnecke”; Helix scabra Salis Marschlins, 1793, intro- duced as a new species, though considered to be the same as that illustrated by Chemnitz; Helix rugosa Lamarck, 1822, and Carocolla gargottae Philippi, 1836, indepen- dently established without reference to Chemnitz.

The oldest available name is Helix corrugata Gmelin, 1791, but it is impossible to use it for the Sicilian species. Only the description fits the Sicilian species, but it also fits many other similarly shelled species. The specimens shown in fig. 51 by Lister and fig. 1208 by Chemnitz might constitute the type series of Helix corrugata, and one of them might be selected as lectotype according to art. 74(c) clarifying the identity of Gmelin’s name. Neverthe- less, the selection of fig. 1208, would not automatically provide a name for the Sicilian species because, despite Pfeiffer’s claim, there is no objective evidence that the two correspond.

The identity of the species called Helix scabra by Salis Marschlins and found at Paestum is also uncertain. Fér- ussac (1821:45) believed that it corresponded to his Helix

Page 6 The Veliger, Vol. 39, No. 1


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Figures 8-11

The three shells (and their original label) cited by Férussac as “Helix groyana” [nomen nudum] and reported as collected by M. de la Groye ‘‘Entre Fiumesino et le case Brusciota, route de Sinigaglia, 4 Ancone,” present in the Férussac collection at the Muséum d’Histoire Naturelle of Paris (France).

G. Manganelli et al., 1996 Page 7

Figures 12-15

Shells of Cernuella rugosa (Lamarck, 1822) from Comuni di Casteluzzo (San Vito Lo Capo, Trapani), 33SUC02, I. Sparacio leg. 3.4.87.

Page 8

The Veliger, Vol. 39, No. 1

(Helicella) simulata (nomen nudum!) [= Trochoidea simulata (Ehrenberg, 1831)] from “Alexandrie d’Egypte.” Again, however, there is no objective evidence, apart from the description (insufficient for diagnosis), to support hypoth- eses on its real nature.

Despite its poor original description, the identity of He- lix rugosa Lamarck, 1822, is certain, as six syntypes have been traced in the Lamarck collection at the Muséum d’Histoire Naturelle of Geneva. The type locality of Helix rugosa, as indicated by Lamarck in the original description (“... en Italie, sur la réute d’Ancéne 4a Sinigaglia’’), is probably derived from Férussac because the oldest label accompanying the syntypes reported only “Italie” (Figure 7), and, apart from the six syntypes, no other material of Helix rugosa exists in the Lamarck collection at Geneva.

Férussac reported that his material was collected by Ménard de la Groye, between Fiumesino and Casa Brus- ciote, along the road from Sinigaglia (Sinigaglia is a mis- spelling of Senigallia and Casa Brusciote of Casa Bruciata) to Ancona. However, it has always been impossible to accept that Menard de la Groye could have found the shells studied by Férussac and Lamarck in this locality. Some Italian malacologists (Benoit, 1862; Paulucci, 1878; Alzona, 1971) and Kobelt (1898), in fact, have stressed that the species was endemic to Sicily and that it had never been found in central Italy. Benoit (1862:184) wrote: “dob- biamo confessarlo, non abbiamo finora contezza della it- aliana H. rugosa [we must confess, we do not yet have clear ideas on the Italian H. rugosa].” The presence of a species with shell corresponding to the syntypes of Helix rugosa of Lamarck in the area between Senigallia and Ancona is also excluded by data in the literature (cf. Piersanti, 1933) and by our research. We visited the site indicated by Fer- ussac, which is between the right bank of the Esino River at its mouth and the town of Falconara Marittima (the area is now in the industrial outskirts of Falconara Mar- ittima). We could find only populations of Cernuella cis- alpina (Rossmassler, 1837), the shells of which correspond to those of well-known forms of the coastal regions of northeastern Italy (i.e., shell rather small, finely and dense- ly ribbed, umbilicus rather small, with traces of peripheral carina).

Thus the fact that H. rugosa was reported in the area can only have two explanations: an exchange of material or a mistake in the original indication of the collecting site.

The hypothesis of an exchange of shell material in the Lamarck collection was first advanced by Philippi (1844). In support of this hypothesis, one could adduce the fact that the syntypes are not accompanied by a label hand- written by Lamarck and that Lamarck’s description of Helix rugosa does not fully fit the Sicilian species, as stressed by Philippi (1844) and as Pfeiffer (1845) was obliged to admit.

When Lamarck died, his collection was bought by Prince Masséna who sold it to B. Delessert in 1840. The collection

was later bought by the Muséum d’Histoire Naturelle of Geneva, where it still resides (Dance, 1986).

An exchange of material in the Lamarck collection is unlikely after 1840. Since Delessert’s (1841-1842) figures of Helix rugosa correspond to the present syntypes (Figures 3-6), the Helix rugosa material in the collection was the same when he acquired it. A previous exchange of material also seems unlikey because in her copy of the Animaux sans Vertébres, Ivet Rosalie de Lamarck listed the number of Helix rugosa syntypes in her father’s collection as six, the same as the present syntypes.

As regards the absence of handwritten label by Lamarck, Y. Finet (personal communication, December 1987) told us that: “the lots of the Lamarck collection rarely contain the original labels with Lamarck’s handwriting.” To ex- plain the poor correspondence between the original de- scription and the syntypes, one can adduce the general lack of accuracy of Lamarck’s (1822) descriptions. This lack of accuracy does not invalidate the descriptions but makes it possible to apply them to different species with similar shells.

We also sought the material cited by Férussac as “Helix groyana.” In the Feérussac collection at the Muséum Na- tional d’Histoire Naturelle of Paris (France) there are two lots. One, labelled ““Helicella groyana, nobis 276. Entre Sinigalia et Ancone”’ contains three shells, slightly smaller than those of the Sicilian species and with less marked ribbing (Figures 8-10). The other lacks the original label (the oldest label states “Helix corrugata. Italie’) and con- tains four specimens (one very juvenile) perfectly corre- sponding to the Sicilian species and Lamarck’s syntypes. The last tube also contains one shell of Xerosecta explanata (Miller, 1774) (a species absent from Italy!), and this is indicative of the frequent exchanges of material in old collections.

Férussac’s specimens from the type locality mentioned in Lamarck’s original description have not the status of type material, the type material being the six syntypes in the Lamarck collection at Geneva.

Helix rugosa Lamarck, 1822, must therefore be consid- ered the same as Carocolla gargottae Philippi, a species the identity of which has never been uncertain and which clearly corresponds to the syntypes in the Lamarck col- lection. This seems to us the only possible solution, and consequently we conclude that Carocolla gargottae Philippi, 1836, is a junior synonym of Helix rugosa Lamarck, 1822, the oldest valid name for this species.

A statement of Deshayes (1830:220) that was completely overlooked by all subsequent authors is evidence that this is the correct interpretation: “C’est a Ménard de la Groye que l’on fut redevable de la connoissance de cette espéce; il la découvrit en Italie, prés de Sinigalia: depuis elle fut retrouvee en Sicilie.” This important statement, published no more than 8 years after Lamarck described Helix rugosa and 6 years before Philippi described Carocolla gargottae

G. Manganelli et al., 1996

Figure 16

Genitalia (gonad excluded) of Cernuella rugosa (Lamarck, 1822) from San Vito Lo Capo (San Vito Lo Capo, Trapani), 33SUC02, V. E. Orlando leg. 10.87. Key: AG, albumen gland; BC, bursa copulatrix; BW, body wall; DBC, duct of bursa copulatrix; DG, digitiform glands; DSC, dart sac complex; E, epiphallus; F, fla- gellum; FHD, first hermaphodite duct; FO, free oviduct; GA, genital atrium; P, penis; POS, prostatic portion of ovispermiduct; PR, penial retractor muscle; T, talon; UOS, uterine portion of ovispermiduct; VD, vas deferens.

(but probably written when Lamarck, old and blind, was still alive) and accompanied by an excellent redescription of the species, clearly demonstrates the perfect correspon- dence between the Lamarckian Helix rugosa and the Si- cilian snail.

Cernuella (Cernuella) rugosa (Lamarck, 1822)

[Figures 3-6, 8-10, 12-26]

Helix rugosa Lamarck, 1822:90. Type locality: the locality reported in the original de- scription: “... en Italie, sur la route d’Ancone a Sini- gaglia” is certainly incorrect and taken from Ferussac.

Page 9

The old label (Figure 7) accompanying the syntypes indicates only “Italie” and, on the basis of available data, the species is known to live only in Sicily. Type series: six syntypes in the Lamarck collection at the Muséum d’Histoire Naturelle of Geneva (Switzer- land) (Figures 3-6).

Carocolla gargottae Philippi, 1836:136-137; pl. 8, fig. 10. Type locality: “prope Panormum et Termini.” Type series: according to R. Kilias (personal commu- nication, May 1994) there is no syntype in the Philippi Collection, at the Zoologisches Museum und Institut fur spezielle Zoologie of Berlin (Germany).

Helix pleurischura Bourguignat, 1876:44-45. Type locality: ‘““Habite la Sicile.” Type series: holotype in the Bourguignat collection at the Muséum d’Histoire Naturelle of Geneva (Switzer- land) (Figure 26).

Helix chonomphala, Bourguignat, 1876:45-46. Type locality: “La Sicile.” Type series: lost? No syntype is found in the Bour- guignat collection at the Muséum d’Histoire Naturelle of Geneva (Switzerland).

Historical material examined: “‘Calatafimini, 1 esempl. ricev. 1877 dal Sig. Benoit col nome di H. gargottae”’ (1 det. H. rugosa var. minor; Coll. Paulucci, Museo di Zool- ogia dell’Universita di Firenze, MZUF 6709); “‘Calataf- imini, 2 esempl. ric. dal Sig. Benoit 1877” (2 det. H. rugosa; Coll. Paulucci, MZUF 6710); ‘“Calatafimini, 2 esempl. ricev. dal Sig. Benoit 1877 col nome di H. gargottae”’ (2 det. H. rugosa var. alba; Coll. Paulucci, MZUF 6711); “Dintorni di Messina, 6 esempl. ricevuti 1870 dal Sig. Benoit a Messina” (6 det. H. rugosa var. minor; Coll. Paulucci, MZUF 6712); ‘“‘Presso Messina” (4 det. Helix gargottae; Coll. Paulucci, MZUF 6713); “Redicofani” (15 det. Helix rugosa; Coll. Paulucci, MZUF 6714); “Sicilia, presso Messina, 12 esempl. 5 ricevuti dal Mse Monterosato 1870 da Trapani, 7 avuti stesso anno dal Sig. Benoit come raccolti presso Messina” (11 det. Helix rugosa; Coll. Pau- lucci, MZUF 6717); “Trapani e Messina, 6 esempl. ri- cevuti meta dal Mse Monterosato come provenienti da Trapani, meta dal Sig. Benoit come raccolti presso Mes- sina- 1870” (6 det. Helix rugosa var. alba; Coll. Paulucci, MZUF 6718).

New material examined: San Vito Lo Capo (San Vito Lo Capo, Trapani), 33SUC02, V. E. Orlando leg. 10.87 (numerous specimens); S. Cianfanelli & M. Calcagno leg. 27.8.91 (5); Comuni di Casteluzzo (San Vito Lo Capo, Trapani), 33SUC02, I. Sparacio leg. 3.4.87 (numerous specimens); Torretta Granitola (Mazara del Vallo-Cam- pobello di Mazara, Trapani), 33STB95, S. Cianfanelli & M. Calcagno leg. 21.8.91 (5).

Diagnosis: A species belonging to Cernuella (Cernuella) easily distinguishable from the two other, currently rec- ognized species of the nominotypical subgenus of Cernuella, C. virgata (Da Costa, 1778) and C. cisalpina (Rossmassler, 1837), by virtue of medium-sized, robust, lenticular, shell,

The Veliger, Vol. 39, No. 1

Page 10

17.20 2mm


G. Manganelli et al., 1996

Table 1

Dimensions of the shell of Cernuella rugosa. For each pa- rameter mean standard deviations and range are given.

Number Shell Shell of spec- Locality diameter height imens San Vito lo Capo 12.62 + 0.58 6.28 + 0.58 5 (12.0-13.3) (5.3-7.0) Comuni di Castelluzzo 12.35 + 0.37 5.91 + 0.66 10 (11.6-13.1) (4.8-6.8) Torreta Granitola 14.50 + 0.75 5.56 + 0.34 5 (13.3-15.4) (5.2-6.2)

with raised, irregularly spaced ribs and evident, cordlike keel at periphery.

Shell (Figures 3-6, 8-10, 12-15): Shell dextral, medium- sized, robust, lenticular, uniformly yellowish-grey or with traces of pale brown bands most evident in lower half, opaque; external surface with evident raised ribs, rather irregularly positioned and spaced: spire depressed, some- times almost flattened, consisting of 5-5’ clearly convex whorls, regularly and slowly enlarging; last whorl large, moderately dilated, sometimes descending near aperture, with evident, cordlike keel at periphery; sutures moder- ately deep, shouldered by keel of preceding whorls; um- bilicus open, very large (about one-third of shell maximum diameter); aperture oval, its external margin angled at keel; peristome interrupted, simple, not reflected, with upper margin sometimes starting at keel, sometimes below keel, with well-developed, yellowish or brownish, internal cal- lous rib.

Dimensions (Table 1). Height: 4.8-7.0 mm; maximum diameter: 11.6-15.4 mm.

Body: Animal blackish, darker above; retractor of right ommatophore independent of penis and vagina; kidney sigmurethrous; jaw odonthognathous; penial nerve origi- nating from right pedal ganglion.

Genitalia (Figures 16-20): General scheme of the semi- diaulic monotrematic type, characterized by: large her-

Page 11

maphrodite gonad (ovotestis) consisting of bunch of acini whose ducts converge into first hermaphrodite duct, its initial portion very slender, then widening to function as seminal vesicle; first hermaphrodite duct ending in clublike “talon” adhering to internal side of large, beanlike albu- men gland; talon