On Cyprian Helicellinae (Mollusca: Gastropoda Pulmonata: Helicidae), making a new start E. Gittenberger Gittenberger, E. On Cyprian Helicellinae (Mollusca: Gastropoda Pulmonata: Helicidae), making a new start. Zool. Med. Leiden 65 (7), 31.vii.1991:99-128, figs. 1-47.— ISSN 0024-0672. Key words: Helicidae; Helicellinae; taxonomy; key; Cyprus. Our fragmentary knowledge concerning the Cyprian Helicellinae is revised and summarized, while new data are added. "HeliceUa (Xerotricha) conspurcata distinguenda" Haas, 1936, is a Helicopsis species that has to be called H. cypriola (Westerlund, 1889) as a consequence of the selection of a neo- type for Westerlund's poorly described nominal taxon. "HeliceUa (Xerotricha) conspurcata arrouxi" sensu Haas, 1934 (not Bourguignat, 1863) is described as HeliceUa juglans spec. nov. This species stands apart among the Helicellinae by the lack of a separate penis. Neither material nor reliable records of Xerotricha conspurcata (Draparnaud, 1801) could be traced; only X. apicina (Lamarck, 1822) is known from Cyprus. The wide-spread "arrouxi" has also been found on the island; for nomenclatural reasons it should be called Microxeromagna armillata (Lowe, 1852). Xeropicta akrotirica spec. nov. is distin guished from small specimens of X. krynickii (Krynicki, 1833). By the selection of a neotype for Helix cyparissias Pfeiffer, 1847, this nominal taxon is stabilized as a junior synonym of Xeropicta krynickii. Xeropicta ledereri (Pfeiffer, 1856) is hypothesized to be a polytypic species with X. /. mavromoustakisi (Haas, 1933) as a subspecies. The status of HeliceUa (Jacosta) syrensis carinatoglobosa Haas, 1934, remains unclear. Two species that are mentioned in the literature under incorrect names, are described as new: Trochoidea (Xerocrassa) nicosiana spec. nov. and Pseudoxerophila confusa spec. nov. E. Gittenberger, Nationaal Natuurhistorisch Museum (Rijksmuseum van Natuurlijke Historie), Postbus 9517, NL-2300 RA Leiden, The Netherlands. Contents Introduction .. 99 Provisional conchological key 100 Systematic review 102 Helicopsis Fitzinger, 1833 102 Xeropicta Monterosato, 1892 107 Pseudoxerophila Westerlund, 1879 113 Xeromunda Monterosato, 1892 115 HeliceUa Ferussac, 1821 115 Xerotricha Monterosato, 1892 119 Microxeromagna Ortiz de Zarate, 1950 120 Trochoidea Brown, 1827 122 Cochlicella Ferussac, 1821 125 Acknowledgements 126 References 126 Index of epithets 128 Introduction In the Helicellinae sensu auct. there is hardly any correlation between the shell characters and the structure of the genitalia. It has been suggested (e.g. Schileyko, 100 ZOOLOGISCHE MEDEDELINGEN 65 (1991) 1978) that these conchological similar species, characterized anatomically by a right ommatophoran retractor that runs next to the genital ducts instead of crossing between penis and vagina, do not constitute a monophyletic group. The present author agrees that most arguments are in favour of a polyphyletic origin indeed. Nevertheless the classical definition of the Helicellinae is maintained, because a sta ble alternative classification and nomenclature cannot be presented yet. This general problem is not dealt with further in the present paper. It suffices to emphasize that the (sub)generic position of Helicellinae species cannot be based reliably on only their shell characters, since it has been demonstrated convincingly that convergent evolution with regard to shell size, shape and sculpture has occurred frequently in this subfamily. While trying to identify some samples of Helicellinae from the island of Cyprus, this turned out to be impossible in several cases. The literature on the subject proved to be highly incomplete and not of much help, especially because data concerning the structure of the genitalia are scarce. Haas (1934, 1936) exemplifies the risk of purely conchological research in Helicellinae by uniting what proved to be species belonging to three different genera (Helicopsis, Xeropicta and HeliceUa) as only two subspecies of a species that most probably does not occur on Cyprus (Xerotricha con spurcata). In the newly collected material by far not enough samples with well preserved specimens were available to enable the composition of a satisfactory monograph concerning the systematics and distribution of the Cyprian Helicellinae. However, some problems can be outlined better now and several confusing errors in the litera ture can be corrected. This is the main purpose of the present paper, in which the taxa are dealt with more or less profoundly, depending upon the amount of easily accessible information in the literature and the state of our actual knowledge. It turned out that even on a relatively small island like Cyprus, distributional patterns may occur that are relevant for the interpretation of taxonomic relation ships. For collections the following abbreviations are used: Bar, collection Z. Bar (Hilversum); Hem, collection J. Hemmen (Wiesbaden); MHNG-Bgt, collection J.R. Bourguignat in Museum d'Histoire Naturelle (Gen&ve); Museum National d'Histoire Naturelle (Paris); Neu, collection W. H. Neuteboom (Heemskerk); NMG, Naturhis- toriska Museet (Goteborg); RMNH, Rijksmuseum van Natuurlijke Historie (Leiden); SMF, Forschungs-Institut Senckenberg (Frankfurt am Main); Smi, collection D. Smits (Woudenberg). Provisional conchological key The key is of limited value because several problematical forms that are mentioned in the text, are not included. It should also be kept in mind that some species are extremely variable in size and shape. Single specimens cannot always be identified, therefore. This is especially so in Xeropicta kryn ickii and X. ledereri. 1. Shell higher than broad 2 Shell width and height about equal, or shell clearly broader than high 4 GITTENBERGER: CYPRIAN HELICELLINAE 101 2. With prominent radial ribs Trochoidea (T.) liebetruti - Without ribs 3 3. Shell clearly more than twice as high as broad Cochlicella acuta Shell clearly less than twice as high as broad Cochlicella conoidea 4. Shell globular, relatively thin; radial colour pattern dominating; umbilicus mea suring less than 1/10 of the shell width Xeromunda candiota - Different 5 5. Spire conical, with straight sides; periphery of at least the final quarter of the body-whorl regularly rounded, and (initial) teleoconch whorls provided with fine regular riblets Xeropicta krynickii Different 6 6. Shell rounded conical in front view; apertural height less than half the total height; umbilicus measuring less than 1/7 of the shell width Trochoidea (T.) pyramidata - Different 7 7. Shell width over 9 mm 8 Shell width less than 9 mm 14 8. Umbilicus measuring less than 1/6 of the shell width 9 - Umbilicus measuring over 1 / 6 of the shell width 12 9. Shell relatively thin, very globular, with a circular aperture Xeropicta akrotirica - Different 10 10. Sculpture obsolete, very irregular 11 Sculpture prominent, rather coarse and not very irregular; width 10-13 mm "HeliceUa" carinatoglobosa 11. Umbilicus measuring 1/10 of the shell width, or less; aperture elliptical, with an internal rib Trochoidea (Xerocrassa) nicosiana Umbilicus more widely open; aperture roundish, with a weak internal rib Trochoidea (Xerocrassa) cretica 12. Body-whorl, both in juvenile and in full-grown specimens, conspicuously keeled; the inner border of the aperture follows the angle of the keel Xeropicta ledereri ledereri - Different 13 13. Whorls increasing very gradually in width, all of them separated by indented sutures Pseudoxerophila confusa Body-whorl dominating the shell, separated from the penultimate one by an indented suture; initial teleoconch whorls with flattened sutures, accompanied by (traces of) a slight indentation, resulting from the keel of young specimens Xeropicta ledereri mavromoustakisi 14. Shell covered with hairs or prominent hair-scars 15 - Neither hairs nor hair-scars are clearly visible 16 15. Hairs very short and densely covering the shell; aperture elliptical Microxeromagna armillata - Hairs long and widely spaced; aperture circular and relatively large Xerotricha apicina 16. Aperture in both juvenile and full-grown specimens with a prominent angle, cor responding with the sharp peripheral keel Xeropicta ledereri ledereri 102 ZOOLOGISCHE MEDEDELINGEN 65 (1991) — Different 17 17. Umbilicus measuring 1/4-1/5 of the shell width 18 — Umbilicus measuring 1 /6-1 /7 of the shell width 19 18. Juvenile shells with a peripheral keel, that is accentuated by (traces of) a slight, upper indentation and a flattened part of the whorls (small specimens, only known from the eastern half of Cyprus) Xeropicta ledereri mavromoustakisi — Juvenile shells angular at the periphery, but whorls evenly rounded between the sutures and (strongly) inflated; western half of Cyprus Helicopsis cypriola 19. Aperture circular Xeropicta akrotirica — Aperture elliptical HeliceUa juglans (or Helicopsis spec.) Systematic review [Following the (sub)generic names, the diagnostic characters of the genitalia are summarized.] Helicopsis Fitzinger, 1833 Genitalia with two dart-sacs, accompanied by accessory sacs. Helicopsis cypriola (Westerlund, 1889) (figs. 1-4,16,17, 20) Helix cyparissias var. cypriola Westerlund, 1889: 252 (without locality); neotype (design, nov.): NMG. Helix (Xerophila) cyparissias var. cypricola [sic!]; Rolle & Kobelt, 1896:55. HeliceUa (Xerotricha) conspurcata distinguenda Haas, 1936: 300, fig. 8 ("Limassol", SMF 10418/holotype, 10419/5 paratypes). Material (fig. 20).— 1.5 km NW. of Coral Bay (= 12 km NW. of Paphos), VD35 (Hem); Cape Lara, VD36 (Hov; RMNH); Ayios Yeorgios, VD36 (Hov; RMNH); 3 km N. of Ayios Yeorgios (= 20 km NNW. of Paphos), VD36 (Hem); "Baths of Aphrodite" (= 4.5 km NW. of Neokhorio, = Neon Chorion), VD38 (Bar; RMNH); Paphos, near the Technical School, VD44 (Neu); Kato Paphos, E. of the main road, VD44 (Bar; RMNH); Peyia, VD45 (Bar); 1 km NE. of Coral Bay (= 10 km N. of Paphos), 100 m alt., VD45 (Hem; RMNH); 0.5 km W. of Kissonerga, 30 m alt., VD45 (Hem); 2 km NW. of Peyia, VD46 (Bar; RMNH); archeological site of Kouklia, 100 m alt., VD53 (Hem; Neu); Ayia Marina (= 2 km N. of Yialia), VD58 (Hem); 5 km WSW. of Pissouri, 100 m alt., VD63 (Hem); "Petra tou Romiou", ravin opposite the "Venus rock" (= 14 km SE. of Paphos), VD63 (Neu); 1 km SW. of Nikoklia, 80 m alt., VD64 (Hem); 2 km NE. of Nikoklia, 200 m alt., VD64 (Hem); 1 km S. of Pissouri, 250 m alt., VD73 (Hem; RMNH); 2 km N. of Pissouri Bay, 100 m alt., VD73 (Hem); 0.5 km N. of Pissouri Bay, VD73 (Hem); archeological site of Kourion, 10 km W. of Limassol, 80 m alt., VD83 (Hem); Episkopi, VD83 (Neu); 1 km NW. of Kantou (= 3.5 km N. of Episkopi), VD83 (Hem); Akrotiri forest, VD92 (SMF 98261/3, 10420b/2 [of 41 "distinguenda"); Limassol, WD03 (NMG/neotype cypriola; RMNH; SMF 10418/holotype distinguenda, 10419/5 paratypes distinguenda). Shell (figs. 1-4).— Shell more or less depressed conical, with 4-5 (very) convex whorls, separated by deep sutures and increasing slowly and regularly in width. The teleoconch is sculptured with coarse, rather irregular transverse ribs. Maybe there are vague hair-pits, but specimens with hairs have not been observed. The body- GITTENBERGER: CYPRIAN HELICELLINAE 103 Figs. 1-4. Helicopsis cypriola (Westerlund). 1-3, Limassol, iv.1935, G. A. Mavromoustakis leg. (actual width 6.4 mm) (RMNH); 4, neotype, idem (actual width 5.1 mm) (NMG). Fig. 5. Helicopsis spec., 1 km W. of Analiondou, 8.iv.l985, A. Hovestadt leg. (actual width 5.8 mm) (RMNH). Figs. 6, 7. Helicella juglans spec, nov., holotype, Nicosia, iv.1935, G. A. Mavromoustakis leg. (actual width 7.3 mm) (RMNH 56404). Figs. 8,9,14,15. Xeropicta akrotirica spec. nov. 8,9, holotype, Akrotiri forest, 1938, G. A. Mavromoustakis leg. (actual width 7.4 mm) (RMNH 56409); 14,15, paratype, Episkopi, 2.xi.l981, W. H. Neuteboom leg. (actual width 7.0 mm) (RMNH 56410). Fig. 10. Microxeromagna armillata (Lowe), Nicosia, under rind of a conifer in the garden of the Armenian School, x.1978, D. Smits leg. (actual width 55 mm) (RMNH). Figs. 11-13. Xeropicta krynickii (Krynicki), neotype of cyparissias, Larnaca, Artemis Avenue, 8.xi.l981, W. H. Neuteboom leg. (actual width 6.7 mm) (RMNH 56412). 104 ZOOLOGISCHE MEDEDELINGEN 65 (1991) Figs. 16-17. Helicopsis cypriola (Westerlund), proximal genitalia. 16, archeological site of Kouklia, 6.xi.l981, W. H. Neuteboom leg. (RMNH slide 958); 17, Cape Lara, 5.iv.l985, A. Hovestadt leg. (RMNH slide 956). Scale lines 1 mm. Abbreviations: A, vas deferens; Ap, appendage (see e.g. fig. 24); D, sper- mathecal duct; Da, accessory sac; Ds, dart-sac; Dt, dart; E, epiphallus; F, flagellum; G, mucous gland; Ga, albumen gland; H, hermaphroditic duct; O, oviduct; P, penis; Pr, prostate; R, penial retractor mus cle; S, spermatheca; So, spermoviduct; V, vagina. whorl may be regularly rounded to more or less clearly angular, or even carinate, at the periphery, especially above the aperture. Near the aperture the body-whorl is not or hardly descending in front. The umbilicus usually measures between 1/4 and 1/5 of the total shell width; it is slightly eccentric to symmetrical. The aperture is slightly broader than high and provided with a simple, moderately prominent, internal rib. The colour pattern is extremely variable, whitish with an irregular corneous spotting, which is often at least partially arranged into one or more interrupted spiral bands. Width 5.0-6.6 mm; height 3.3-4.3 mm. Generally this species can be recognized by its wide, symmetrical umbilicus and GITTENBERGER: CYPRIAN HELICELLINAE 105 the convex whorls with a coarse sculpture. Very small specimens of Xeropicta ledereri mavromoustakisi, known from near Cape Greco, may look very similar, differing only by the shallower sutures that are accompanied by a slight, parallel indentation. Genitalia (figs. 16, 17).— The penis is gradually increasing in width towards the genital atrium; its proximal part is about twice as long as the distal part and in line with a somewhat inflated part of the genital atrium. The flagellum is as long as the penis (fig. 16) or shorter (fig. 17), measuring less than 1/3 of the length of the epiphallus. The vagina is as long as the penis or slightly shorter. The spermathecal duct is somewhat shorter than the epiphallus. The dart-sacs insert close to the genital atrium. Notes.— Syntypes of Helix cyparissias var. cypriola could be traced neither in the Naturhistoriska Museet, Goteborg (Von Proschwitz, in litt. 4.10.1988), nor in the Naturhistoriska Riksmuseet, Stockholm (A. Waren, in litt. 6.11.1988). The original description of the taxon is insufficient; the variety is said to be smaller than "cyparis sias", with a rather conical spire, an angular periphery accompanied by a spiral band just above it and some finer spiral banding below. This may apply to more than one Cyprian species. Therefore, to stabilize the nomenclature of the small Cyprian He licellinae, a neotype is selected for cypriola from a sample from Limassol and deposit ed in the part of the Westerlund collection housed in Goteborg. While introducing HeliceUa (Xerotricha) conspurcata distinguenda, which becomes a junior synonym by this action, Haas (1936) referred neither to Helix cyparissias nor its variety cypriola. Haas (1936: 300) mentioned a specimen of distinguenda also from the Akrotiri for est (SMF 10420/paratype). In an earlier paper (Haas, 1934: 19, fig. 6) this specimen had been figured. The shell in question is very globular, with a circular aperture and a relatively large body-whorl. It could be restudied in SMF and proved to belong to the sympatric Xeropicta akrotirica. A sample from Kato Paphos (shells only) contains very small, keeled specimens. The smallest probably full-grown shell, with 3 1/4 whorls, measures only 3.8 mm in width and 2.3 mm in height. In the sample from Ayia Marina the umbilicus is relatively narrow, measuring c. 1/6 of the shell width. Specimens collected 1.5 km NW. of Coral Bay are more de pressed than usual and have an umbilical width of nearly 1/3 of the total shell width. Helicopsis spec, (figs. 5,18-20) Material.— 2 km S. of Vizalea, WD07 (Hov/5; RMNH/1); 1 km W. of Analiondou, WD27 (Hov/4; RMNH/1); Moni Ayios Heracleidos, WD27 (Hov/4; RMNH/1). Shell (fig. 5; see "Notes") — The shells have 3 1/2 to 5 whorls. They are relatively fragile and provided with a sculpture which is less prominent than that of H. cyprio la. The whorls of the conical spire are much more flattened and separated by less deep sutures. The body-whorl is angled. The umbilicus measures 1/9 to less than 1/6 of the total shell width and the aperture may be somewhat more descending in front. Width 5.3-7.2 mm; height 3.4-5.4 mm. Genitalia (figs. 18, 19).— The genitalia that could be studied are very similar to those of H. cypriola. 106 ZOOLOGISCHE MEDEDELINGEN 65 (1991) Figs. 18-19. Helicopsis spec., proximal genitalia. 18,1 km W. of Analiondou, 8.iv.l985, A. Hovestadt leg. (RMNH slide 960); 19, Moni Ayios Heracleidos, 8.iv.l985, A. Hovestadt leg. (RMNH slide 959). Scale lines 1 mm. For abbreviations, see legends to figs. 16,17. Notes.— From three localities (fig. 20) outside the range of Helicopsis cypriola specimens of a Helicopsis spec, are known, which differ conspicuously from the pre ceding species. A name for this form could not be traced in the literature. The shells look hardly adult, but the animals dissected from them proved to be full-grown, or nearly so, anatomically. Although the shells listed above look all very similar, the possibility that a more variable taxon is involved, has to be taken into consideration. At two of the localities some additional, somewhat deviating, empty shells were found: two specimens col lected near Analiondou that differ slightly in having less markedly flattened whorls (resembling Helicopsis cypriola) and two specimens collected at Moni Ayios Hera cleidos (more similar to HeliceUa juglans). Xeropicta Monterosato, 1892 Genitalia with two dart-sacs, accompanied by accessory sacs that surpass the dart-sacs in length, and a conspicuous appendage. GITTENBERGER: CYPRIAN HELICELLINAE 107 Xeropicta akrotirica spec. nov. (figs. 8,9,14,15,21,26) HeliceUa (Xerotricha) conspurcata arrouxi; Haas, 1934: 20 [part.], fig. 6 (Akrotiri forest). Not arrouxi Bourguignat, 1863. Material (fig. 21).— Holotype: Akrotiri forest, VD92 (RMNH 56409 "HeliceUa akrotirica Tomlin"; G. A. Mavromoustakis leg.). Paratypes: type locality (RMNH 56411/2 [with the holotype]; SMF 10420a/3, 10420b/2 [of 4] "distinguenda")', Episkopi, VD83 (Neu/14; RMNH 56410/2); Hala Sultan Tekkesi (= 5 km SW. of Larnaca), WD55 (Hem/4); 3 km NE. of Paralimni (= c. 5 km SE. of Famagusta), WD97 (Hem/1). Figs. 20-23. UTM 10 km-grid distribution maps. 20, Helicopsis cypriola (Westerlund) [dots] and Helicopsis spec, [triangles]; 21, Xeropicta akrotirica spec, nov.; 22, X. krynickii (Krynicki); 23, X. ledereri mavromoustakisi (Haas) [dots], with aulostoma [A], andrewi [An], ledereri sensu stricto [L], tremithensis [T], and "HeliceUa" carinatoglobosa [C]. 108 ZOOLOGISCHE MEDEDELINGEN 65 (1991) Shell (figs. 8, 9,14,15).— Shell globular, with 4 1/4-5 inflated whorls, sculptured with irregular, rather coarse riblets. Aperture hardly descending in front, circular, with a simple, (very) weakly developed internal rib. Umbilicus symmetrical, measur ing 1/6-1/7 of the total shell width. Colour pattern variable, spirally arranged. Shell width 6.0-9.8 mm; height 4.6-7.4 mm. X. akrotirica differs from the congeneric species by the globular, relatively small shells, with an irregular sculpture, and a narrow umbilicus. Genitalia (fig. 26).— The proximal part of the genitalia of the holotype was stud ied superficially after having softened dry remains of the animal with sodium phos phate. The flagellum is relatively short, as in X. ledereri and X. derbentina (Krynicki, 1836). The appendage is very prominent. Notes.— The species has been found in sympatry with X. ledereri mavromoustak- isi. As far as known, the epithet "akrotirica" has never been published in connection with this species. Xeropicta krynickii (Krynicki, 1833) (figs. 11-13, 22,24,33) Helix cyparissias Pfeiffer, 1847: 32 ("insulo Cypro"): neotype (design, nov.): RMNH 56412. Helix lauta sensu Bourguignat, 1853: 29. Not Lowe, 1831. Helix larnacensis Rolle & Kobelt, 1896: 55, pi. 22 figs. 10,11. Helix cretica var. littoralis Mousson, 1854:378 ("Cypre"). HeliceUa q/parissias var. cypriola sensu Hesse, 1934:29, pi. 6 fig. 51 (genitalia). Not Westerlund, 1889. HeliceUa larnacensis; Hesse, 1934:30, pi. 6 fig. 52 (genitalia). Material (fig. 22).— Kato Paphos, (a) "Apostle Paulus Avenue", (b) "Tombs of the Kings", and (c) near the "Odeion Gymnasium", VD44 (Neu); Paphos, near the Technical School, VD44 (Neu); Paphos, near the lighthouse, 20-30 m alt., VD44 (Hem); 1 km N. of Peyia, 7 km NNW. of Paphos, 300 m alt., VD45 (Hem); Peyia, VD45 (Bar); 2.5 km W. of Polis, 50-80 m alt., VD47 (Hem); Polis, direction "Baths of Aphrodite", VD47 (RMNH; Smi); archeological site of Kouklia, 100 m alt., VD53 (Hem); Yeroskipos, VD54 (Bar); "Petra tou Romiou", ravin opposite the "Venus rock" (= 14 km SE. of Paphos), VD63 (Neu); 2 km NE. of Nikoklia, 200 m alt., VD64 (Hem); 1 km S. of Pissouri, 250 m alt., VD73 (Hem); 2 km N. of Pissouri Bay, 100 m alt., VD73 (Hem); 1 km N. of Evdhimou Bay, 3 km E. of Pissouri, 100 m alt., VD73 (Hem); archeological site of Kourion, 10 km W. of Limassol, 80 m alt., VD83 (Hem); Limassol, WD03 (RMNH "cretica littoralis, G. A. Mavromoustakis leg."); Nicosia, WD39 (RMNH); Limassol — Larnaca, 10 km SW. of Larnaca near Agkleisides, WD45 (RMNH; Smi); 2 km W. of Kalokhorio (= Kalon Chorion) (= 10 km W. of Larnaca), WD46 (Hem); Larnaca, WD56 (Bar; RMNH; SMF 67374/1, 289611/6, 67378/10 "cyparissias"); Larnaca, "Artemis Avenue", WD56 (Neu; RMNH 56412/neotype "cyparissias"); 1 km NW. of Phrenaros, 3.5 km W. of Paralimni, 100 m alt., WD87 (Hem); 3 km S. of Liopetri (= 10 km W. of Ayia Napa), WD87 (Hem); Famagusta, WD88 (SMF 67367/1, 67371/3, 67376/31, 67379/48 "cyparissias"); Ayia Napa, northern side, WD97 (Hem); 2.5 km NE. of Agia Napa, WD97 (Hem). Shell (figs. 11-13, 33).— The conical spire of the shell is usually somewhat elevat ed, not strongly depressed; the whorls are separated by indented sutures. Aperture roundish to slightly elliptical, not or shortly descending in front; palatal lip inserting at the periphery or just below it. The initial teleoconch whorls are sculptured with regular, sharp and fine, transverse riblets, similar to those of for example the western European Candidula gigaxii (Pfeiffer, 1850). On the body-whorl this very characteristic sculpture may be somewhat less regular. The eccentric umbilicus measures less than