BASTERIA, 125-155, 1998 Greece, in Particular On
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BASTERIA, Vol 62: 125-155, 1998 New data the fauna of Clausiliidae of in on Greece, particular on Albinaria from Attica and the Peloponnese (Gastropoda Pulmonata: Clausiliidae) Eike Neubert und Naturmuseum D-60325 Forschungsinstitut Senckenberg, Senckenberganlage 25, Frankfurt am Main, Germany Initiated a short nomenclatural of H. Nordsieck by note (1995) an attempt is made to supply information the nomenclatural which more on confusing problems arose from different of the interpretation specific names grisea Deshayes and maculosa Deshayes as used by the most A important authors in the last century. table is added summarizing the use of this names to facilitate the understanding of the publications ofthese authors. The polytypic species hitherto has bear the oldest known as Albinaria hohorsti H. Nordsieck, 1984, to group name available which is Clausilia 1878. The bigibbosula Westerlund, following taxa are described as new to science: Albinaria grisea micromaculata, Albinaria litoraria,Albinaria mixta inflexa, Albinaria argynnis geraniensis Albinaria solicola solicola , andAlbinaria solicola leonidiae. for the taxa , Lectotypes following Clausilia are designated: Clausilia grisea Deshayes, Clausilia (Albinaria) heterochroa Westerlund, (Albinaria) heterochroa f. werneri Clausilia Sturany, argynnis Westerlund. Arguments are presented from Attica and the The for the grouping of Albinaria species Peloponnese. most common in the is Albinaria which is here species area grisea, considered to encompass five subspecies. the Several conchologically stable forms are described which probably are result ofhybridization ofA. with grisea ssp. species living nearby. Probable hybridizationzoneswith related species are located, intermediate forms are illustrated. Key words: Gastropoda, Pulmonata, Clausiliidae, Albinaria, taxonomy, Greece, Attica, Peloponnese. INTRODUCTION The Clausiliidae, in particular the genus Albinaria von Vest, 1864, exhibit a remark- able diversity throughout the eastern Mediterranean region. Although the interest of many malacologists focused on this family since the early years of the last century, new taxa are still to be discovered in this area today. This is mainly caused by the tendency ofseveral ofthe involved into forms with genera to split up local highly restricted areas of distribution. It is also furthered by geological and geographical factors like various substrata or extreme variation in habitat structures within a few kilometres. One of the most common Albinaria species in the central to eastern Peloponnese and the like Attica is A. neighbouring areas grisea (Deshayes, 1833). This polytypic species is often misidentified and confused with forms of other related Albinaria very closely which makes it this work into nomenclature and species necessary to split a part on another with of one dealing questions systematics. Additionally, a conchological ap- proach is attempted again to provide clear definitions of species and subspecies. A of revised material is and the distri- compilation given maps are supplied illustrating bution of the taxa in the area under discussion. 126 BASTERIA, Vol. 62, No. 3-4, 1998 MATERIAL AND METHODS Traditionally, several technical terms were used for describing details of the clausilial apparatus derived from publications of German students of Clausiliidae. Some of these be the dentition of structures, however, are supposed to homologous to apertural the Pupilloidea. As a consequence, several scientists started to use the terms 'columellaris' for 'lower lamella' (Unterlamelle) and 'angularis' for 'upper lamella' (Oberlamelle). Although this is correct, the author prefers to use the more traditional names here. The here is based the material collected investigation presented mainly on by the author during several travels to Greece and the collection of the Forschungsinstitut und Naturmuseum Senckenberg in Frankfurt/Main. Material from several private collec- tions is added. Collections: GNM = Goteborgs Naturhistoriska Museum MNHNP = Musee National d'Histoire Naturelle Paris NEUB = private collection E. Neubert NMNH — National Museum of Natural History Leiden NMW = Naturhistorisches Museum Wien NORDS = private collection H. Nordsieck SMF = Senckenberg Museum Frankfurt/Main SUB = private collection P. Subai = = = Abbreviations: D — diameter; H height; PD diameter of peristome; PH height of peristome; W = number of whorls; rkm = road kilometre. NOMENCLATURE to One ofthe first describe terrestrial molluscs from Greece was Deshayes in St. Bory de who about Vincent, reported the scientific results of an expedition to the Pelopon- nese. In his part, Deshayes described and/or figured four species of Clausiliidae and of the Buliminidae. of one species Recently, most the original material of Deshayes was rediscovered by Dr. Ph. Bouchet in the collection of the Ecole des Mines (Paris) and transferred the MNHNP to (pers. comm. Bouchet, 1996). The syntypes of the taxa described by Deshayes in 1833 had been checked by H. Nordsieck, who published about the nomenclatural of this in consequences investigation 1995. A short review ofthe extremely confusing situation following Deshayes' work is given here to facilitate understanding of the problematic use ofthe taxa Clausilia maculosa and C. grisea. described several details of Deshayes the shells, but unfortunately, most of these are not that important or refer to intraspecific variation. The nomenclature and homology of features of the clausilial single apparatus were not yet defined. As a result, one only can assume which lamella or fold he actually described. In his C. maculosa for example, he described the which is aperture as 'tridentata', the case in most of the Clausiliidae and lower In C. he also that (upper lamella, subcolumellaris). grisea noted, there may be another 'fold' diverging from the lower lamella. This is no fold but the inner rim of the columella which is often somewhat callous and thus seems to be bifurcate. A few later of years (1838), Deshayes changed parts the description of his species focusing the of even more on morphology the columellarfolds and callosities and thus producing the base for misinterpretation. Neubert: Greek Clausiliidae, particularly Albinaria in Attica/Peloponnese 127 In Roth the results 1839, published of his expedition in the eastern Mediterranean and and what he believed be C. region reports figures subsequently to grisea Deshayes 2 It is evident from both and that he dealt with (pp. 21-22, pi. fig. 6). text figure a mixture of and A. both, A. bigibbosula true grisea (in the sense they are used today). On he describes be found variation within p. 22, two palatal pliculae to as a the specimens of his collection, a character, which can often be found in A. bigibbosula subspecies but never in the true A. grisea. Roth cites Deshayes supposing that he also found two palatal folds in his C. Fact that in 1833 wrote 'on voit deux grisea. is, Deshayes gros plis sur la columelle' which then was obviously misunderstood by Roth. Moreover, Roth's magnified figure (pi. 2 fig. 6) shows a last whorl with flat-topped and evenly wide-spaced riblets as is characteristic for A. bigibbosula and never found in the true A. grisea. The of these characters in description some specimens proves, that he dealt at least with two species. Roth's mistake from 1839 was followed up by Pfeiffer (1848: 457), who had problems to recognize A. grisea from the description of the original author and thus refers to Roth concerning the variation within this species. Bourguignat (1853: 48) identified without any comment C. grisea sensu Roth with C. maculosa Deshayes and records C. grisea from Athens and Deshayes (correct) Sparta (confusion with A. g. immensa (O. Boettger, 1889)?). At the same time, Charpentier (1852: 371) in his work on Clausilia differentiates between C. from grisea (he had original material Deshayes) and a var. B. In the variation, finds he among other characters two palatal folds which makes it evident, that he also had true specimens of the A. bigibbosula. Kiister's C. grisea (1852: 81, pi. 9 figs. 1-4) was later O. C. schuchii Rossmassler. This recognised by Boettger (1878) to represent point of view can be followed as Kiister's figure 1 clearly shows the prominent subcolumellaris of this species which is characteristic of A. schuchii. Roth himself recognised his mistake in 1855 when reporting about his second trip the Orient. he attributed the maculosa to Now, erroneously name C. Deshayes to the in which species the second palatalis is missing and persists in using the name C. grisea for the species displaying this fold. This confusion could not be clarified by Martens used (1872: 43), who still the name C. grisea in Roth's sense. It is worth noting, that he mentions where this collected E. Nauplia (= Nafplion), species was by Raymond (the original lot is still housed in SMF). The whole situation in was re-investigated by O. Boettger 1878. He was the first who described the involved and who definitions of character clearly taxa gave exact states. He the first understand that the of was to position the lunella relative to the aperture and presence or absence of a basalis were useful criteria for species discrimination in this He that there which intermediate group. points out, are specimens seem to be but always can be separated by these characters (1878: 141). This point of view could be reconfirmed completely by the investigation presented here. Unfortunately, O. Boettger had no opportunity to check the original material of Deshayes. Otherwise he would easily have recognized the misinterpretation of preceding authors, in particular Roth. Fact that C. maculosa three later described Rossmassler is, Deshayes was years by under the Clausilia schuchii from name Greece. This is here proven by the illustration of the types of both taxa. The problem arises, that C. maculosa Deshayes pre-dates C. schuchii Rossmassler of and thus, according to the rule priority, C. schuchii is a junior of C. maculosa. As outlined the C. maculosa synonym before, name was frequently and used. erroneously Although the author supports the consequent use of the rule of it to be useful to the older C. maculosa in favour of C.