On altitude dependent characters in Albinaria idaea (L. Pfeiffer, 1849), with a revision of the species (Gastropoda Pulmonata: Clausiliidae) G.H. Engelhard & J.W.F. Slik Engelhard, G.H. & J.W.F. Slik. On altitude dependent characters in Albinaria idaea (L. Pfeiffer, 1849), with a revision of the species (Gastropoda Pulmonata: Clausiliidae). Zool. Med. Leiden 68 (3), 15.vii.1994: 21-38, figs. 1-31, 1 table.— ISSN 0024-0672. G.H. Engelhard & J.W.F. Slik, c/o Dept. of Mollusca, Nationaal Natuurhistorisch Museum, Postbus 9517, NL-2300 RA Leiden, The Netherlands. Key words: Gastropoda, Pulmonata, Clausiliidae, Albinaria, taxonomy, altitudinal variation, Greece. Several forms of Albinaria idaea (L. Pfeiffer, 1849) have been described as (sub)species from various sites on Mt. Ida (Central Crete). In recent years, however, much additional material has been collect- ed, which made clear that no sharp boundaries exist between these forms. Moreover, several charac- ters proved to be altitude dependent to a certain extent. It may be concluded that on Mt. Ida only a single evolutionary entity can be distinguished, viz. A. i. idaea. On the isolated Mt. Kedros, south-west of Mt. Ida, an undescribed allopatric form was found, which because of its smaller size and differenc- es in altitude dependent characters was given subspecific rank as A. i. amabilis ssp. nov. On the Paxi- madia Isles, south of Central Crete, a percostate and a finely ribbed subspecies occur allopatrically. The finely ribbed one seems consubspecific with A. i. idaea; the percostate one was recently described as the separate subspecies A. i. pichcapunlla Schultes & Wiese, 1991. Introduction In Europe, no land snail genus has radiated as strongly as Albinaria Vest, 1867. A wealth of species and subspecies is found from the Ionian islands to southern Asia Minor. For an understanding of the taxonomy and speciation scenarios of the species of Crete, where the genus is extremely speciose, detailed distributional data are high ly relevant. Until recently, little was known about the snails that occur south-west of Mt. Ida (2546 m), in the vicinity of the Mts. Kedros (1777 m), Siderotas (1136 m) and Vouvala (947 m). During an expedition to this area, a new, allopatric form of A. idaea was found on Mt. Kedros. While comparing the form from Mt. Kedros with the alleged subspecies of A. idaea from Mt. Ida, the existing classification seemed unsatisfactory. According to Nord- sieck (1977), the species is distributed over the latter mountain with four subspecies. However, many samples that have been collected on Mt. Ida in recent years cannot be classified with one of these subspecies, but seem to be intermediate in characters. This implies that either additional subspecies should be described or the diagnoses of currently recognized ones revised. This paper deals with these questions. Materials and methods In this analysis, the samples of A. idaea present in the Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie, = RMNH), Leiden were used. The RMNH inventory K-numbers and the corresponding UTM ( l xl km) 22 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) codes are listed below; ale. refers to material in alcohol 70%. Additional material from other collections is referred to with the following abbreviations: BM = British Museum (Natural History); HCN = Haus der Natur, Cismar; Ma = W. J. M. Maassen, Duivendrecht; NHMW = Naturhistorisches Museum, Wien; No = H. Nordsieck, Vil lingen-Schwenningen; PKI = Panepistimio Kritis, Iraklion; SMF = Senckenberg Museum, Frankfurt am Main; Su = P. Subai, Aachen. Whenever possible, ten specimens per sample were analysed. The holo- or lecto- types of all the nominal taxa within Albinaria idaea were also studied. The following characters were used: 1, number of whorls; 2, height of the shell; 3, width of the shell; 4, outside height of the aperture; 5, outside width of the aperture; 6, height of the parietalis, measured in front; 7, the shortest distance between the umbilical chink and the apertural margin, seen from the side, as a measure of protru sion of the aperture; 8, number of spiral ribs on the dorsal side of the penultimate whorl, measured across 2 mm; 9, number of cervical ribs along the suture, measured across 2 mm; 10, number of cervical ribs along the midline of the cervix, measured across 2 mm; 11, number of smooth apical whorls; 12, prominence of growth lines. For each character, mean values were calculated per sample. The geographical dis tribution of the characters was studied by plotting these averages on a geographical map of the area. The relation between characters and altitude was analysed by scat- terplots and Spearman rank correlation coefficients. Results In Albinaria idaea, three geographically isolated groups of populations can be dis tinguished (see fig. 1). One covers most of Mt. Ida, another is confined to a relatively small area on Mt. Kedros, and the third is found on the Paximadia Isles. On Mt. Ida, the species is quite variable. The populations on Mt. Kedros show little variation. On the Paximadia Isles, two distinct forms can be distinguished. On Mt. Ida the species occurs on the east, south, west and north-west slopes from approximately 300 to 1600 m above sea level. So far no specimens have been found on the highest parts of the mountain. The variability on Mt. Ida is great, but gradual for most measured characters. When compared with altitude (see table 1, figs. 2-12), the following characters show significant (p < 0.05) and positive correlations: width of the shell, height and width of the aperture, height of the parietalis, spiral and cer vical rib density. The spiral and cervical rib densities are the characters that are most clearly correlated with the altitude at which the snails in question occurred. On Mt. Kedros Albinaria idaea lives from 1200 m above sea level up to the summit (1777 m). Here no altitude dependent characters have been detected (see table 1). The samples have generally lower character values than Ida samples of similar altitudes (figs. 2-12). On the Paximadia Isles two allopatrically distributed forms of Albinaria idaea can be distinguished. One of these is percostate and is only found on limestone cliffs of the eastern part of the eastern islet. The other is finely ribbed and resembles the Mt. Ida forms; it lives all over both islets, on schist. The two forms are interconnected by a narrow hybrid zone (Schultes & Wiese, 1992). We have measured typical specimens of both forms from a single sample (NMW 80255), containing intermediate shells as well (figs. 29,30). Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) 23 Fig. 1. Map of central Crete, showing the location of the studied samples of Albinaria idaea. The 10 χ 10 km UTM grid and the 500,1000,1500 and 2000 m contour lines have been indicated. Records of other Albinaria species within this area have been omitted. 24 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) Table 1. Spearman rank correlation coefficients and corresponding p- values of shell characters with altitude in samples from Mt. Ida and Mt. Kedros. Significant correlations (p < 0.05) are marked with *. Mt. Ida Mt. Kedros 25 samples 9 samples number of whorls -0.2698 (P == 0.1862) 0.5588 (P == 0.1140) height of shell 0.2341 (P == 0.2514) 0.4352 (P == 0.2184) width of shell 0.6592 * (P == 0.0012) - 0.1841 (P == 0.6026) height of aperture 0.5237* (P == 0.0103) 0.0672 (P == 0.8492) width of aperture 0.6147* (P == 0.0026) - 0.0840 (P == 0.8121) height of parietalis 0.6484* (P == 0.0015) 0.1172 (P == 0.7404) distance umbilicus - apertural margin 0.2895 (P == 0.1561) 0.5607 (P == 0.1128) ribs/mm on penultimate whorl 0.7556 * (P == 0.0002) - 0.0084 (P == 0.9811) ribs/mm near suture of cervix 0.8805 * <P == 0.0000) - 0.3613 (P == 0.3068) ribs/mm near periphery of cervix 0.8369* (P == 0.0000) - 0.5021 (P == 0.1556) number of unribbed initial whorls 0.2309 (P == 0.2579) 0.1172 (P == 0.7404) The following list summarizes quantitative data concerning 35 samples (34 in RMNH, with Κ numbers; 1 in NNM) with altitudes of the localities in question. Abbreviations used: NW = number of whorls; HS = height of the shell (mm); WS = width of the shell (mm); HA = height of the aperture (mm); WA = width of the aper­ ture (mm); HP = height of the parietalis (mm); UA = distance from umbilicus to apertural margin (mm); RP = rib density of the penultimate whorl (per mm); RCS = rib density of the cervix along the suture (per mm); RCP = rib density of the cervix near the periphery (per mm); UIW = number of unribbed initial whorls; GL = promi­ nence of growth lines. For each character (except for GL) we give: mean ± standard deviation, minimum-maximum (number). The prominence of growth lines is indicat­ ed, giving numbers of specimens with or without this microsculpture; this distinc­ tion cannot always be made objectively, however. K1040 (550 m): NW = 12.8 ± 0.9,11-14 (10); HS = 19.8 ± 1.4,17.8-21.6 (10); WS = 4.1 ± 0.1, 4.0-4.3 (10); HA = 4.1 ± 0.3, 3.6-4.6 (10); WA = 3.3 ± 0.2,3.0-3.8 (10); HP = 0.30 ± 0.06, 0.19-0.38 (10); UA = 1.1 ± 0.1, 0.9-1.3 (10); RP = 4.6 ± 0.8, 3.7-5.8 (10); RCS = 3.0 ± 0.5, 2.6-3.7 (10); RCP = 2.6 ± 0.4, 2.1-3.1 (10); UIW = 1.6 ± 0.3,1.3-2 (10); GL = absent (10). K1042 (550 m): NW = 12.6 ± 0.5,12-13 (10); HS = 20.4 ± 1.0,18.8-22.6 (10); WS = 4.2 ± 0.2, 3.9-4.5 (10); HA = 4.2 ± 0.3, 3.8-4.5 (10); WA = 3.3 ± 0.3,2.7-3.6 (10); HP = 0.26 ± 0.08, 0.06-0.32 (10); UA = 1.1 ± 0.1, 0.9-1.2 (10); RP = 3.7 ± 0.4,3.1-4.7 (10); RCS = 2.4 ± 0.3, 2.1-3.1 (10); RCP = 2.0 ± 0.2,1.6-2.1 (10); UIW = 1.3 ± 0.2,1-1.5 (10); GL = absent (10). K1043 (650 m): NW = 12.7 ± 0.6,12-14 (10); HS = 19.4 ± 0.9,18.0-21.3 (10); WS = 4.0 ± 0.2, 3.6-4.4 (10); HA = 4.0 ± 0.3, 3.6-4.6 (10); WA = 3.2 ± 0.1, 3.0-3.4 (10); HP = 0.21 ± 0.04,0.14-0.28 (10); UA = 1.1 ± 0.1, 0.96-1.4 (10); RP = 4.3 ± 0.3,3.8-5.0 (10); RCS = 2.8 ± 0.2,2.5-2.9 (10); RCP = 2.2 ± 0.2,1.7-2.5 (10); UIW = 2.1 ± 0.4,1.5-3 (10); GL = present (9), absent (1). K1044 (400 m): NW = 12.2 ± 0.2,12-12.5 (3); HS = 18.9 ± 0.1,18.8-19.0 (2); WS = 3.9 ± 0.1, 3.8^.2 (7); HA = 3.5 ± 0.2, 3.2-3.7 (3); WA = 2.9 ± 0.1, 2.8-2.9 (2); HP = 0.09 ± 0.02, 0.06-0.12 (4); UA = 1.2 ± 0.1,1.1-1.3 (3); RP = 3.9 ± 0.4,3.1-4.2 (7); RCS = 2.0 ± 0.3,1.5-2.5 (7); RCP = 1.5 ± 0.3,1.0-1.9 (7); UIW = 1.3 ± 0.2,1- 1.5 (5); GL = present (8). K1078 (520 m): NW = 12.9 ± 0.2,12.5-13 (5); HS = 18.2 ± 0.6,17.2-18.8 (5); WS = 3.9 ± 0.2, 3.5-4.4 (10); HA = 3.7 ± 0.2, 3.4-3.9 (5); WA = 3.0 ± 0.1, 2.9-3.1 (5); HP = 0.12 ± 0.04, 0.07-0.16 (6); UA = 1.0 ± 0.07, 0.88-1.1 (7); RP = 4.3 ± 0.3,3.8-5.0 (9); RCS = 2.5 ±0.3,2.1-2.9 (9); RCP = 1.9 ± 0.2,1.5-2.1 (9); UIW = 1.4 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) 25 ± 0.2,1-1.5 (7); GL = present (8). K1079 (430 m): NW = 12.5 ± 0.4,12-13 (5); HS = 17.7 ± 0.8,16.4-18.4 (5); WS = 3.7 ± 0.1, 3.5-3.9 (9); HA = 3.6 ± 0.2, 3.4-4.0 (5); WA = 2.9 ± 0.1, 2.7-3.0 (4); HP = 0.09 ± 0.01, 0.08-0.10 (7); UA = 1.2 ± 0.2,1.1-0.4 (6); RP = 3.4 ± 0.4,2.9-4.2 (9); RCS = 1.7 ± 0.1,1.5-1.7 (6); RCP = 1.2 ± 0.2,1.0-1.3 (6); UIW = 1.4 ± 0.3,1-2 (8) ; GL = present (9). K1080 (350 m): NW = 12.3 ± 0.3,12-12.5 (2); HS = 17.8 ± 0.05,17.7-17.8 (2); WS = 3.6 ± 0.2, 3.5-3.9 (4); HA = 3.4 ± 0.05, 3.3-3.4 (2); WA = 2.5 ± 0.3, 2.2-2.7 (2); HP = 0.09 ± 0.02, 0.08-0.12 (3); UA = 1.1 ± 0.1, 1.0- 1.2 (3); RP = 3.9 ± 0.7,3.3-5.0 (4); RCS = 1.8 ± 0.2,1.5-2.1 (3); RCP = 1.5 ± 0.2,1.3-1.7 (3); UIW = 1 ± 0, 1-1 (3); GL = present (4). K1081 (1550 m): NW = 11.1 ± 0.7,10-12 (10); HS = 18.3 ± 1.2,16.3-20.2 (10); WS = 4.2 ± 0.1,3.9-4.3 (10); HA = 4.0 ± 0.5, 3.3-5.2 (10); WA = 3.3 ± 0.3, 2.6-3.6 (10); HP = 0.30 ± 0.09, 0.13-0.45 (10); UA = 1.0 ± 0.1, 0.9-1.3 (10); RP = 6.2 ± 0.9,5.2-8.4 (10); RCS = 4.6 ± 1.0, 3.7-6.3 (10); RCP = 3.2 ± 0.6, 2.6-4.7 (10); UIW = 2.3 ± 0.5,1.5-3.0 (10); GL = present (2), absent (8). K1084 (1270 m): NW = 12.5 ± 0.5, 12-13 (8); HS = 21.2 ± 1.7,18.1-23.6 (7); WS = 4.6 ± 0.3, 4.1-5.3 (10); HA = 4.3 ± 0.4,3.8-5.1 (9); WA = 3.5 ± 0.3,3.1-4.0 (9); HP = 0.31 ± 0.05,0.25-0.38 (9); UA = 1.2 ± 0.1,1.1- 1.3 (10); RP = 6.2 ± 1.0,4.7-8.4 (10); RCS = 4.2 ± 0.4,3.7-5.2 (10); RCP = 2.9 ± 0.4,2.1-3.7 (10); UIW = 1.8 ± 0.2,1.5-2.3 (8); GL = present (3), absent (7). K1192 (480 m): NW = 12.5 ± 0,12.5-12.5 (3); HS = 18.1 ± 0.4,17.7-18.4 (2); WS = 4.1 ± 0.2, 3.7-4.4 (6); HA = 3.8 ± 0.1, 3.6-3.9 (3); WA = 2.9 ± 0.2, 2.6-3.0 (3); HP = 0.15 ± 0.01, 0.14-0.16 (3); UA = 1.1 ± 0.1,1.0-1.3 (5); RP = 4.4 ± 0.6, 3.8-5.8 (7); RCS = 2.4 ± 0.3, 2.1-2.9 (6); RCP = 2.3 ± 0.3,1.9-2.9 (6); UIW = 2.1 ± 0.4, 1.5-2.5 (4); GL = present (5). K1193 (400 m): NW = 12.6 ± 0.6,12-13.5 (6); HS = 20.1 ± 0.7,18.9-20.4 (5); WS = 4.1 ± 0.2, 3.8-4.4 (10); HA = 3.8 ± 0.3,3.4-4.5 (8); WA = 2.9 ± 0.1,2.7-3.0 (8); HP = 0.15 ± 0.05,0.08-0.22 (8); UA = 1.0 ± 0.1,0.8- 1.1 (8); RP = 3.8 ± 0.3,3.3-4.2 (10); RCS = 2.2 ± 0.2, 2.1-2.5 (10); RCP = 1.7 ± 0.2,1.3-2.1 (10); UIW = 1.6 ± 0.5,1-2.5 (7); GL = present (10). K1196 (750 m): NW = 11.1 ± 0.6,10-12 (8); HS = 17.8 ± 0.6,16.8-18.7 (7); WS = 4.2 ± 0.2, 4.0-4.4 (9); HA = 3.9 ± 0.1, 3.8-4.1 (9); WA = 3.2 ± 0.1, 2.9-3.4 (9); HP = 0.29 ± 0.03, 0.25-0.32 (9); UA = 1.2 ± 0.2,1.0-1.6 (9) ; RP = 5.2 ± 0.6,4.2-5.2 (10); RCS = 2.8 ± 0.5,2.1-3.7 (10); RCP = 2.3 ± 0.2,2.1-2.6 (10); UIW = 1.3 ± 0.1, 1- 1.5 (8); GL = present (9). K1201 (860 m): NW = 12 ± 0,12-12 (3); HS = 17.8 ± 0.4,17.3-18.1 (3); WS = 3.9 ± 0.2, 3.6-4.1 (4); HA = 3.8 ± 0.3, 3.6-4.3 (4); WA = 2.8 ± 0.1,2.7-2.9 (4); HP = 0.22 ± 0.03,0.19-0.25 (4); UA = 1.1 ± 0.1,1.0-1.2 (4); RP = 6.2 ± 0.6, 5.2-6.8 (4); RCS = 4.1 ± 0.2, 3.7-4.2 (4); RCP = 3.4 ± 0.6, 2.6-4.2 (4); UIW = 1.1 ± 0.1,1-1.3 (3); GL = present (3), absent (1). K1203 (1000 m): NW = 12.9 ± 0.5,12-14 (10); HS = 19.4 ± 1.1,18.0-21.0 (10); WS = 4.4 ± 0.2,4.0-4.7 (10); HA = 4.0 ± 0.2, 3.8-4.3 (10); WA = 3.2 ± 0.1, 3.2-3.4 (10); HP = 0.18 ± 0.07, 0.06-0.32 (10); UA = 1.3 ± 0.1, 1.1- 1.5 (10); RP = 6.2 ± 0.6,5.2-7.3 (10); RCS = 3.9 ± 0.6, 3.1-4.7 (10); RCP = 2.4 ± 0.4, 2.1-3.1 (10); UIW = 1.4 ± 0.2,1-1.8 (10); GL = present (7), absent (1). K1218 (1050 m): NW = 12.7 ± 0.5, 12-13 (7); HS = 20.1 ± 0.6,18.8-21.0 (7); WS = 4.2 ± 0.2, 3.7-4.4 (10); HA = 39 ± 0.3, 3.5-4.5 (7); WA = 3.1 ± 0.2, 2.8-3.4 (8); HP = 0.19 ± 0.1, 0.13-0.32 (9); UA = 1.1 ± 0.2, 0.5- 1.3 (9); RP = 5.7 ± 0.7,4.7-6.8 (10); RCS = 3.5 ± 0.8, 2.6-5.2 (10); RCP = 3.0 ± 0.7, 2.1-4.2 (10); UIW = 1.4 ± 0.2,1.3-1.8 (7); GL = present (8), absent (1). K1219 (1150 m): NW = 12.5 ± 0.5,12-13 (2); HS = 17.4 (1); WS = 4.5 ± 0.4, 3.8-5.0 (4); HA = 3.9 ± 0.3,3.6- 4.3 (3); WA = 3.2 ± 0.2, 2.9-3.4 (3); HP = 0.21 ± 0.03, 0.19-0.25 (3); UA = 1.3 ± 0.1,1.1-1.4 (3); RP = 6.3 ± 0.4, 5.8-6.8 (4); RCS = 4.0 ± 0.3, 3.7-4.2 (4); RCP = 2.7 ± 0.2, 2.6-3.1 (4); UIW = 1.5 ± 0,1.5-1.5 (2); GL = present (4). K1220 (1130 m): NW = 12.4 ± 0.5,12-13 (7); HS = 19.5 ± 0.9,18.5-20.7 (7); WS = 4.5 ± 0.2, 4.2-4.7 (10); HA = 4.1 ± 0.1,3.9-4.4 (9); WA = 3.1 ± 0.3,2.6-3.7 (9); HP = 0.20 ± 0.06,0.13-0.32 (9); UA = 1.2 ± 0.1,1.1- 1.4 (9); RP = 5.8 ± 0.9,4.7-7.9 (10); RCS = 3.5 ± 0.5, 2.6-4.7 (10); RCP = 2.6 ± 0.4, 2.1-3.1 (10); UIW = 1.6 ± 0.3,1-2 (7); GL = present (9), absent (1). K1332 (750 m): NW = 12.3 ± 0.6,11-13 (9); HS = 18.0 ± 0.6,17.1-18.9 (6); WS = 3.9 ± 0.2,3.6-4.3 (10); HA = 3.5 ± 0.1, 3.4-3.8 (6); WA = 2.9 ± 0.1, 2.7-3.0 (6); HP = 0.18 ± 0.04,0.12-0.24 (8); UA = 1.0 ± 0.1, 0.88-1.2 (8); RP = 5.1 ± 0.4,4.2-5.8 (10); RCS = 2.6 ± 0.4,1.9-3.3 (10); RCP = 2.1 ± 0.3,1.7-2.5 (10); UIW = 2.6 ± 0.3, 2- 3 (6); GL = present (5), absent (2). 26 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) K1502 (1000 m): NW = 13 ± 0,13-13 (10); HS = 21.0 ± 1.2,19.8-24.1 (10); WS = 4.3 ± 0.3, 3.7-4.7 (10); HA = 4.2 ± 0.2, 3.9-4.4 (10); WA = 3.2 ± 0.1, 3.1-3.5 (10); HP = 0.19 ± 0.03, 0.13-0.25 (10); UA = 1.3 ± 0.1,1.2- 1.6 (10); RP = 5.2 ± 0.2,4.7-5.8 (10); RCS = 2.8 ± 0.4,2.1-3.7 (10); RCP = 2.2 ± 0.4,1.6-2.6 (10); UIW = 1.3 ± 0.2,1-1.5 (10); GL = present (10). K1545 (1400 m): NW = 11.9 ± 0.6,11-13 (7); HS = 18.3 ± 1.4, 16.6-20.5 (7); WS = 4.1 ± 0.2, 3.8-4.4 (10); HA = 3.8 ± 0.3, 3.3-4.4 (10); WA = 3.1 ± 0.3, 2.7-3.7 (10); HP = 0.28 ± 0.04, 0.25-0.38 (10); UA = 1.1 ± 0.1, 0.8-1.3 (10); RP = 6.1 ± 1.3,4.7-8.9 (10); RCS = 3.8 ± 0.5, 3.1-4.7 (10); RCP = 3.0 ± 0.6, 2.1^.2 (10); UIW = 2.0 ± 0.3,1.5-2.5 (7); GL = present (7), absent (3). K1546 (1400 m): NW = 11.8 ± 0.6,11-13 (10); HS = 19.9 ± 1.4,18.0-21.6 (10); WS = 4.4 ± 0.2,4.0-4.6 (10); HA = 4.1 ± 0.3, 3.6-4.6 (10); WA = 3.5 ± 0.2, 3.2-3.8 (10); HP = 0.31 ± 0.07, 0.19-0.44 (10); UA = 1.2 ± 0.1, 1.1-1.4 (10); RP = 4.8 ± 0.5,4.2-5.8 (10); RCS = 3.2 ± 0.3, 2.6-3.7 (10); RCP = 2.9 ± 0.5, 2.6-4.2 (10); UIW = 1.7 ± 0.4,1-2.3 (10); GL = present (8), absent (1). K2545 (1210 m): NW = 10.7 ± 0.5,10-11 (10); HS = 15.8 ± 0.7,14.9-16.8 (10); WS = 3.6 ± 0.1, 3.5-3.7 (10); HA = 3.5 ± 0.1, 3.3-3.7 (10); WA = 2.9 ± 0.1, 2.6-2.9 (10); HP = 0.13 ± 0.06, 0.032-0.25 (10); UA = 0.97 ± 0.07, 0.89-1.1 (10); RP = 5.6 ± 0.9, 4.2-7.3 (10); RCS = 4.1 ± 0.8, 2.6-4.7 (10); RCP = 3.2 ± 0.7, 2.1-4.2 (10); UIW = 2.0 ± 0.2,1.3-2.3 (10); GL = present (9). K2546 (1300 m): NW = 10.8 ± 0.6,10-11.5 (3); HS = 16.1 ± 1.3,14.3-16.6 (3); WS = 3.4 ± 0.1, 3.2-3.5 (6); HA = 3.4 ± 0.05, 3.3-3.4 (6); WA = 2.7 ± 0.2, 2.5-2.9 (7); HP = 0.12 ± 0.06, 0.04-0.24 (7); UA = 0.87 ± 0.08, 0.80-0.96 (7); RP = 4.3 ± 0.4,3.8-5.0 (7); RCS = 2.5 ± 0.4,1.7-2.9 (7); RCP = 2.2 ± 0.4,1.3-2.1 (7); UIW = 2.3 ± 0.2,2-2.5 (6); GL = present (7). K2547 (1400 m): NW = 12.0 ± 0.6,11-13 (10); HS = 16.6 ± 1.0,15.0-18.2 (10); WS = 3.4 ± 0.1, 3.3-3.6 (10); HA = 3.2 ± 0.1, 3.1-3.4 (10); WA = 2.7 ± 0.1, 2.6-2.8 (10); HP = 0.16 ± 0.03, 0.11-0.23 (10); UA = 0.97 ± 0.07, 0.88-1.1 (10); RP = 5.2 ± 0.6, 4.2-6.3 (10); RCS = 3.3 ± 0.7, 2.3-4.6 (9); RCP = 2.3 ± 0.5,1.7-3.3 (10); UIW = 2.5 ± 0.4,1.5-3 (10); GL = present (9). K2548 (1400 m): NW = 10.4 ± 0.5,10-11 (5); HS = 16.2 ± 0.9,14.9-17.5 (5); WS = 3.8 ± 0.1,3.7-4.0 (7); HA = 3.6 ± 0.2, 3.4-3.9 (5); WA = 3.1 ± 0.1, 3.0-3.3 (5); HP = 0.18 ± 0.06, 0.12-0.26 (5); UA = 0.95 ± 0.06, 0.88- 1.0 (5); RP = 6.4 ± 0.9,5.8-7.9 (7); RCS = 3.3 ± 0.2,2.9-3.7 (6); RCP = 2.2 ± 0.2,1.9-2.5 (6); UIW = 2.2 ± 0.2, 2-2.5 (6); GL = present (3), absent (2). K2549 (1480 m): NW = 11.1 ± 0.3,11-12 (10); HS = 15.6 ± 0.6,14.7-16.5 (10); WS = 3.5 ± 0.1, 3.4-3.8 (10); HA = 3.4 ± 0.1, 3.3-3.5 (10); WA = 2.9 ± 0.1, 2.6-3.0 (10); HP = 0.16 ± 0.06, 0.09-0.26 (10); UA = 0.90 ± 0.09, 0.80-1.04 (10); RP = 6.2 ± 0.8, 5.0-7.1 (10); RCS = 3.5 ± 0.8, 2.5-5.4 (10); RCP = 2.3 ± 0.3,1.7-2.9 (10); UIW = 2.3 ± 0.2,2-2.5 (10); GL = present (3), absent (7). K2551 (1700 m): NW = 11.8 ± 0.5, 11-12.5 (10); HS = 16.9 ± 1.1, 15.3-18.7 (10); WS = 3.5 ± 0.1, 3.4-3.6 (10); HA = 3.4 ± 0.2, 3.1-3.8 (9); WA = 2.8 ± 0.1, 2.6-3.0 (9); HP = 0.15 ± 0.06, 0.07-0.24 (10); UA = 0.96 ± 0.13, 0.88-1.20 (9); RP = 5.1 ± 0.9, 3.8-6.7 (10); RCS = 2.9 ± 0.7, 2.1-4.2 (10); RCP = 1.8 ± 0.4,1.3-2.3 (10); UIW = 2.1 ± 0.4,1.5-2.5 (10); GL = present (3), absent (4). K2552 (1777 m): NW = 11.8 ± 0.4,11-12 (6); HS = 16.2 ± 1.0, 14.8-17.3 (4); WS = 3.4 ± 0.1, 3.1-3.6 (10); HA = 3.5 ± 0.1, 3.3-3.6 (7); WA = 2.8 ± 0.2, 2.6-3.0 (6); HP = 0.14 ± 0.05, 0.04-0.24 (9); UA = 1.1 ± 0.1, 0.88-1.4 (10); RP = 6.0 ± 0.7,5.0-7.5 (10); RCS = 3.3 ± 0.3, 2.9-3.8 (9); RCP = 2.2 ± 0.3,1.7-2.5 (10); UIW = 2.3 ± 0.4,1.5-3 (8); GL = present (3), absent (2). K2553 (1600 m): NW = 11.3 ± 0.4,11-12 (8); HS = 16.4 ± 1.0, 15.3-18.1 (8); WS = 3.5 ± 0.2, 3.0-3.6 (10); HA = 3.5 ± 0.2, 3.2-3.8 (10); WA = 2.9 ± 0.2, 2.6-3.2 (10); HP = 0.14 ± 0.03, 0.08-0.20 (10); UA = 1.0 ± 0.1, 0.8-1.2 (10); RP = 5.0 ± 0.6,4.2-6.2 (10); RCS = 2.7 ± 0.4, 2.1-3.3 (10); RCP = 2.0 ± 0.3,1.5-2.5 (10); UIW = 2.4 ± 0.3,2-3 (8); GL = present (6), absent (2). K2554 (1500 m): NW = 12.0 ± 0.7,10.5-13 (10); HS = 17.6 ± 1.4,15.0-19.4 (10); WS = 3.6 ± 0.2, 3.3-3.9 (10); HA = 3.5 ± 0.2,3.2-3.8 (10); WA = 2.9 ± 0.2,2.6-3.2 (10); HP = 0.13 ± 0.04,0.09-0.19 (10); UA = 1.1 ± 0.1, 0.9-1.2 (10); RP = 5.5 ± 1.0, 4.2-7.9 (10); RCS = 3.2 ± 0.5, 2.3-3.8 (10); RCP = 2.1 ± 0.2, 1.7-2.5 (10); UIW = 2.4 ± 0.2,2-2.5 (10); GL = present(3), absent(4). K2583 (500 m): NW = 12.0 ± 0,12-12 (8); HS = 20.2 ± 0.9,18.6-21.6 (8); WS = 4.3 ± 0.3, 3.9-4.9 (10); HA = 4.3 ± 0.4, 3.9-4.8 (10); WA = 3.1 ± 0.2,2.9-3.4 (10); HP = 0.30 ± 0.06, 0.19-0.38 (10); UA = 1.0 ± 0.1, 0.8-1.1 (10); RP = 4.2 ± 0.5, 3.7-5.2 (10); RCS = 2.4 ± 0.3, 2.1-2.6 (10); RCP = 1.8 ± 0.2,1.6-2.1 (10); UIW = 1.6 ± 0.4,1.3-2.5 (6); GL = absent (10). K2584 (500 m): NW = 13.0 ± 0.7,12-14 (4); HS = 19.4 ± 0.4,19.1-20.0 (4); WS = 4.2 ± 0.2, 3.9-4.4 (4); HA Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) 27 = 4.0 ± 0.2, 3.8-4.3 (4); WA = 3.2 ± 0.2, 2.9-3.4 (4); HP = 0.37 ± 0.03, 0.32-0.38 (4); UA = 1.2 ± 0.1,1.1-1.2 (4); RP = 4.7 ± 0.5, 4.2-5.2 (4); RCS = 2.6 ± 0.4, 2.1-3.1 (4); RCP = 2.4 ± 0.3, 2.1-2.6 (4); UIW = 1.5 ± 0.2, 1.3-1.8 (4); GL = absent (4). K2597 (300 m): NW = 13.1 ± 0.6,12-14 (9); HS = 19.2 ± 0.8,17.6-20.1 (9); WS = 4.0 ± 0.3, 3.5-4.4 (10); HA = 3.7 ± 0.2, 3.4-4.0 (10); WA = 2.9 ± 0.1, 2.7-3.1 (9); HP = 0.06 ± 0.04, 0.02-0.16 (10); UA = 1.2 ± 0.1,1.0- 1.3 (10); RP = 4.5 ± 0.3,4.2-5.0 (10); RCS = 2.4 ± 0.3, 2.1-2.9 (10); RCP = 1.7 ± 0.1,1.5-1.9 (10); UIW = 1.8 ± 0.4,1.5-2.5 (9); GL = present (6), absent (4). K2601 (360 m): NW = 11.8 ± 0.5,11-12.5 (10); HS = 17.6 ± 0.8,16.2-19.2 (10); WS = 3.8 ± 0.1, 3.7-4.0 (10); HA = 3.6 ± 0.1, 3.4-3.8 (9); WA = 2.8 ± 0.1, 2.7-3.0 (9); HP = 0.13 ± 0.02, 0.08-0.16 (9); UA = 0.96 ± 0.08, 0.80-1.1 (9); RP = 5.2 ± 0.5,4.6-6.2 (10); RCS = 2.4 ± 0.2, 2.1-2.9 (10); RCP = 2.1 ± 0.2,1.7-2.5 (10); UIW = 1.6 ± 0.5,1-2.5 (10); GL = present (3), absent (4). NMW 80255, finely ribbed form: NW = 13.1 ± 1.0, 12-15 (7); HS = 17.8 ± 2.1, 16.0-22.3 (7); WS = 3.5 ± 0.5, 3.1-4.8 (10); HA = 3.5 ± 0.3, 3.1-4.2 (9); WA = 2.7 ± 0.3, 2.3-3.2 (8); HP = 0.20 ± 0.06, 0.13-0.31 (10); UA = 1.0 ± 0.1, 0.9-1.2 (10); RP = 3.7 ± 0.4, 3.2-4.3 (10); RCS = 1.9 ± 0.4,1.6-2.7 (10); RCP = 2.0 ± 0.6,1.6- 3.2 (10); UIW = 1.4 ± 0.2,1.3-1.8 (7); GL = present (4), absent (5). NMW 80255, percostate form: NW = 13.1 ± 0.6,12.5-14 (5); HS = 20.3 ± 0.7,19.6-21.4 (5); WS = 3.7 ± 0.2, 3.5-4.1 (10); HA = 3.9 ± 0.3, 3.5-4.3 (10); WA = 3.1 ± 0.2, 2.9-3.4 (9); HP = 0.22 ± 0.04, 0.16-0.25 (10); UA = 1.2 ± 0.1,1.0-1.6 (10); RP = 1.1 ± 0.1,1.0-1.4 (10); RCS = 0.7 ± 0.2, 0.4-1.0 (10); RCP = 0.7 ± 0.2, 0.4-1.3 (10); UIW = 2.1 ± 0.2,2-2.5 (4); GL = present (9). • • 0 200 400 600 800 1000 1200 1400 1600 1800 2 altitude (m) Figs. 2-12. Scatterplots showing the distribution of various shell characters in samples of Albinaria idaea in relation to the altitudes at which the samples were taken. Filled squares: A. i. idaea (Mt. Ida, Paximadia Isles); open squares: A. i. amabilis (Mt. Kedros); triangles: A. I pichcapunlla (Paximadia East Isle). For A. i. idaea a regression line is shown in cases where significant correlations between character state and altitude were found (see table 1). Note that A. I amabilis of Mt. Kedros does not fit well into the clinal variation of A. i. idaea. 28 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) 29 30 Engelhard & Slik. Albinaria idaea. Zool. Med. Leiden 68 (1994) 0.4 0.35 - Ε Ε. 0.3 m • (Λ m .1 0.25 CO Û. • * φ • • £ 0.2 -C m D ° Ο) φ • JZ 0.15 • 0.1 ••• 0.05 200 400 600 800 1000 1200 1400 1600 1800 altitude (m) 1.4 • • E 1.3 - E Φ V­ • • D 1.2 — •c φ α co ι CO 1.1 D • • 15 Ε Β ϋΦ • D • C CO t£o 0.9 ι ι Τ3 • 0.8 0 200 400 600 800 1000 1200 1400 1600 1800 8 altitude (m)