Lindholmiola corcyrensis ( ROSSMÄSSLER, 1838 )

Subai, Peter & Neubert, Eike, 2014, Revision of the genus Lindholmiola HESSE, 1931 (Gastropoda: Pulmonata: Helicodontidae), Contributions to Natural History 23, pp. 1-94 : 39-56

Lindholmiola corcyrensis ( ROSSMÄSSLER, 1838)

Figs 20–23 , 24–25 , 27 , 29

1828 Helix contorta , – Menke, Synopsis methodica molluscorum generum omnium et specierum earum, quae in Museo Menkeano adservantur: 10 (nomen nudum, homonym of Helix contorta LINNAEUS, 1758 ).

1837 Helix (Caracollina) corcyrensis BECK , Index molluscorum praesentis aevi musei principis augustissimi Christiani Frederici: 28 (nomen nudum).

1838 Helix (Helix) corcyrensis , – Anton, Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard Anton befinden: 39, No. 1428 (nomen nudum).

1838 Helix (Helix) canalifera ANTON , Verzeichniss der Conchylien welche sich in der Sammlung von Hermann Eduard Anton befinden: 39, No. 1427. Locus typicus: "Corfu" (nomen oblitum).

1838 Helix contorta [ H. corcyrensis ] ROSSMäSSLER, Iconographie der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der europäischen noch nicht abgebildeten Arten, (1) 2 (1/2): 40, Taf. 39, Fig. 538. Locus typicus: "Corfu". Type material: lost [(partim, not the variety minor = girva FRIVALDSZKY, 1835) (homonym of Helix contorta LINNAEUS, 1758 )].

1839 Helix corcyrensis , – Deshayes in Férussac & Deshayes, 1820–1851: Histoire naturelle générale et particulière des mollusques terrestres et fluviatiles, 1: 21 (Taf. 69 E, Fig. 1–5 published in, 1840).

1846 Helix corcyrensis , – L. Pfeiffer, In: Martini & Chemnitz, Systematisches Conchylien-Cabinet, (1) 12 (II): 103, Taf. 16 Fig. 23–24 .

1848 Helix corcyrensis , – L. Pfeiffer, Monographia heliceorum viventium, 1: 415.

1848 Helix corcyrensis var. γ. major, – L. Pfeiffer, Monographia heliceorum viventium, 1: 415.

1848 Helix canalifera , – L. Pfeiffer, Monographia heliceorum viventium, 1: 415.

1853 Helix corcyrensis , – L. Pfeiffer, Monographia heliceorum viventium, 3: 262.

1853 Helix canalifera , – L. Pfeiffer, Monographia heliceorum viventium, 3: 262.

1854 Helix corcyrensis , – Reeve, Conchologia Iconica, 7: species 1048.

1859 Helix corcyrensis , – Mousson, Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich, 4: 17, 256.

1859 Helix corcyrensis var. octogyrata MOUSSON , Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich, 4: 257.

1859 Helix corcyrensis var. cefalonica MOUSSON , Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich, 4: 29.

1859 Helix corcyrensis var. canalifera , – Mousson, Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich, 4: 17, 257.

1859 Helix corcyrensis var. girva , – Mousson, Vierteljahrsschrift der Naturforschenden Gesellschaft in Zürich, 4: 268 [non girva FRIVALDSZKY ].

1859 Helix corcyrensis , – L. Pfeiffer, Monographia heliceorum viventium, 4: 312.

1859 Helix canalifera , – L. Pfeiffer, Monographia heliceorum viventium, 4: 312.

1868 Helix corcyrensis , – L. Pfeiffer, Monographia heliceorum viventium, 5: 415.

1873 Helix corcyrensis var. canalifera , – Martens, Malakozoologische Blätter, 20: 33.

1876 Helix corcyrensis , – L. Pfeiffer, Monographia heliceorum viventium, 7: 467.

1879 Helix (Trigonostoma) corcyrensis , – Westerlund & Blanc, Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 36.

1879 Helix (Trigonostoma) corcyrensis var. cefalonica , – Westerlund & Blanc, Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 36.

1879 Helix (Trigonostoma) corcyrensis var. octogyrata , – Westerlund & Blanc, Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 37.

1879 Helix (Trigonostoma) corcyrensis var. canalifera , – Westerlund & Blanc, Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 37.

1879 Helix (Trigonostoma) corcyrensis var. girva , – Westerlund & Blanc, Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 37 [non girva FRIVALDSZKY ].

1882 Helix corcyrensis , – Hesse, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 9: 321.

1883 Helix (Anchistoma) corcyrensis , – O. Boettger, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 10: 316, 331.

1883 Helix (Anchistoma) corcyrensis cefalonica , – O. Boettger, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 10: 323.

1886 Helix (Gonostoma) corcyrensis var. octogyrata , – O. Boettger, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 13: 52.

1887 Helix (Caracolina) corcyrensis , – Tryon, Manual of Conchology, 3: 117.

1887 Helix (Caracolina) corcyrensis var. cephalonica , – Tryon, Manual of Conchology, 3: 118.

1887 Helix (Caracolina) corcyrensis var. octogyrata , – Tryon, Manual of Conchology, 3: 118.

1887 Helix (Caracolina) corcyrensis var. canalifera , – Tryon, Manual of Conchology, 3: 118.

1889 Helix (Caracolina) corcyrensis , – Westerlund, Fauna der in der paläarctischen Region lebenden Binnenconchylien, 2: 19.

1889 Helix (Caracolina) corcyrensis var. octogyrata , – Westerlund, Fauna der in der paläarctischen Region lebenden Binnenconchylien, 2: 19.

1889 Helix (Caracolina) corcyrensis var. cefalonica , – Westerlund, Fauna der in der paläarctischen Region lebenden Binnenconchylien, 2: 19.

1889 Helix (Caracolina) corcyrensis var. canalifera , – Westerlund, Fauna der in der paläarctischen Region lebenden Binnenconchylien, 2: 19.

1889 Helix (Caracollina) corcyrensis , – Martens, Archiv für Naturgeschichte, 55 (1): 172.

1891 Helix (Gonostoma) corcyrensis , – O. Boettger, Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 23 (5/6): 83.

1894 Helix (Gonostoma) corcyrensis , – Sturany, Annalen des Kaiserlich-Königlichen Naturhistorischen Hofmuseums, 9 (3): 371 [partim].

1894 Helix (Caracollina) corcyrensis , – Pilsbry, Manual of Conchology, 9: 288.

1894 Helix (Caracollina) corcyrensis var. cephalonica , – Pilsbry, Manual of Conchology, 9: 288.

1894 Helix (Caracollina) corcyrensis var. octogyrata , – Pilsbry, Manual of Conchology, 9: 288.

1894 Helix (Caracollina) corcyrensis var. canalifera , – Pilsbry, Manual of Conchology, 9: 288.

1905 Gonostoma corcyrensis , – Sturany, in Paganetti-Hummler: Verhandlungen der kaiserlich-königlichen Zoologisch-Botanischen Gesellschaft in Wien, 1905: 119.

1915 Helicodonta (Caracollina) contorta , – Sturany & Wagner, Denkschriften der Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse, 91: 31, Taf. 2, Fig. 7 , a–c.

1918 Caracollina corcyrensis , – Hesse, Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 50: 109.

1918 Caracollina corcyrensis octogyrata , – Hesse, Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 50: 109.

1918 Caracollina corcyrensis cephalonica , – Hesse, Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft, 50: 109.

1924 Helicodonta (Caracollina) contorta , – Polinski, Annales zoologici Musei Polonici Historiae Naturalis, 3 (3/4): 130.

1924 Helicodonta (Caracollina) contorta var. canalifera , – Polinski, Annales zoologici Musei Polonici Historiae Naturalis, 3 (3/4): 130.

1930 Caracollina contorta contorta , – Käufel, Sitzungsberichte der Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse, 139 (3/4): 183.

1930 Caracollina contortata cephalonica , – Käufel, Sitzungsberichte der Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse, 139 (3/4): 183.

1931 Lindholmiola corcyrensis , – Hesse, Zoologica, 31 (81) 1/2: 50.

1936 Lindholmiola corcyrensis , – Fuchs & Käufel, Archiv für Naturgeschichte, N.F. 5: 651. Fig. 79 [partim, only shells from Podgorica].

1941 Lindholmiola corcyrensis cephalonica , – Käufel & Fuchs, Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien, 88/89: 201.

1957 Lindholmiola corcyrensis , – Jaeckel, Klemm & Meise, Abhandlungen und Berichte aus dem Staatlichen Museum für Tierkunde in Dresden, 23 (2): 165 [partim].

1962 Lindholmiola corcyrensis , – Klemm, Sitzungsberichte der Österreichischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Abt. I, 171 (6/7): 252.

1962 Lindholmiola corcyrensis octogyrata , – Klemm, Sitzungsberichte der Österreichischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Abt. I, 171 (6/7): 253.

1962 Lindholmiola corcyrensis cephalonica , – Klemm, Sitzungsberichte der Österreichischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Abt. I, 171 (6/7): 253.

1962 Lindholmiola corcyrensis , – Jaeckel, Die Tierwelt Mitteleuropas, p.188 (partim).

1962 Lindholmiola corcyrensis , – Paget, Memorie di Biogeografia Adriatica, 4: 182, 184–185, 195, Fig. 3–5 [shell].

1965 Lindholmiola corcyrensis , – Forcart, Verhandlungen der Naturforschenden Gesellschaft in Basel, 76 (1): 141.

1971 Lindholmiola corcyrensis , – Alzona, Atti della Società Italiana di Science Naturali e del Museo Civico di Storia Naturale di Milano, 111: 185.

1973 Lindholmiola (Lindholmiola) corcyrensis cephalonica , – Hudec & Vašátko, Acta scientiarum naturalium Academiae Scientiarum Bohemoslovacae, 7 (9): 13.

1989 Lindholmiola corcyrensis , – Reischütz & Stummer, Malakologische Abhandlungen aus dem Staatlichen Museum für Tierkunde Dresden, 14 (12): 106.

1990 Lindholmiola corcyrensis , – Reischütz & Sattmann, Annalen des Naturhistorischen Museums in Wien, 91 (B): 258.

1990 Lindholmiola girva , – Fechter & Falkner, Steinbachs Naturführer: 222, Abb. 6 (partim) (non girva FRIVALDSZKY, 1835 ).

1996 Lindholmiola girva , – Welter-Schultes, Schriften zur Malakozoologie, 9: 23–31 (non girva FRIVALDSZKY, 1835 ).

1996 Lindholmiola girva , – Dhora & Welter-Schultes, Schriften zur Malakozoologie, 9: 157, 190, Taf. 13, Fig. 165–166 (non girva FRIVALDSZKY, 1835 ).

2002 Lindholmiola girva , – Dhora, Studime mbi molusqet e Shqipërisë: 99, 156–158 (non girva FRIVALDSZKY, 1835 )

Diagnosis: shell depressed, shell surface covered by granulation, whorls slightly shouldered, penial lumen with a number of small, v-shaped pilasters only.

Description of shell: shell depressed, spire only slightly raised, shell colour light to chestnut brown; first ½ to ¾ protoconch whorl smooth, then with a few fine radial stripes; the following teleoconch whorls with increasingly stronger stripes, but often reduced on the penultimate and last whorls; stripes may vary in strength to even form small riblets; on the subsurface of the shell, stripes usually weak and blurred; complete shell surface covered by a fine granulation giving the shell a dull appearance; sometimes with small hair warts (mainly on subsurface); shell covered by hairs, hairs in the umbilicus usually shorter than those on the rest of the shell, hair length at the shell's shoulder ca 0.4–0.7 mm; 5¾–7¾ regularly increasing, slightly shouldered whorls; last whorl slightly descending before the aperture, then abruptly curved downwards, the bent part in a brighter to white colour; suture deep; umbilicus of oval shape, cylindrical to slightly perspective, only slightly increasing at the beginning, with a rapid, almost double increase in the last whorl, reaching a diameter of 1.4–3.75 mm; aperture oblique, ear-shaped, apertural insertion with a distance of 2.3–5 mm, connected by a very thin, almost indiscernible callus; peristomial rim sharp, forming a strong lateral and weaker basal lip, with a small peristomial callus covering ca 1/10 of the umbilicus.

Measurements: H: 3.75–7.5; D: 7.8–15.9; aH: 3.4–7.2 (2–4.7); aW: 3.6–7.8.

Details of body (8 animals; eastern boundary of Stroúni/1, Island Lefkáda, Evgiros/3, Island Kefallinia, 1.5 km E of Sámi/4): head and dorsal body cream to grey or brown, with many intermediate colourations, becoming brighter in direction to the tail; foot sole cream to beige; mantle with small, elongate grey dots; secondary ureter opens 1–1.5 mm before the pneumostome, where it splits into two canals.

Morphology of the genital organs ( Fig. 27 ): penis short, slightly inflated, ca half of its length is covered by penial glandular tissue, penial lumen with a number of small, v-shaped pilasters; vagina relatively long, glandula narrow, scarcely folded, reaching 2/3 of the length of the bursa copulatrix, pedunculus and vesicle of the bursa copulatrix of the same length; atrial lumen filled by two to three elevated and elongated pilasters with a few short secondary pilasters running in parallel, the major pilasters unite to form a large single pilaster filling almost the complete vaginal lumen.

Differential diagnosis: Conchologically, this species is close to L. pirinensis . However, it differs from that species by its more flattened whorls, its shallow suture, and its less pronounced surface sculpture. Its radial riblets are quite regular, and its hairs are longer (although less permanent). As in L. pirinensis , the aperture is only slightly bent downwards, but in frontal view, the aperture of L. pirinensis is broader and larger. The apertural rim of L. corcyrensis is only slightly reinforced, and the umbilical shield less pronounced. In L. pirinensis ( Fig. 26 ), the penial lumen shows only two large pilasters with one of them curved backwards forming a third, smaller pilaster.

Lindholmiola girva is quite similar, but the shell is smaller on average with more flattened whorls. However, in this species, the teleoconch whorls are more distinctly shouldered with the radial riblets being more pronounced. The last whorl is narrower. The umbilicus formation in L. girva is similar, but here it is less perspective (almost circular) and not oval as in L. corcyrensis . It aperture is narrower and sometimes shows a tooth-like swelling. As in L. pirinensis , L. girva shows two to three elongated penial pilasters, and thus can easily be distinguished from L. corcyrensis .

Similarly sized shells of L. barbata have a half to one additional whorl, L. corcyrensis is clearly more densely coiled. In addition, the shell of L. barbata is covered by small hair warts, its umbilicus is partially to completely covered by a columellar shield, and the aperture is less bent downwards with a weaker labial callus. The penial lumen of L. barbata shows two parallel elongated pilasters and thus differs profoundly from the v-shaped pilasters of L. corcyrensis .

Type specimens: Greece, Kefallinia Island, "aux environs d’Argostoli,...à Frangata et Grisata", 38.17°N 20.49°E, syntypes var. cefalonica : ZMZ 508737/6 ; "aux environs de Préveza ", 38.96°N 20.75°E, syntypes var. octogyrata : ZMZ 508745/5 .

Additional specimens examined:

Croatia, Pelješac–Peninsula , sea floating debris near Prapatna, UTM YH 14 , 42 .81°N 17.67°E, leg. Maassen 4.1989, Maa /6 ; "Dalmatien, Zirona Grande", UTM n. det., SMF 284844 /2 ; Badija Island (= Sv. Scoglia Badia) near Korčula Island, sea floating debris, UTM XH 75 , 42 .95°N 17.15°E, SMF 207804 /1.

Italy, Prov. Foggia, Promontorio del Gargano, San Menáio, UTM WG 74, 41.93°N 15.94°E, ( Paget, 1962: 182; Forcart, 1965: 141) ; Promontorio del Gargano, Péschici, UTM WG 84, 41.94°N 16.01°E, ( Forcart, 1965: 141).

Montenegro, fortress near Vranjina, UTM CM 48 , 42 .27°N, 19.11°E, leg. Maassen 5.1984, Maa/1 ; Busovnik (= Mountain W of Podgorica), UTM CM 49 , 42 .44°N, 19.14°E, SMF 284837 /3, 284865/1 ; Ulcinj, UTM CM 54 , 42 .93°N, 19.21°E, leg. Podani 20.8.1976, HNHM 21472 /1 (juv.) ; Podgorica (= Titograd), UTM CM 59 , 42 .44°N, 19.26°E, SMF 101901 /11 ; Šasko Lake , castle ruins, UTM CM 64 , 41 .97°N, 19.33°E, leg. Erőss & Fehér leg. 20.4.2000, HNHM 85624 /8 ; Spuž, UTM CN 50 , 42 .51°N, 19.19°E, leg. Wohlberedt, 1933, SMF 101900 /4 ; Zelenik (= Zelnik ), UTM n. det., leg. Wohlberedt, 1905, SMF 284836 /2, 284866/2.

Albania, valley of Povlë River S of Greshicë (= 15 + 1 km SE of Ballsh), approx. 140–200 m alt., UTM CK 98 , 40 .54°N, 19.77°E, leg. Erőss, Fehér & Kovács 14.4.2001, HNHM 85619 /4 ; Durrës (= Durazzo), UTM CL 67–77 , 41 .32°N, 19.44°E, SMF 207808 /3 ; 24.9.1959, HNHM 83641 /1 ; beach of Durrës, UTM CL 77 , 41 .31°N, 19.47°E, leg. Clauss, SMF 207805 /3 ; Krujë , on castle walls, UTM CL 99 , 41 .5075°N, 19.7933°E, leg. Erőss & Fehér 2.7.1996, HNHM 85135 /1 ; Shkodër (= Skutari), UTM CM 75 , 42 .07°N, 19.52°E, 1905, SMF 101928 /11 ; leg. Barbanic, 1905, SMF 284873 /15 ; leg. Wohlberedt, 1905, SMF 284863 /1 ; leg. Flasch, 1906, SMF 284871 /10 ; leg. Kormos, 1907, SMF 101926 /4 ; Shkodër, castle hill, UTM CM 75 , 42 .0475°N, 19.4936°E, leg. Erőss & Fehér 1.7.1996, HNHM 85140 /1 ; leg. Erőss & Fehér & Kovács 7.4.2001, HNHM 85623 /3 ; Vau i Dejes (= approx. 20 km SE of Shkodër), UTM CM 85 , 42 .00°N, 19.63°E, leg. Erőss & Fehér 2.7.1996, HNHM/1 ; Pllanë (= N of Milot), 4 km E from bridge of Mat River , approx. 100 m alt., UTM CM 91 , 41 .70°N, 19.72°E, leg. Erőss & Fehér & Kovács 9.4.2001, HNHM 85620 /3 ; rock walls at Borsh, UTM DK 03 , 40 .06°N, 19.85°E, leg. Erőss & Fehér 6.7.1996, HNHM 85142 /1 ; Tepelenë , on castle walls, UTM DK 16 , 40 .29°N 20.02°E, leg. Erőss & Fehér 6.7.1996, HNHM 85141 /1 ; leg. Erőss & Fehér & Kovács 14.4.2001, HNHM 85637 /1 ; Tepelenë, Lavdi Deshmorevë cemetery on the mountain, approx. 260 m alt., UTM DK 16 , 40 .29°N 20.02°E, leg. Erőss 17.8.1993, S, 18263/2 ; leg. Erőss & Fehér & Kovács 14.4.2001, HNHM 85628 /7 ; 4 km S of Tërpan (= 28 km from Berat direction Këlcyrë ), approx. 700 m alt., UTM DK, 18, 40.54°N 20.00°E, leg. Erőss & Fehér & Kovács 13.4.2001, HNHM 85625 /3 ; 4 km N of Tërpan (= 20 km from Berat direction Këlcyrë ), approx. 700 m alt., 40.57°N 20.01°E, UTM DK, 19, leg. Erőss & Fehér & Kovács 13.4.2001, HNHM 85636 /2 ; left bank of Dëshnicë River 5 km N of Këlcyrë, approx, 270 m alt., UTM DK 26 , 40 .35°N 20.16°E, leg. Erőss & Fehér & Kovács 13.4.2001, HNHM 85621 /8 ; Osum Gorge 4 km SE of Çorovodë (= direction Zogas), approx. 400 m alt., UTM DK 38 , 40 .47°N 20.25°E, leg. Erőss & Fehér & Kovács 12.4.2001, HNHM 85622 /1 ; Mal e Melesinit (= 6 km S of Leskovik), UTM DK 64 , 40 .10°N 20.59°E, leg. Erőss & Fehér 7.7.1996, HNHM 85132 /1 ; Leskovik (= Lescovic ), 800 m alt., UTM DK 64 , 40 .14°N 20.59°E, SMF 207807 /1 + 1 (juv.) ; Mal i Dajtit , path direction relais station, on rocks, approx. 1200 m alt., UTM DL 07 , 41 .36°N, 19.91°E, leg. Erőss & Fehér 4.7.1996, HNHM 85131 /1 ; Mal i Dajtit , 1 km above Linzë, UTM DL 07 , 41 .36°N, 19.88°E, leg. Fehér 27.7.1993, HNHM 85136 /4 ; Tiranë River Valley 1 km N from Turkish bridge between Ferraj and Shtish, UTM DL 08 , 41 .39°N, 19.86°E, leg. Erőss & Fehér & Kovács 10.4.2001, HNHM 85630 /3 ; Tiranë River Valley 4 km N from Turkish bridge between Ferraj and Shtish, UTM DL 08 , 41 .43°N, 19.85°E, leg. Erőss & Fehér & Kovács 10.4.2001, HNHM 85633 /2 ; Berat, castle hill SW slope, approx. 100 m alt., UTM DL 10 , 40 .70°N, 19.94°E, leg. Erőss & Fehér & Kovács 12.4.2001, HNHM 85627 /7 ; Berat, left bank of Osum River opposite castle hill, approx. 100 m alt., UTM DL 10 , 40 .70°N, 19.94°E, leg. Erőss & Fehér & Kovács 13.4.2001, HNHM 85626 /5 ; S of Gracen (= Elbasan – Tiranë country road, 7 km N from bifurcation to Cërrik), approx. 500 m alt., UTM DL 15 , 41 .14°N, 19.96°E, leg. Erőss & Fehér & Kovács 14.4.2001, HNHM 85632 /2 ; Erzen Gorge NW of Ibë, approx. 300 m alt, UTM DL 16 , 41 .25°N, 19.91°E, leg. Erőss & Fehér & Kovács 11.4.2001, HNHM 85635 /2 ; hills S of Ibë , UTM DL 16 , 41 .23°N, 19.93°E, leg. Erőss & Fehér 15.9.1994, HNHM 85137 /2 ; Fangul Pass (= 9 km from Ibë direction Kllojkë), approx. 700 m alt., UTM DL 16 , 41 .27°N 20.05°E, leg. Erőss & Fehér & Kovács 11.4.2001, HNHM 85631 /1 ; Elbasan, UTM DL 25 , 41 .11°N 20.09°E, leg. Fehér 23.7.1993, HNHM 85039 /1 ; Mirakë, Ura e Kamarës , left bank of Shkumbin River , approx. 210 m alt., UTM DL 36 , 41 .16°N 20.24°E, leg. Erőss & Fehér & Kovács 14.4.2001, HNHM 85629 /1 ; on rocks above Tushemist at Ohrid Lake , UTM DL 72 , 40 .90°N 20.72°E, leg. Erőss & Fehér 7.7.1996, HNHM 85138 /1 ; Pustec, bank of Prespa Lake , UTM DL 91 , 40 .79°N 20.91°E, leg. Fehér, 18.9.1992, HNHM 85139 /1 ; Djelcic, UTM n. det., SMF 284845 /1.

Fyrom, Crni Drim River left bank, N of Lukovo, on limestone rocksand in the soil, 630 m alt., UTM DL 67 , 41 .3544°N 20.6107°E, leg. Pintér, E. & P. Subai & Szigethy 17.07.1972, HNHM 49545 /1 (juv.) ; Crni Drim River left bank, some km S of Lukovo, 630 m alt., UTM DL 67 , 41 .3398°N 20.6282°E, leg. Pintér, E. & P. Subai & Szigethy 17.07.1972, HNHM 49543 /1, 85618/1, S 3848/1 ; 1 km SE of Džepišta (= direction Lukovo ), 700 m alt., UTM DL 68 , 41 .4432°N 20.5374°E, leg. Pintér, E. & P. Subai & Szigethy 17.07.1972, HNHM 49544 /1, S 3847/1 ; Sv.Naum, under stones, UTM DL 72 , 40 .9141°N 20.7412°E, leg. Maassen 5.1976, Maa/27, NNM/3 ; Galičica Mountains , in forest 10 km from the crossing Carina– Sv. Naum – Ohrid direction Carina, on limestone rocks, 1560 m alt., UTM DL 83 , 40 .9618°N 20.8165°E, leg. Pintér, E. & P. Subai & Szigethy 14.07.1972, HNHM 49542 /1 (juv.), S 3845/1 ; Galičica Mountains , pass 17 km from bifurcation at Trpejca direction Oteševo, 1600 m alt., UTM DL 83 , 40 .9553°N 20.8146°E, leg. Maassen 5.1976, Maa/16 ; at boat harbour of Trpejca , on limestone rocksand from Lake Ohrid, 700 m alt., UTM DL 83 , 40 .9584°N 20.7781°E, leg. Pintér, E. & P. Subai & Szigethy 16.07.1972, HNHM 49533 /5 + 4 (juv.+ fragment), 85634/1, S 3841/6 + 1 (juv.) ; rocks N of boat harbour of Trpejca , UTM DL 83 , 40 .9591°N 20.7781°E, leg. Maassen 5.1975, Maa/1 ; Sv.Naum – Ohrid highway, bifurcation at Trpejca , UTM DL 83 , 40 .9439°N 20.7895°E, leg. Kiss & Pintér 4.7.1985, HNHM 30854 /1 + 4 (juv. + fragment) ; Trpejca , UTM DL 83 , 40 .96°N 20.78°E, leg. Kiss & Pintér 5.7.1985, HNHM 30855 /1 ; between Trpejca and Peštani, UTM DL 83 , 40 .98°N 20.80°E, leg. Kiss & Pintér 5.7.1985, HNHM 30856 /1 ; on limestone rocks at shore of Lake Ohrid approx. 2.5 km S of Peštani, 710 m alt., UTM DL 83 , 40 .9981°N 20.8007°E, leg. Pintér, E. & P. Subai & Szigethy 16.07.1972, HNHM 49531 /1 + 4 (juv.), S 3840/3 ; leg. Maassen 5.1975, Maa/1 ; 1 km S of Peštani, 800–1000 m alt., UTM DL 83 , 40 .9980°N 20.8025°E, leg. Maassen 5.1975, Maa/10 ; bank of Ohrid Lake S of Peštani, on limestone rocks, UTM DL 83 , 41 .0065°N 20.8050°E, leg. Pintér, E. & P. Subai & Szigethy 16.07.1972, HNHM 49534 /1 + 1 (juv.), S 3843/3 ; Galičica Mountains W side, on limestone rocks in "Elešec" camping, 4 km N of Peštani and of Lake Ohrid before the camping, 750 m alt., UTM DL 84 , 41 .0386°N 20.8079°E, leg. Subai 11.–14.08.1971, S 3846/2 ; leg. Pintér, Subai & Szigethy 15.7.1972, HNHM 49532 /3 + 4 (juv.), S 3844/2 ; above Sveti Stefan (= S of Ohrid), on rocks, 1100 m alt., UTM DL 84 , 41 .0757°N 20.8289°E, leg. Maassen 5.1975, Maa /2 ; tunnel near Orce Nikolov (= 6.5 km S of Ohrid), UTM DL 84 , 41 .05°N 20.80°E, leg. Maassen 5.1975, Maa /1 ; mountain slope at E border of Ohrid, UTM DL 85 , 41 .13°N 20.85°E, leg. Mikuska & Sipos 2.10.1975, HNHM 49549 /1 (juv.).

Greece, Macedonía, 300 m N of Pefkófito , moist place in a small river gorge, approx. 1000 m alt., UTM DK 96 , 40 .2972°N 20.9566°E, leg. Subai 10.05.1995, S 15826 /1 ; E slope of the mountain W of Pefkófito , on dry lime rocks, about 1200–1400 m alt., UTM DK 96 , 40 .2967°N 20.9433°E, leg. Subai 10.05.1995, S 15823 /2 ; mountain E slope between Epáno and Káto Aréna (= W of Pefkófito), on limestone rocks, 1650 m alt., UTM DK 96 , 40 .2948°N 20.9381°E, leg. Subai 11.05.1995, S 15840 /5 ; Grámmos Mountains, E side of Epáno Aréna mountain (= NW of Pefkófito), near the crest, 2000 m alt., UTM DK 96 , 40 .3455°N 20.8621°E, leg. Subai 15.07.1990, S 15872 /4 ; Epáno Aréna Mountain (= NW of Pefkófito), mountain slope and ridge, on limestone rocks and under stones, 1750–2100 m alt., UTM DK 96 , 40 .3140°N 20.8972°E, leg. Subai & Szekeres 17.05.1997, S 15848 /1 ; 15 km SW von Grevená, Landstrasse Zákas–Spíleo, an Kalkfelsen, 925 m alt., UTM EK 23 , 40 .0104°N 21.2832°E, leg. Gittenberger & Uit de Weerd 11.5.2000, NNM/4 ; Kastoriá , N part of peninsula, 620 m alt., UTM EK 28 , 40 .52°N 21.27°E, leg. Neuteboom 28.7.1979, NNM/1 ; Kastoriá, Monastir Panagia Mavriotissa , UTM EK 28 , 40 .5043°N 21.2804°E, leg. Neuteboom 28.7.1979, NNM/4 ; on the peninsula in Kastoriá , on limestone rocks approx. 50–100 m W of Monastir Panagia Mavriotissa, 640 m alt., UTM EK 28 , 40 .5050°N 21.2786°E, leg. Subai 14.07.1990, S 15877 /46 ; at bridge 4 km SE of Vogatsikó , on limestone rocks, 680 m alt., UTM EK 37 , 40 .3983°N 21.4143°E, leg. Subai 10.05.1995, S 15824 /28 ; in valley W of Gérmas , on limestone rocks, 850 m alt., UTM EK 37 , 40 .4528°N 21.4149°E, leg. Subai & Szekeres 17.05.1997, S 15849 /51 ; Mikrá Prespa Lake , behind the biological station, approx. 200 m W of the W border of Mikrolímni, on limestone rocksand in a stone quarry, 885 m alt., UTM EL 01 , 40 .7419°N 21.1078°E, leg. Subai 13.07.1990, S 15878 /2 ; leg. Subai leg. 10.5.1995, S 15825 /2. — Thessalía, 3–4 km from Diáva towards Koromiliá (= 4 km SW of Kalambáka), additional 400 m on a right path from the bifurcation, on limestone rocks, 550 m alt., UTM EJ 49 , 39 .68°N 21.58°E, leg. Gittenberger & Uit de Weerd 13.5.2000, NNM/3 ; NW border of Píli (= 18 km SW of Tríkala), on rocks at both sides of the Turkish arched bridge, 240 m alt., UTM EJ 56 , 39 .4597°N 21.6121°E, leg. Pintér, E. & P. Subai 28.07.1976, S 3812/1 ; Theópetra (= Kuvelci , 6 km SSE of Kalambáka), on limestone blocks of rocks at river bank, 260 m alt., UTM EJ 59 , 39 .6798°N 21.6810°E, leg. E. & P. Subai 26.07.1975, S 3813/1 ; Kalambáka, on conglomerate rocks beside the highway from Kalambáka to the Meteóra monasteries, 280 m alt., UTM EJ 59 , 39 .7103°N 21.6164°E, leg. Pintér, E. & P. Subai 29.07.1976, S 3814/1 ; Kastráki near Kalambáka , UTM EJ 59 , 39 .71°N 21.62°E, leg. Neuteboom leg. 25.7.1979, NNM/2 ; at S border of Dafnospiliá , on limestone rocks beside the cave, 260 m alt., UTM EJ 84 , 39 .2274°N 21.9427°E, leg. Subai & Szekeres 12.05.1997, S 15814 /2. — Kérkira Island (= Korfu), Paleokastrítsa , limestone rocks on elevations, 30 m alt., UTM CJ 89 , 39 .6708°N, 19.7019°E, leg. E. & P. Subai, 18.07.1975, S 3822/5 ; Neuteboom 6.10.1991, NNM/10 + 1 (juv.) ; Paleokastrítsa, Villa Reale, UTM n. det., SMF 284829 /5 ; Pélekas (= Pelleka), UTM CJ 98 , 39 .59°N, 19.82°E, SMF 101933/2 ; on limestone rocks at S border of Ermónes bay, in rock debris, UTM CJ 98 , 39 .6086°N, 19.7758°E, leg. Subai 03.08.1995, S 15843 /3 ; Vátos, bei Monastir Mirtiotíssa, UTM CJ 98 , 39 .5972°N, 19.7936°E, leg. Neuteboom 2.10.1991, NNM/1 ; Rand von Sokráki in Richtung Zigós , UTM CJ 99 , 39 .7168°N, 19.7977°E, leg. Neuteboom 3.10.1991, NNM/4 + 4 (juv.) ; Auffahrtweg zum Pantokrátor, 700 m alt., UTM CJ 99 , 39 .7499°N, 19.8428°E, leg. Neuteboom 3.10.1991, NNM/1 ; Áno Korakiana, UTM CJ 99 , 39 .70°N, 19.79°E, leg. Neuteboom 3.10.1991, NNM/1 (juv.) ; mountain approx. 1 km NW of Strinilas , on limestone rocks, 580 m alt., UTM CJ 99 , 39 .7464°N, 19.8294°E, leg. Subai 07.08.1995, S 15841 /3 ; Análipsis (= Analepsis), UTM CJ 99 , 39 .70°N, 19.82°E, SMF 284827 /1 ; Ágios Márkos, UTM CJ 99 , 39 .71°N, 19.82°E, leg. Maassen 3.7.1980, Maa /1 ; 500 m E of Troumpéta (= Pandeleimonas ), N exposed rock holes at border of highway and on S exposed limestone rocks, 460 m alt., UTM CJ 99 , 39 .7092°N, 19.7531°E, leg. Subai 06.08.1995, S 14275 /1 (juv.) ; in olive grove 1 km N of Nímfes , on limestone rocks beside the highway, 150 m alt., UTM CK 90 , 39 .7609°N, 19.7829°E, leg. Subai 01.08.1995, S 15837 /31 ; 1.2 km S of Ág. Panteleimonas (= direction Episkepsis ), on limestone rocks beside the highway, 240 m alt., UTM CK 90 , 39 .7734°N, 19.8267°E, leg. Subai 01.08.1995, S 15836 /8 ; from Ágios Martínos direction Acharávi , UTM CK 90 , 39 .78°N, 19.83°E, leg. Neuteboom 3.10.1991, NNM/1 ; near road between Xathátes and Nímfes , UTM CK 90 , 39 .75°N, 19.77°E, leg. Neuteboom 4.10.1991, NNM/1 (juv.) ; 1.2 km ESE of Akharave , on corrugated karst rocks, 110 m alt., UTM CK 90 , 39 .7905°N, 19.8336°E, Subai 30.07.1995, S 15844 /25 ; Ahillio, UTM DJ 07 , 39 .56°N, 19.90°E, SMF 101932 /1 ; gegenüber Ahillio, UTM DJ 07 , 39 .56°N, 19.90°E, SMF 101930 /2 ; Gardíki, byzantinische Festung, UTM DJ 07 , 39 .4770°N, 19.8848°E, leg. Neuteboom 12.10.1991, NNM/1 + 2 (juv.) ; 1.5 km N von Benítses , UTM DJ 07 , 39 .55°N, 19.91°E, leg. Gittenberger 9.1996, NNM/2 (alk.) ; 2 km S of Benitzes , on rocks in the vicinity of the sea, 20 m alt., UTM DJ 07 , 39 .5309°N, 19.9203°E, leg. E. & P. Subai, 19.07.1975, S 3818/5 ; "Santi Deka", UTM DJ 07 , 39 .55°N, 19.88°E, SMF 101934 /2 ; Ag. Déka Mountain (= W of Ag. Déka Village), on limestone rocks and under stones, 290 m alt., UTM DJ 07 , 39 .5548°N, 19.8819°E, leg. E. & P. Subai, 19.07.1975, S 3817/5 ; Kérkira Town (= Korfu), UTM DJ 08 , 39 .62°N, 19.91°E, HNHM 49548 /1, SMF 284821 /11, 284822/2, 284823/5, 284824/3, 284825/1, 284826/1, 284828/8 (juv.), 284830/1, 284840/12, 284841/9, 285672/2, 285673/1 ; Brömme leg., SMF 284848 /1 ; leg. C. R. Boettger, 1906, SMF 101941 /6 ; leg. Conéménos, SMF 101937 /5 ; Krüper, 1883, SMF 101939 /1 ; Stadt Kérkira , Fort (= Fortezza), UTM DJ 08 , 39 .62°N, 19.91°E, SMF 101931 /2 ; Klosterruine bei Gouviá, UTM DJ 08 , 39 .65°N, 19.83°E, SMF 209373 /5 ; Kanóni (= " Al Kanone "), UTM DJ 08 , 39 .59°N, 19.92°E, SMF 284842 /8 ; "Mon Repos", UTM DJ 08 , 39 .60°N, 19.92°E, SMF 101935 /1 ; between Dassia and Káto Korakiána , among roots of olive trees, 40 m alt., UTM DJ 09 , 39 .6830°N, 19.8314°E, leg. E. & P. Subai, 18.07.1975, S 3821/23 ; Pantokrátor [Mountain], UTM DJ 09 , DK 00, 39.74°N, 19.87°E, leg. Brömme, SMF 284861 /1 ; Pantokrátor mountain slope between Barbáti and Nissáki, among rock debris, 200 m alt., UTM DJ 09 , 39 .7249°N, 19.8776°E, leg. E. & P. Subai 20.07.1975, S 3819/4 ; Pantokrátor S slope above Nissáki , approx. 100 m W of Katsoúri cave, 200–250 m alt., UTM DJ 09 , 39 .7299°N, 19.8975°E, leg. Subai 06.08.1995, S 14276 /7 ; S border of road 1.5 km from Petália direction peak of Pantokrátor , limestone rocks, 720 m alt., UTM DJ 09 , 39 .7521°N, 19.8452°E, leg. Subai 05.08.1995, S 14250 /3 (juv.) ; mountain SE of Loútses , on broken karst rocks, 550 m alt., UTM DK 00, 39.7722°N, 19.8928°E, leg. Subai 02.08.1995, S 15842 /3 ; Pantokrátor Mountain slope between Petália and Láfki, on limestone rocks, 480 m alt., UTM DK 00, 39.7601°N, 19.8325°E, leg. E. & P. Subai 07.08.1975, S 3820/9 ; E slope of Pantokrátor karst plateau, 700 m alt., UTM DK 00, 39.74°N, 19.88°E, leg. Gittenberger 10.1978, NNM/1 (alk.). — Paxí Island (= Paxos), UTM n. det., SMF 284832 /2 ; Gaïou (= Gaion), UTM DJ 23 , 39 .19°N 20.18°E, leg. K. L. Pfeiffer 4.1932, SMF 101911 /11 ; rocks between Gaïou and Gaïou port, UTM DJ 23 , 39 .19°N 20.18°E, leg. Megens & Moorsel 14.5.2000, NNM/3 . — Lefkáda Island (= Lefkas ), between road and Evgiros (= 25 km S Lefkáda), 150–175 m alt., UTM DH 67 , 38 .62°N 20.65°E, leg. Gittenberger 21.10.1981, NNM/4 (alk.) ; Megáli Óros plateau, UTM DH 68 , 38 .72°N 20.62°E, SMF 279322 /4 ; S border of Sivros , on limestone rocks along border of road, 170 m alt., UTM DH 68 , 38 .6668°N 20.6433°E, leg. E. & P. Subai 03.08.1975, S 3802/59 ; valley at Stavrotas Mountain (= W of Sivros), on limestone rocks, 200 m alt., UTM DH 68 , 38 .6704°N 20.6373°E, leg. E. & P. Subai 03.08.1975, S 3804/25 ; 7 km SE of Sivros (= direction Póros ), on rocks beside a small chapel, 320 m alt., UTM DH 77 , 38 .6524°N 20.6850°E, leg. E. & P. Subai 03.08.1975, S 3806/2 ; above Póros , on limestone rocks, 300 m alt., UTM DH 77 , 38 .6419°N 20.7089°E, leg. E. & P. Subai 03.08.1975, S 3807/2 ; leg. Pintér & Subai 30.7.1976, S 3808/7 ; mountain slope 500–1000 m S of Nídri , under stones, 20 m alt., UTM DH 78 , 38 .6915°N 20.6979°E, leg. E. & P. Subai 03.08.1975, S 3805/43 ; 2 km from Kariá direction Lefkáda , on limestone rocks beside the highway, 440 m alt., UTM DH 79 , 38 .7756°N 20.6554°E, leg. E. & P. Subai 05.08.1975, S 3803/11 ; Fríni (= 2.5 km W of Lefkáda Town), UTM DH 79 , 38 .82°N 20.68°E, SMF 279321 /6 ; leg. Gittenberger 22.10.1981, NNM/2 (alk.) ; border of Frini (= direction Tsoukaládes), on limestone rocks beside asphalt road, 140 m alt., UTM approx. DH 69, 38.8236°N 20.6800°E, leg. E. & P. Subai 05.08.1975, S 6677/38 ; Kaligóni , between roots of olive trees, UTM DH 79 , 38 .8148°N 20.7124°E, leg. E. & P. Subai 31.07.1975, S 3811/15 ; Kaligóni , from the spring "Megali Vrysi" and on limestone rocks beside the chapel, UTM DH 79 , 38 .8147°N 20.7119°E, leg. E. & P. Subai 01.08.1975, S 3809/21, 3810/8. — Kefaloniá Island , UTM n. det., SMF 101910 /4, 284857/1 ; leg. Heldreich leg., SMF 284858 /1 ; 6 km WNW of Lixoúri , UTM DH 43 , leg. Visser 4.11.1985, NNM/1 (alk.) ; 1.5 km E of Sámi , UTM DH 73 , leg. Visser 6.11.1985, NNM/5 (alk.). — Ípiros (= Epirus), S border of Igoumenítsa , on limestone rocks and from a spring, 40 m alt., UTM DJ 37 , 39 .4858°N 20.2818°E, leg. E. & P. Subai 21–22.07.1975, S 3828/1 ; above Pléssio , on lime conglomerate rocks, 450 m alt., UTM DJ 38 , 39 .6493°N 20.2811°E, leg. Subai 13.05.1995, S 15832 /2 ; mountain slope approx. 2 km N of Kestríni (= NW of Igoumenítsa ), on limestone rocks, 20 m alt., UTM DJ 38 , 39 .5745°N 20.1943°E, leg. Subai, 19.07.1990, S 15876 /4 ; S border of Kótsika , on limestone rocks, under stones, 300 m alt., UTM DJ 39 , 39 .6592°N 20.2471°E, leg. Subai 13.05.1995, S 15834 /4 ; Párga, Valtos Beach , on rocks near castle, UTM DJ 44 , 39 .2832°N 20.3970°E, leg. Megens & Moorsel 14.5.2000, NNM/1 ; 1.4 km N of Agiá (= N of Párga ), on dry limestone rocks, 300 m alt., UTM DJ 45 , 39 .3202°N 20.3443°E, leg. Subai 15.05.1995, S 15830 /1 ; Párga–Agiá country road, 600 m N of Anthoūssa , UTM DJ 45 , 39 .3044°N 20.3679°E, leg. Hoenselaar 8.6.1996, Maa /2 ; Nekromanteion (= Ephyra ), UTM n. det., leg. Hoenselaar 9.6.1996, Maa /2 ; 1 km N of Kartéri (= highway Margaríti–Igoumenítsa ), on limestone rocks, 170 m alt., UTM DJ 46 , 39 .3953°N 20.3702°E, leg. Subai 20.07.1990, S 15829 /2 ; leg. Subai & Szekeres 14–15.5.1997, S 15850 /3 ; 5 km E of Igoumenítsa, mountain slope between Krióvrisi and Lákka Villages , under stones, 370 m alt., UTM DJ 47 , 39 .4882°N 20.3274°E, leg. E. & P. Subai 22.07.1975, S 3837/7 ; 2 km from Parapótamos direction Ioánina (= highway Igoumenítsa–Ioánina ), on limestone rocks, 200 m alt., UTM DJ 47 , 39 .5297°N 20.3426°E, leg. E. & P. Subai 23.07.1975, S 3836/4 ; cave chapel 5 km from highway Filiátes–Igoumenítsa, direction Mavronéri , on limestone rocks, approx. 150–200 m alt., UTM DJ 48 , 39 .6434°N 20.3125°E, leg. Subai, 19.07.1990, S 15875 /1 ; leg. Subai 13.5.1995, S 15833 /4 ; leg. Subai & Szekeres 15.5.1997, S 15852 /4 ; about 5 km SE of the highway Filiátes–Igoumenítsa (= direction Kipárisso, Paleohóri ), approx. 100 m W of the bifurcation to Astós, beside the Thíamis river , on limestone rocks, 50 m alt., UTM DJ 48 , 39 .5673°N 20.3700°E, leg. Subai & Szekeres 15.05.1997, S 15853 /3 ; 9.5 km from the highway Filiátes–Igoumenítsa direction Keramitsa , 900 m from the bifurcation to Rizó (= 1 km E of Dafni), on limestone rocks in the street bend, 300 m alt., UTM DJ 48 , 39 .6429°N 20.4095°E, leg. Subai & Szekeres 15.05.1997, S 15851 /3 ; bifurcation of highway Filiátes–Keramítsa direction Rizó , from here still 2 km eastwards in a river bed, on limestone rocks, 250 m alt., UTM DJ 48 , 39 .6435°N 20.4164°E, leg. Subai, 19.07.1990, S 15874 /6 ; S of Mórfi, junction to Párga , UTM DJ 54 , 39 .2958°N 20.4779°E, leg. Eröss 10.8.1977, HNHM 9003 /4 ; 500 m from Ag. Kiriáki direction Párga (= bifurcation of highway Préveza–Igoumenítsa direction Párga ), dry hill slope, on rocks, 160 m alt., UTM DJ 54 , 39 .2900°N 20.4415°E, leg. E. & P. Subai 06.08.1975, S 3830/5 ; 13 km from Vrossína direction Igoumenítsa, E slope of limestone mountain, 650 m alt., UTM DJ 58 , 39 .5944°N 20.4918°E, leg. Riedel & Subai 13.04.1988, S 15820 /3 ; N of Glikí, along highway to Koukoulií , on limestone rocks, approx. 160 m alt., UTM DJ 65 , 39 .3322°N 20.6030°E, leg. Subai 15.05.1995, S 15831 /1 ; small chapel at N border of Tsangári , on limestone rocks, 480 m alt., UTM DJ 66 , 39 .4202°N 20.6127°E, leg. Subai 15.05.1995, S 15827 /3 ; river gorge at S border of Dovlá , on limestone rocks, 400 m alt., UTM DJ 68 , 39 .5679°N 20.5481°E, leg. Subai 13.05.1995, S 15835 /1 ; Préveza , UTM DJ 71 , SMF 284846 /1, 284855/3 ; leg. Conéménos, 1883, SMF 284850 /4 ; leg. Conéménos, 1899, SMF 101912 /7 ; Camping Kalamitsi Beach , 3 km N of Préveza, UTM DJ 71 , 38 .98°N 20.70°E, leg. Hoenselaar 4.6.1996, Maa /5 ; wall remains of Nikopolis (= N of Préveza), 20 m alt., UTM DJ 71 , 39 .0235°N 20.7367°E, leg. E. & P. Subai 06.08.1975, S 3831/2 ; along old highway Préveza–Igoumenítsa, mountain slope 500 m from road junction to Zálongo direction Igoumenítsa , on limestone rocks, 90 m alt., UTM DJ 73 , 39 .1248°N 20.6568°E, leg. E. & P. Subai 06.08.1975, S 3829/7 ; on high rocks nearly monuments of Zálongo and on the mountain behind Moni Ágios Dimítrios (= 21.5 km NNW of Préveza), 650–725 m alt., UTM DJ 73 , 39 .14°N 20.68°E, leg. Gittenberger & Uit de Weerd 15.5.2000, NNM/1 ; Skafidáki Village near Préveza and at other localities on Skafidáki Peninsula, UTM DJ 81 , 38 .9877°N 20.7830°E, leg. Pintér, E. & P. Subai 31.07.1976, S 3824/2 ; 2 km along the highway Ioánina–Árta opposite the rock hole near Ágios Geórgios , 130 m alt., UTM DJ 84 , 39 .2723°N 20.8446°E, leg. Fauer & Subai 28.09.1989, S 15867 /56 ; on limestone rocks along border of road in Nikolítsi Village , 210 m alt., UTM DJ 85 , 39 .3037°N 20.7676°E, leg. Subai 15.05.1995, S 15828 /4 ; gorge 5 km N of Térovo (= E of highway Ioánina–Árta ), on limestone rocks, 430 m alt., UTM DJ 86 , 39 .4497°N 20.8790°E, leg. E. & P. Subai 30.07.1975, S 3827/1 ; bifurcation of the highway Ioánina–Árta to Térovo , on limestone rocks, 285 m alt., UTM DJ 86 , 39 .4161°N 20.8459°E, leg. Fauer & Subai 28.09.1989, S 15863 /1 ; ruins of the antic amphitheater near Dodoni (= SSW of lake Ioánina), 640 m alt., UTM DJ 87 , 39 .5461°N 20.7879°E, leg. E. & P. Subai 30.07.1975, S 3832/9 ; leg. Neuteboom 26.7.1979, NNM/16 + 1 (juv.) ; Ioánina, UTM DJ 89 , 39 .66°N 20.84°E, HNHM 40244 /1 ; Kria (= N of Pérama near Ioánina ), lime rocks above the upper (= southeastern) cemetery, 610 m alt., UTM DJ 89 , 39 .7226°N 20.8409°E, leg. Fauer & Subai 27.09.1989, S 15857 /49 ; E border of Amfithea (= Stroúni , E of Pérama), on limestone rocks, 480 m alt., UTM DJ 89 , 39 .6860°N 20.8727°E, leg. E. & P. Subai 24.07.1975, S 6676/27 + 1 (alk.) ; leg. Pintér & Subai 30.7.1976, S 3833/2 ; 2 km E of Amfithea (= Stroúni, E of Pérama ), on limestone rocks, 480 m alt., UTM DJ 89 , 39 .6802°N 20.8871°E, leg. E. & P. Subai 24.07.1975, S 3834/2 ; NE border of Pérama , on limestone rocks behind the houses, 490 m alt., UTM DJ 89 , 39 .6986°N 20.8482°E, leg. Fauer & Subai 28.09.1989, S 15862 /39 ; leg. Subai 12.5.1995, S 15845 /34 ; Pérama , on limestone rocks at exit of the stalactite cave, 500 m alt., UTM DJ 89 , 39 .6949°N 20.8457°E, leg. E. & P. Subai 29.07.1975, S 3823/7, 3835/6 ; leg. Pintér & Subai 30.7.1976, HNHM 41385 /15, 49537/4, S 3825/36 ; leg. Neuteboom 27.7.1979, NNM/4 + 2 (juv.) ; leg. Gittenberger & Uit de Weerd 14.5.2000, NNM/3 ; road between Amótopos and Nisista , UTM DJ 94 , 39 .27°N 20.97°E, leg. Neuteboom 26.7.1979, NNM/6 ; Velás monastery (= approx. 2 km S of Kalpáki), on limestone rocks above the spring and in a small cave, 450 m alt., UTM DK 61 , 39 .8705°N 20.6253°E, leg. Fauer & Subai 27.09.1989, S 15858 /2 ; leg. Subai 16.7.1990, S 15871 /4 ; Kalpáki , UTM DK 61 , 39 .88°N 20.62°E, leg. Neuteboom 27.7.1979, NNM/1 ; 1 km NW of Asfáka , UTM DK 70 , 39 .78°N 20.72°E, leg. Kiss & Pintér 25.7.1986, S 29880 /11 + 5 (juv.) ; Kalpáki–Monodéndri highway, dry lime rocks at road junction to Elafótopos , 860 m alt., UTM DK 71 , 39 .8753°N 20.6951°E, leg. E. & P. Subai 30.07.1975, S 3826/1 ; Arísti , UTM DK 72 , 39 .93°N 20.67°E, leg. Neuteboom 27.7.1979, NNM/1 ; gorge entrance at bridge across Sarantáporos River between Exohí and Amárantos, dry rock slope, 445 m alt., UTM DK 74 , 40 .1114°N 20.7218°E, leg. Subai 16.07.1990, S 15869 /24 ; leg. Subai & Szekeres 16.5.1997, S 15846 /1 ; Loutrá (= 5 km N of Amárantos), W slope of limestone mountain, 1270 m alt., UTM DK 74 , 40 .1745°N 20.7297°E, leg. Riedel & Subai 12.04.1988, S 15821 /1 ; Loutrá (= N of Amárantos), limestone mountain SW side, 1120 m alt., UTM DK 74 , 40 .1676°N 20.7321°E, leg. Fauer & Subai 25.09.1989, S 15860 /2 ; mountain E of Loutrá near Amárantos , on N exposed limestone rocks and in forest approx. 500 m SE of Loutra, 1250 m alt., UTM DK 74 , 40 .1745°N 20.7297°E, leg. Subai & Szekeres 16.05.1997, S 15847 /3 ; Tímfi Mountains , limestone grove after 3h 30' walking E of Pápigo village (= approx. 1–2 km NE of refuge hut), on limestone boulders and under stones, 1650–1700 m alt., UTM DK 82 , 39 .9961°N 20.7709°E, leg. Subai 24.05.1991, S 15819 /5 ; Tímfi Mountains, near Drakólimni Lake , 2215 m alt., UTM DK 82 , 39 .98°N 20.77°E, leg. Gittenberger 26.7.1997, NNM/1 ; 6.6 km from Kónitsa direction Pádes , on limestone rocks in dry mixed forest, 1000 m alt., UTM DK 83 , 40 .0547°N 20.7898°E, leg. Subai 16.07.1990, S 15870 /2 ; Pádes (= approx. 28 km E of Kónitsa), on limestone rocks at W and N border of the village, 1150–1250 m alt., UTM DK 93 , 40 .0408°N 20.9054°E, leg. Subai 16.07.1990, S 15868 /2 ; rocks at S border of Agía Paraskeví Village , 995 m alt., UTM DK 94 , 40 .1438°N 20.8965°E, leg. Fauer & Subai 25.09.1989, S 15859 /1 ; approx. 1 km E of Agía Paraskeví (= in N part of Smólikas Mountains ), on rocks beside the highway, 1060 m alt., UTM DK 94 , 40 .1411°N 20.9069°E, leg. Fauer & Subai 25.09.1989, S 15861 /2 ; 3 km N of Drossopigí, meadow and rocks (no lime) approx. 50 m E of the old arched bridge above the Sarantáporos river , 705 m alt., UTM DK 95 , 40 .2281°N 20.9133°E, leg. Subai 15.07.1990, S 15873 /1 + 1 (juv.) ; 17.5 km NE von Árta, an Felsen neben der Landstrasse, 180 m alt., UTM EJ 14 , leg. Gittenberger & Uit de Weerd 14.5.2000, NNM/2 . — Evritanía, in spruce forest 6.8 km S of Proussós (= direction Agrínio ), under stone plates, 1300 m alt., UTM EH 58 , 38 .7164°N 21.6533°E, leg. Subai 17.05.1991, S 15818 /13 ; bridge at Mégdova river (= Tavropós , 26 km W of Karpeníssi), on limestone rocks, 340 m alt., UTM EJ 51 , 38 .9415°N 21.6809°E, leg. Fauer & Subai 30.09.1989, S 15854 /3 ; small valley at mountain slope E of Kalesméno (= 16 km W of Karpeníssi), on limestone rocks, 880 m alt., UTM EJ 60 , 38 .9212°N 21.7281°E, leg. Fauer & Subai 30.09.1989, S 15856 /1 ; 1.5 km N of Vráha, approx. 500 m from Moni Vráha , mountain N and S slope, on limestone rocks, approx. 1100 m alt., UTM EJ 62 , 39 .0776°N 21.7930°E, leg. Subai & Szekeres 12.05.1997, S 15815 /2 + 1(alk.) ; Kaliakoúda Mountains, Timfristós Mountain (= " Kaljakuda Berge, Veluchi "), UTM n. det., leg. Krüper, 1890, SMF 101913 /4, 101914/17, 284854/3. — Etoloakarnanía, UTM n. det., SMF 101936/3 ; Etoloakarnanía, on the narrow land strip leading to Lefkáda Island, on limestone rocks and on ruins of a small castle, UTM DH 79 , 38 .8434°N 20.7213°E, leg. E. & P. Subai 06.08.1975, S 3816/4 ; Vónitsa, UTM DJ 90 , 38 .92°N 20.88°E, leg. Conéménos, 1890, SMF 284856 /2 ; Veloútsa Mountain SE slope above Astakós , on limestone rocks, approx., 185 m alt., UTM EH 06 , 38 .5370°N 21.0692°E, leg. Subai, 19.05.1991, S 15817 /19 ; small valley 1 km E of Astakós , on limestone rocks, UTM EH 06 , 38 .5323°N 21.0998°E, leg. Subai, 19.05.1991, S 15816 /10 ; Lisimáhia (= Murstiana, Mushtiona), UTM EH 26 , 38 .54°N 21.33°E, leg. Krüper, 1928, SMF 101916 /32 ; Messolóngi, UTM EH 34 , 38 .36°N 21.42°E, SMF 285671 /2 ; leg. Krüper, 1899, SMF 101915 /7, above Kefalóvrisso , on limestone rocks, 30 m alt., UTM EH 35 , 38 .4598°N 21.3670°E, leg. Fauer & Subai 30.09.1989, S 15855 /6 ; Klisura gorge between Agrínio and Etolikó, on limestone rocks above Ag. Eleusis monastery, 290 m alt., UTM EH 36 , 38 .5006°N 21.3674°E, leg. Pintér, E. & P. Subai 31.07.1976, S 3815/3 ; Berg Varássova (= Karvassara), UTM EH 54 , 38 .36°N 21.59°E, leg. Conéménos, 1884, SMF 284849 /2 ; Gurica (= Goritza, between Mirtéa and Ágios Nikólaos monastery) at Trihonída Lake (= Lake of Vrachori ), UTM EH 57 , 38 .58°N 21.61°E, leg. Krüper, 1891, SMF 101938 /5, 101940/17, 207802/1, 284831/4, 284843/6 ; N of Amfilohía , on limestone rocks 100–200 m from the bifurcation direction Sardínia, 60 m alt., UTM EJ 10 , 38 .8833°N 21.1740°E, leg. Fauer & Subai 29.09.1989, S 15865 /16 ; 5.9 km from Empessós direction Perdikáki , on limestone rocks in valley of two lime mountains, approx. 600 m alt., UTM EJ 31 , 39 .0058°N 21.3606°E, leg. Fauer & Subai 29.09.1989, S 15866 /2 ; 2.5 km E of Halkiópouli , on limestone rocks, 600 m alt., UTM EJ 31 , 38 .9829°N 21.3700°E, leg. Subai 17.05.1995, S 15839 /2 ; 9.5 km from Halkiópouli direction Tríklino , on limestone rocks, 950 m alt., UTM EJ 31 , 38 .9666°N 21.3964°E, leg. Fauer & Subai 29.09.1989, S 15864 /1 ; 11.5 km E of Halkiópouli , on small limestone rocks, 750 m alt., UTM EJ 31 , 38 .9702°N 21.4000°E, leg. Subai 17.05.1995, S 15838 /2. Fokída, Strómi Village (= 13.5 km SW of Paúliani), on limestone rocks, 850 m alt., UTM FH 08 , 38 .7089°N 22.2568°E, leg. Subai 12.04.1986, S 15822 /1. — Ahaïa , Pátra (= Patras), UTM EH 63 , 38 .23°N 21.73°E, SMF 101942 /1.

Remarks: According to Cernohorsky (1978: 299), the publication of Anton was issued "around the middle of the year, 1838", but therein, the name Helix corcyrensis remained a nomen nudum. Thus, Helix canalifera ANTON, 1838 is the first correct and valid description of the Lindholmiola species from Corfu. This rarely used name (last mention by Polinski (1924: 130)) is here considered a nomen oblitum. However, Rossmässler (1838: 40, Taf. 39, Fig. 538) validated the name H. corcyrensis (ICZN 23.9 cannot be applied), and it was used under his authority in the subsequent literature. The type specimen of Rossmässler is missing in his collection (note of the late Dr Zilch).

In its shell form, L. corcyrensis is quite stable, but it varies considerably in size. The populations on the Ionian Islands and the western Epirus are small to medium-sized with a narrower umbilicus and a flat spire (cefalonica sensu auct.). In the central parts of Epirus, shells with a broader umbilicus occur. Similarly sized shells, but with an even larger umbilicus and flatter spire are recorded from the Tímfi and Grammos Mountains , where they live at altitudes between 1200–2000 m.

The eastern part of the distribution area of L. corcyrensis is inhabited by populations with a strikingly different shell. Here, shells grow larger, with a more elevated spire and a large umbilicus (octogyrata sensu auct.). However, in the same area, shells have been recorded, which do not differ from those in the Epirus, and thus, octogyrata cannot be considered a separate taxon.

Very small specimens can be found on the island of Lefkáda in Greece and in some places on the eastern shore of Lake Ohrid.

Distribution ( Fig. 29 ): Lindholmiola corcyrensis inhabits the south-western Balkan Peninsula. There are two isolated localities known, the Pelješac Peninsula and the Island Badija in Croatia, and the Monte Gargano in Italy. South of the Croatian locality, there is an area of almost 200 km length without any Lindholmia species at all. The main distribution area of L. corcyrensis starts in south-eastern Montenegro and covers the whole of Albania and western Greece. In the south, it reaches the island of Kefallinia and the northern shores of the Bay of Korinthos. The southernmost record from close to Pátra in Ahaïa could not be verified by recent collecting efforts. The eastern boundary of its distribution area follows the eastern rim of the Pindos Mountains.

Figures

Figs 20–23. Lindholmiola corcyrensis. Fig. 20: NMBE 515620, Island Kérkira between Dassía and Káto Korakiána (form with narrow umbilicus, Ionian Islands), D = 10.8 mm; Fig. 21: NMBE 515622, Island Lefkáda, above Póros (small form), D = 9.19 mm; Fig. 22: NMBE 515621, Ipiros, Tímfi Mts., hiking path of Pápigo, 1–2 Km N of the shelter (large umbilicus, flat shell), D = 11.23 mm; Fig. 23: syntype Helix (Trigonostoma) corcyrensis var. cefalonica, ZMZ 508737, Island Kefaloniá, "aux environs d'Argostoli, ...à Frangata et Grisata", D = 12.0 mm. — All photos Neubert/Bochud, × 3.

Figs 24–25. Lindholmiola corcyrensis. Fig. 24: syntype Helix corcyrensis var. octogyrata, ZMZ 508745, Ípiros, surroundings of Préveza, D = 14.36 mm; Fig. 25: NMBE 515623, Evritanía, mountain ca. 5 Km NE of Vráha (large form), D = 13.73 mm. — All photos Neubert/Bochud, × 3.

Figs 26–28. Genital organs of Lindholmiola species. Fig. 26: Lindholmiola pirinensis, a: situs, b: lumina of genital organs, Greece, Makedonía, Falakró Mts., 3 Km from the crossing Vólakas in direction to Skiing centre; Fig. 27: Lindholmiola corcyrensis, a: situs, Greece, Island Kefaloniá, 1.5 Km E of Sámi, b: lumina of genital organs, Greece, Island Lefkáda, 25 Km S von Lefkáda village; Fig. 28: Lindholmiola reischuetzi, a: situs, b: lumina of genital organs, Greece, Makedonía, 4 Km of Kehrókambos in direction to Stavroúpoli.

Fig. 29. Distribution of Lindholmiola corcyrensis and L. girva.

Figs. 1–2. Genital organs of Helicodontidae. Fig. 1: Lindholmiola barbata; Greece, Kríti, Nomos Haniá, Bay of Soúda, 100–150 m from the mouth of the river S of the English cemetery. Fig. 2: Helicodonta obvoluta (O. F. MüLLER, 1774); Belgium, valley of the Meuse, 500 m from Annevoie in direction to Dinant, 2a) situs of genital organs, 2b) interior structure of penis. — Abbreviations used for the description of the genital organs: A = atrium, Al = albumen gland, E = Epiphallus, G = distal glandular part of epiphallus, Gl = Glandula, Hd = hermaphroditic duct, Hg = hermaphroditic gland, P = Penis, R = retractor muscle, Sp = Spermoviduct, V = vesicle, Vag = Vagina, Vd = Vas deferens.

Figs 3–6. Lindholmiola lens. Fig. 3: syntype Helix lentiformis, ZMZ 508656, "Thessalie et Attique", D = 11.2 mm; Fig. 4: NMBE 515542, Attika, Methana, 500 m before village, 12.07.1996, leg. E. Neubert, D = 12.77 mm; Fig. 5: syntype Helix lens var. callojuncta, NHMG 372, D = 12.0 mm; Fig. 6: syntype Helix lens var. elia, SMF 101961, Kumani, Elis, coll. Jetschin ex Boettger, 1883, D = 11.94 mm. — All photos Neubert/Bochud, × 3.

Figs 7–11. Lindholmiola lens. Fig. 7: NMBE 515616, Greece, Ilía (= Elis), Alfiós canal between Olimbía and Kréstena (nominotypical form), D = 11.78 mm; Fig. 8: NMBE 515617, Greece, Fokída, Delphi, archaeological site (flat keeled form,broad whorls, wide umbilicus), D = 14.85 mm; Fig. 9: NMBE 515614, Greece, Thessalía, W boundary of Morfovoúni (densely coiled, blunt keel, narrow umbilicus), D = 12.88 mm; Fig. 10: NMBE 515615, Greece, Makedonía, Kassándra Peninsula, Loutrá (small form, blunt keel, moderately wide umbilicus), D = 7.95 mm; Fig. 11: NMBE 515629, Greece, Makedonía, Canyon 2 Km N of Neohóri (= NE of Aridéa) (flat form, blunt keel, wide umbilicus), D= 14.16 mm. — All photos Neubert/Bochud, × 3.