Revision of the genus Lindholmiola HESSE, 1931 (Gastropoda: Pulmonata: Helicodontidae)

Abstract

A revision of the land snail genus Lindholmiola that is mainly found on the Balkan Peninsula is presented. Until now, 9 valid species are considered to form the genus: Lindholmiola lens (A. Férussac, 1832), L. barbata (A. Férussac, 1821), L. corcyrensis (Rossmässler, 1838), L. girva (Frivaldszky, 1835), L. gyria (Roth, 1839), L. pirinensis Jaeckel, 1954, L. regisborisi (A. J. Wagner, 1927), L. reischuetzi Falkner, 1995 and L. spectabilis Urbanski, 1960. All species are characterized by their shells and the anatomy of their genital organs; their distribution patterns are shown in maps. All available type specimens and characteristic shell-forms are illustrated.

Full text

CONTRIBUTIONS TO NATURAL HISTORY No. 23: 1–94 · 3 April 2014
Revision of the genus Lindholmiola Hesse, 1931
(Gastropoda: Pulmonata: Helicodontidae)
Peter Subai & Eike Neubert
ABSTRACT Contrib. Nat. Hist. 23: 1–94
A revision of the land snail genus Lindholmiola that is mainly found on the Balkan
Peninsula is presented. Until now, 9 valid species are considered to form the genus:
Lindholmiola lens (A. Férussac, 1832), L. barbata (A. Férussac, 1821), L. corcyrensis
(Rossmässler, 1838), L. girva (Frivaldszky, 1835), L. gyria (Roth, 1839), L. pirinen-
sis Jaeckel, 1954, L. regisborisi (A. J. Wagner, 1927), L. reischuetzi Falkner, 1995
and L. spectabilis Urbanski, 1960. All species are characterized by their shells and
the anatomy of their genital organs; their distribution patterns are shown in maps.
All available type specimens and characteristic shell-forms are illustrated.
Key words: Taxonomy, Lindholmiola, anatomy, distribution.
Kurzfassung: Eine Revision der hauptsächlich auf der Balkan Halbinsel beheimate-
ten Landschneckengattung Lindholmiola wird präsentiert. Hierbei werden folgende
9 Arten als valide anerkannt: Lindholmiola lens (A. Férussac, 1832), L. barbata (A.
Férussac, 1821), L. corcyrensis (Rossmässler, 1838), L. girva (Frivaldszky, 1835),
L. gyria (Roth, 1839), L. pirinensis Jaeckel, 1954, L. regisborisi (A. J. Wagner, 1927),
L. reischuetzi Falkner, 1995 sowie L. spectabilis Urbanski, 1960. Die Gehäusemerk-
male sowie die Merkmale der Genitalorgane aller Arten werden dargestellt, ihre Ver-
breitung wird anhand von Karten illustriert. Alle verfügbaren Typusexemplare sowie
charakteristische Formen werden abgebildet.
Schlüsselwörter: Taxonomie, Lindholmiola, Anatomie, Verbreitung.

2Peter Subai & Eike Neub ert
Introduction
The revision of the genus Lindholmiola was initiated by several Dutch col-
leagues, who contributed by basic bibliographic work (L. J. M. Butot and E.
Gittenberger), and the extensive collection of both, shells and preserved
specimens by W. Neuteboom and W. J. M. Maassen. A preliminary step was
made by Gittenberger & Groh (1986), who clarified the status of the two old-
est available names in the genus by designating lectotypes for Helix barbata
A. Férussac, 1821 and Helix lens A. Férussac, 1832. The unfinished version
of this manuscript was then handed on by Gittenberger to the present authors
including his photographic documentation of specimens.
The list of synonyms contains only those references where the recorded
species can be identified. For all other records, the specimens have been
checked by the authors personally. The localities follow an order from North
to South, and from West to East; if possible, all localities are written using
their respective national notation. A few localities could not be traced; their
spelling is copied from the original labels or reference.
In some species of Lindholmiola, the aperture is bent downwards, which
makes it difficult to measure its absolute height. For this reason, all measure-
ments of the apertural height presented here use the following method: The
shell is fixed to the ground exposing the ventral side to enable the exact
measurement of the shell height; the value of the apertural height in fron-
tal view is added in parentheses. The number of whorls is counted following
the method of Ehrmann (1933: 21). Genital organs are described using the
gonads as proximal and the atrium as distal end. All drawings are to the same
scale, all measurements in mm.
Material
Collections used:
E&F Private collection Z. Erőss and Z. Fehér, Budapest
HNHM Hungarian Natural History Museum, Budapest
Maa Private collection J. M. W. Maassen, Duivendrecht
NHMG Naturhistoriska Museet, Göteborg
NMNH National Museum of Natural History, Sofia
NNM Nationaal Natuurhistorisch Museum, Leiden
S Private collection P. Subai, Aachen (now at the NMBE)
SMF Senckenberg Museum, Frankfurt a. M.
ZMB Zoologisches Museum, Berlin

3
Revision of the genus Lindholmiol a Hesse, 1931
ZMZ Zoologisches Museum the Universität Zürich-Irchel
ZSM Zoologische Staatssammlung, München
Other abbreviations:
aH apertural height
aW apertural width
D shell diameter
H shell height
n. id. UTM-Code not identified
UTM UTM-Code for registration of the European invertebrates
Systematic account
Lindholmiola Hesse, 1931
1931 Lindholmiola Hesse, Zoologica, 18: 50 [type species by original designation
Helix lens A. Férussac, 1832].
Description: shell flat to lentiform (H: 3.1–7.5 mm, D: 7.2–16.5 mm); spire
slightly elevated (exception: L. gyria with a concave spire); colour brown to
yellowish-brown; teleoconch surface usually with hairs; teleoconch sculpture
of fine to coarse radial riblets; whorls regularly increasing, 5¼–7¾ whorls;
periphery of last whorl rounded, bluntly shouldered or sharply keeled; last
whorl strongly bends downwards before reaching the aperture; umbilicus nar-
row to moderately wide, cylindrical to perspective, usually only slightly cov-
ered by the peristome (exception: L. barbata, where the umbilicus is almost
completely covered); aperture narrow, oblique and curved (in lateral view);
peristome narrow to broad, reflexed.
Genital organs (Fig. 1) simple; atrium short; penis a club-shaped tube,
epiphallus distally strongly contracted (often, this area is covered by a glan-
dular tissue); penial retractor muscle attaching at the central part of epiphal-
lus, which connects to vas deferens without any visible distinction; vagina
short (completely reduced in L. reischuetzi); glandula a single tube with a
narrow stem and a broader hose often sculptured by crossing furrows; bursa
copulatrix with a narrow pedunculus and a more broadened vesicle (often not
clearly visible) (Schileyko 2006: 1919, Fig. 2439).
Interior of the terminal genital organs with a long fold starting at the genital
pore and ending at the distal vagina; atrial part of the fold often subdivided
by small crossing furrows (an additional semicircular crested fold in the atrium

4Peter Subai & Eike Neub ert
Figs. 1–2. Genital org ans 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: Helicodon-
ta obvoluta ( O. F. Mül ler, 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 glan-
dular 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.

5
Revision of the genus Lindholmiol a Hesse, 1931
can be found in L. regisborisi); characters of the penial lumen differ according
to the species, usually with 2–3 raised longitudinal pilasters filling at least the
distal half of the penial lumen (exception: In L. corcyrensis, the longitudinal
pilasters are replaced by v-shaped, obliquely arranged, short folds).
It has to be stressed that the characters of the terminal genital organs as
described above can only be seen in sexually mature specimens. In juveniles,
the folds are missing or very weak.
Differentiating remarks: The shells of Helicodonta are also depressed
but never lentiform and usually concave. In species of this genus, the whorls
increase faster in width, particularly the ultimate and the penultimate whorl,
which are always broader than those in Lindholmiola. Additionally, they are
laterally rounded and only slightly shouldered. The umbilicus of Helicodonta
spp. is broader and more perspective if compared to Lindholmiola, and their
apertures have often three indentations which are even thickened to form
teeth. The peristome is usually broadened to form a lip.
Both genera differ in the morphology of their genital organs: Helicodonta spe-
cies have a considerably longer penis and vagina and a short tubular organ
of unknown function that branches off distally to glandua and bursa copula-
trix (Fig. 2) (not present in Lindholmiola). In the interior of the terminal genital
organs, there are elongated narrow pilasters in the vagina (different in Lind-
holmiola), and in the middle of the penis, the penial lumen is filled by two
thickened pilasters (different in Lindholmiola). In the distal part of the penis,
an additional pilaster covered by small punctiform granules is found (not present
in Lindholmiola). The glandular tissue at the distal end of the epiphallus of
Lindholmiola spp. is missing in Helicodonta species.
Lindholmiola lens (A. Férussac, 1832) Figs 3–6, 7–11, 12, 15
1821 Helix barbata A. Férussac, Tableaux systématiques des animaux mol-
lusques classés en familles naturelles: 41 (Folio edition) or 37 (Quarto edi-
tion), Nr. 152 [partim] .
1832 Helix lens A. Férussac (in A. Férussac & Deshayes), Histoire naturelle
générale et particulière des mollusques terrestres et fluviatiles, (Atlas),
22–27: Taf. 66, F. 2 [Lectotype, design. Gittenberger & Groh 1986] .
1832 Helix barbata, – Deshayes, Exped. Moree Moll., p. 162 (non barbata
Férussac).
1838 Helix lens, – Rossmässler, Iconographie der Land- & Süsswasser-Mollusken
mit vorzüglicher Berücksichtigung der europäischen noch nicht abgebildeten
Arten, (1) 2 (1/2): 10, Taf. 32, Fig. 450.

6Peter Subai & Eike Neub ert
1846 Helix lens, – L. Pfeiffer, In: Martini & Chemnitz: Systematisches Conchylien-
Cabinet, (1) 12 (II): 101, Taf. 13, Fig. 16–17.
1848 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 1: 209.
1850 Helix lens, – Deshayes (in Férussac & Deshayes), Histoire naturelle géné-
rale et particulière des mollusques terrestres et fluviatiles, (Text), 1: 110
[partim, nec descr. et terr. typ.] .
1853 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 3: 162.
1854 Helix lens, – Reeve, Conchiologica Iconica, 7: species 1221.
1855 Helix lens, – Roth, Malakozoologische Blätter, 2: 30.
1859 Helix lens, – Mousson, Vierteljahrsschrift der Naturforschenden Gesell-
schaft in Zürich, 4: 258.
1859 Helix lentiformis Mousson, Vierteljahrsschrift der Naturforschenden
Gesellschaft in Zürich, 4: 258. Locus typicus: "Thessalie et Attique".
1859 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 4: 187.
1868 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 5: 260.
1868 Helix lentiformis, – L. Pfeiffer, Monographia heliceorum viventium, 5: 260.
1873 Helix barbata, – Martens, Malakozoologische Blätter, 20: 32.
1873 Helix lens, – Martens, Malakozoologische Blätter, 20: 32.
1874 Helix lens, – Martens, Malakozoologische Blätter, 21: 122.
1876 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 7: 295.
1876 Helix lens, – Westerlund, Fauna der in der paläarctischen Region lebenden
Binnenconchylien, Prodromus, I: 40.
1876 Helix lens var. aliostoma Westerlund, Fauna der in der paläarctischen
Region lebenden Binnenconchylien, Prodromus, I: 41.
1876 Helix lens var. lentiformis, – Westerlund, Fauna der in der paläarctischen
Region lebenden Binnenconchylien, Prodromus, I: 41.
1877 Helix lens, – L. Pfeiffer, Monographia heliceorum viventium, 8: 576.
1879 Helix (Trigonostoma) lens, – Westerlund & Blanc, Aperçu sur la faune
malacologique de la Grèce inclus l’Epire et la Thessalie: 34.
1879 Helix (Trigonostoma) lens var. lentiformis, – Westerlund & Blanc, Aperçu
sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 35.
1879 Helix (Trigonostoma) lens var. piligera Westerlund & Blanc, Aperçu sur
la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 36. Taf. 1,
Fig. 10.
1879 Helix (Trigonostoma) lens var. callojuncta Westerlund & Blanc, Aperçu
sur la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 36.
Taf. 2, Fig. 11.
1882 Helix lens, – Hesse, Jahrbücher der Deutschen Malakozoologischen Gesell-
schaft, 9: 320.

7
Revision of the genus Lindholmiol a Hesse, 1931
1882 Helix lens var. lentiformis, – Hesse, Jahrbücher der Deutschen Malako-
zoologischen Gesellschaft, 9: 320.
1883 Helix (Anchistoma) lens, – O. Boettger, Jahrbücher der Deutschen Malako-
zoologischen Gesellschaft, 10: 323.
1883 Helix (Anchistoma) lens var. elia O. Boettger, Jahrbücher der Deutschen
Malakozoologischen Gesellschaft, 10: 330.
1884 Helix lens, – Hesse, Jahrbücher der Deutschen Malakozoologischen
Gesellschaft, 11: 233, Taf. 4, Fig. 5, Fig. 5 a.
1885 Helix (Gonostoma) lens var. piligera, – O. Boettger, in Stussiner & O. Boett-
ger, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 12: 165.
1885 Helix (Gonostoma) lens var. lentiformis, – O. Boettger, in Stussiner & O. Boett-
ger, Jahrbücher der Deutschen Malakozoologischen Gesellschaft, 12: 165.
1887 Helix (Caracolina) lens, – Tryon, Manual of Conchology, 3: 119.
1887 Helix (Caracolina) lens var. lentiformis, – Tryon, Manual of Conchology, 3: 119.
1887 Helix (Caracolina) lens var. piligera, – Tryon, Manual of Conchology, 3: 119.
1887 Helix (Caracolina) lens var. callojuncta, – Tryon, Manual of Conchology, 3: 119.
1887 Helix (Caracolina) lens var. aliostoma, – Tryon, Manual of Conchology, 3: 119.
1889 Helix (Caracollina) lens, – Martens, Archiv für Naturgeschichte, 55 (1): 172.
1889 Helix (Gonostoma) lens var. lentiformis, – Martens, Archiv für Naturge-
schichte, 55 (1): 173.
1889 Helix (Caracolina) lens, – Westerlund, Fauna der in der paläarctischen
Region lebenden Binnenconchylien, 2: 22.
1889 Helix (Caracolina) lens var. lentiformis, – Westerlund, Fauna der in der
paläarctischen Region lebenden Binnenconchylien, 2: 22.
1889 Helix (Caracolina) lens var. piligera, – Westerlund, Fauna der in der palä-
arctischen Region lebenden Binnenconchylien, 2: 23.
1889 Helix (Caracolina) lens var. callojuncta, – Westerlund, Fauna der in der
paläarctischen Region lebenden Binnenconchylien, 2: 23.
1889 Helix (Caracolina) lens var. elia, – Westerlund, Fauna der in der paläarcti-
schen Region lebenden Binnenconchylien, 2: 23.
1891 Helix (Gonostoma) lens var. callojuncta, – O. Boettger, Nachrichtsblatt
der Deutschen Malakozoologischen Gesellschaft, 23 (5/6): 83.
1892 Helix lens, – Schuberth, Archiv für Naturgeschichte, 58 (1): 8, Taf. 1 Fig. 6.
1892 Helix lens var. lentiformis, – Schuberth, Archiv für Naturgeschichte, 58 (1): 8,
Taf. 1, Fig. 7–8.
1894 Helix (Caracollina) lens, – Pilsbry, Manual of Conchology, 9: 288.
1894 Helix (Caracollina) lens var. lentiformis, – Pilsbry, Manual of Conchology,
9: 288.
1894 Helix (Caracollina) lens var. piligera, – Pilsbry, Manual of Conchology, 9: 288.

8Peter Subai & Eike Neub ert
1894 Helix (Caracollina) lens var. callojuncta, – Pilsbry, Manual of Conchology,
9: 288.
1894 Helix (Caracollina) lens var. aliostoma, – Pilsbry, Manual of Conchology,
9: 288.
1894 Helix (Caracollina) lens var. elia, – Pilsbry, Manual of Conchology, 9: 288.
1894 Helix (Anchistoma) lens, – O. Boettger, Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 26 (1–2): 2.
1902 Helix (Gonostoma) lens, – Sturany, Denkschriften der Kaiserlichen Aka-
demie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche
Klasse Wien, 52: 403.
1902 Helix (Gonostoma) barbata, – Sturany, Denkschriften der Kaiserlichen
Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaft-
liche Klasse Wien, 52: 403 (non barbata Férussac).
1918 Caracollina lens, – Hesse, Nachrichtsblatt der Deutschen Malakozoologi-
schen Gesellschaft, 50: 109.
1918 Caracollina lens lentiformis, – Hesse, Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 50 : 109.
1918 Caracollina lens aliostoma, – Hesse, Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 50 : 109.
1918 Caracollina lens piligera, – Hesse, Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 50 : 109.
1918 Caracollina lens callojunc ta, – Hesse, Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 50: 109.
1918 Caracollina lens elia, – Hesse, Nachrichtsblatt der Deutschen Malakozoo-
logischen Gesellschaft, 50: 109.
1928 Helicodonta lens, – Adensamer & Käufel, Sitzungsberichte der Akademie
der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse,
137 (10): 793.
1928 Helicodonta lens piligera, – Adensamer & Käufel, Sitzungsberichte der
Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaft-
liche Klasse, 137 (10): 793.
1928 Helicodonta lens lentiformis, – Adensamer & Käufel, Sitzungsberichte der
Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaft-
liche Klasse, 137 (10): 793.
1929 Helicodonta (Caracollina) lens, – Gambetta, Archivio Zoologico Italiano,
13 (1/2): 51.
1930 Caracollina lens, – Käufel, Sitzungsberichte der Akademie der Wissen-
schaften in Wien, Mathematisch-Naturwissenschaftliche Klasse, 139: 184.
1931 Lindholmiola lens, – Hesse, Zoologica, 31 (81) 1/2: 50.

9
Revision of the genus Lindholmiol a Hesse, 1931
1936 Lindholmiola lens, – Fuchs & Käufel, Archiv für Naturgeschichte, N.F. 5 (4):
651. Fig. 77 [partim, not in the Angista canyon close to Drama]
1941 Lindholmiola (Helicodonta) lens lens, – Käufel & Fuchs, Verhandlungen
der Zoologisch-Botanischen Gesellschaft in Wien, 88/89: 201.
1960 Lindholmiola lens, – Zilch, Euthyneura, p. 693, Abb. 2419.
1961 Lindholmiola lens, – Jaeckel & Plate, Zoologische Abhandlungen, Abhand-
lungen and Berichte aus dem Staatlichen Museum für Tierkunde in Dresden,
26 (1): 3.
1962 Lindholmiola lens, – Klemm, Sitzungsberichte der Österreichischen Aka-
demie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse,
Abt. I, 171 (6/7): 252.
1976 Lindholmiola lens, – Frank, Mitteilungen der Zoologischen Gesellschaft
Braunau, 2 (9/11): 261.
1986 Lindholmiola lens, – Gittenberger & Groh, Archiv für Molluskenkunde, 116
(4/6): 220. Abb. 1, a (= 2).
1986 Lindholmiola lens, – Reischütz, Malakologische Abhandlungen aus dem
Staatlichen Museum für Tierkunde Dresden, 11 (9): 94.
1990 Lindholmiola lens, – Fechter & Falkner, Steinbachs Naturführer, p. 222
Abb. 1–3, 5.
1996 Lindholmiola lens, – Welter-Schultes, Schriften zur Malakozoologie, 9:
22, 31.
1996 Lindholmiola lens, – Dhora & Welter-Schultes, Schriften zur Malakozoo-
logie, 9: 156, 190, Taf. 13, Fig. 167.
2002 Lindholmiola lens, – Dhora, Studime mbi molusqet e Shqipërisë: 99: 193.
Diagnosis: shell flat, lower whorls with a blunt to sharp keel, umbilicus
cylindrical to broadly perspective, penial lumen with two longitudinal pilasters.
Description of shell: shell almost flat, shell colour light brown to yellow-
ish brown; protoconch whorls smooth; initial whorls of the teleoconch with
fine radial stripes, developing to small riblets on the subsequent whorls, but
decreasing in height on the last teleoconch whorls; on the subsurface, radi-
al riblets weak to almost missing; sculpture of teleoconch of fine granules;
below the suture a small stripe devoid of hairs, the remaining shell including
the umbilicus covered by usually impermanent hairs (often only the periphery
of the shell with hairs); hairs yellowish-brown, 0.56–0.75 mm long; 5½–7¾
regularly increasing whorls; lower whorls usually flattened with a blunt to
sharp keel (sometimes the whorls are more rounded forming an inconspicu-
ous keel); last whorl only slightly descending, then shortly bent towards the
aperture; suture moderately deep; umbilicus shape variable, either initially
cylindrical and rapidly increasing with the last whorl, or initially broadly per-

10 Peter Subai & Eike Neub ert
spective and regularly increasing, reaching a diameter of 1.6–3.9 mm; aper-
ture oblique, in lateral view slightly to strongly bent; peristomial insertions
2.2–5.9 mm apart, connected by an inconspicuous callus (except in callojuncta);
peristome sharp to blunt, sometimes slightly reflected; columellar peristomial
callus weak, covering the umbilicus ca 10%.
Measurements: H: 3.6–7; D: 7.2–15.6; aH: 2–3.9 (3–6.1); aW: 3.4–7.8.
Details of body (n = 12 from 5 populations: 1 specimen from 2 km N of
Neohóri/1; 4 specimens from the pass at Haniá; 1 specimen from E of Glifá-
da; 2 specimens from 3 km S of Ptéri; and 4 specimens between Olymbía and
Kréstena): head and dorsum grey to brown, more bright at the flanks and the
posterior body; foot sole cream, sometimes darker at the fringes; mantle on
the last 1½ whorls with small irregular dots; dots are more elongate between
lung vein and suture and larger than those on the lower side of the mantle;
intensity of pigmentation increasing towards the mantle collar, pigments often
fuse to a large brown spot on the last whorl; ureter opens 1.5–1 mm before
the respiratory pore.
Morphology of the genital organs: penis relatively short, moderately thick-
ened, glandular tissue covering one third to half of the total of the penial tube;
penial lumen filled by two longitudinal pilasters of equal length starting as
thin threads on the distal end of the penis and ending in the proximal third of
the penis with thickened tissue; sometimes a few small folds at the distal end
of the penis parallel to the major pilasters; vagina moderately long, usually
thickened; glandula with a short stem and a long distal part with deep cross-
ing furrows; bursa copulatrix as long as or somewhat longer than the glandula,
the pedunculus reaches ca 60% of the total length of the organ, usually well
separated from the vesicle; atrium with a broad and flat pilaster starting at
the genital pore, often subdivided by deep longitudinal furrows; vagina filled
by a single sometimes broadened pilaster.
Differential diagnosis: Its sisterspecies seems to be L. spectabilis, which
has a shell that is similar to the form callojuncta of L. lens. It differs from this
form by its more slowly increasing and less perspective umbilicus. Generally,
the shell surface in L. spectabilis is more coarsely granulated, hairs are almost
missing (if present, then below the shell's periphery), its aperture is narrower
but higher, its lower rim is more wavy and larger, and its peristomial insertions
are closer. In the penial lumen, L. spectabilis shows three longitudinal pilas-
ters (instead of two in L. lens); according to Fechter & Falkner (1990 : 222),
its pre-mating behaviour is shorter, and the lack of several behavioural elements
proved to establish a mating barrier.
Broader specimens of L. girva are similar to that form of L. lens, which
inhabits the Island Évvia and the easternmost parts of continental Greece

11
Revision of the genus Lindholmiol a Hesse, 1931
Figs 3–6. Lindholmiola lens. Fig. 3: syntype Helix lentiformis, ZMZ 508 656, "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. Jet schin ex Boettger, 1883, D = 11.94 mm. — All
photos Neubert/Bochud, × 3.

12 Peter Subai & Eike Neub ert
(= lentiformis). However, the shell of L. girva is less flattened, it has more
rounded and less shouldered whorls; its radial riblets are less regular, and its
aperture is more strongly bent in lateral view; its penis is longer, but the glan-
dular tissue is less developed; the penial lumen is filled with 2–3 pilasters
which are not broadened at their proximal end.
Shells of L. barbata are usually more rounded with a blunt keel. Shells of
the same size differ by having an additional whorl, and the umbilicus is almost
completely covered by the peristomial columellar reflection (for more details
refer to L. barbata).
Type specimens: Greece, Ilía (= Elis), Koúmanis (= Kumani), UTM EG 68, 37.78°N
21.74°E, leg. Brenske, 1883, syntypes lens var. elia : SMF 101961/6, 225646/1,
225647/2. — Etoloakarnanía, Náfpaktos (= Lèpante, Lepanto), UTM EH 74, 38.39°N
21.82°E, syntype lens var. callojuncta: NHMG 372.
Additional specimens examined:
Greece, Makedonía, Vérmio Mountains, about 6 km from the highway Véria–Kozáni
direction Séli (= Áno Vérmio), conglomerate rocks at river bed, 530 m alt., UTM EK 98,
40.4971°N 22.1253°E, leg. Fauer & Subai 24.09.1989, S 15686/1; gorge 2 km N of Neo-
hóri (= NE of Aridéa), on limestone rocks, 250 m alt., UTM EL 94, 41.0475°N
22.0916°E, leg. Subai 09.05.1995, S 15922/13 + 1 (alk.); leg. Subai & Szekeres,
19.5.1997, S 15923/5; 8.5 km NNE of Aridéa, on limestone rocks 3 roadkm. N of Vorinó,
250 m alt., UTM EL 94, 41.06°N 22.09°E, leg. Gittenberger & Uit de Weerd 9.5.2000,
NNM/6; 9 km NNE of Aridéa, isolated rocks 6.3 roadkm. N of Vorinó, 440 m alt., UTM
EL 94, 41.08°N 22.06°E, leg. Gittenberger & Uit de Weerd 9.5.2000, NNM/2 + 1 (juv.);
in Véria, on loess wall beside the rivulet, 135 m alt., UTM FK 08, 40.5260°N 22.1997°E,
leg. Fauer & Subai 24.09.1989, S 15925/3 + 1 (juv.); forest in rocky gorge near Néa
Pandeleimónas (= 18 km NW of Stómio), 100 m alt., UTM FK 32, 40.00°N 22.59°E, leg.
Gittenberger & Uit de Weerd 24.5.1999, NNM/1 ( juv.); Halkidikí Peninsula, 500 m N of
Eleohória, on low limestone rocks, 170 m alt., UTM FK 86, 40.3434°N 23.1613°E, leg.
Subai 05.05.1995, S 15684/11; Halkidikí Peninsula, in valley beside the cave of
Petrálona, 360 m alt., UTM FK 87, 40.3759°N 23.1681°E, leg. Subai 05.05.1995, S
15924/24; leg. Subai 29.5.1997, S 15682/2; rocky mountain slope 11.5 km ESE of
Aridéa, NE border of Theodorákio, 450 m alt., UTM FL 03, 40.95°N 22.19°E, leg. Gitten-
berger & Uit de Weerd 10.5.2000, NNM/2; 500 m W of Gríva (= WSW of Axioúpoli), on
limestone rocks, 540 m alt., UTM FL 13, 40.9562°N 22.4009°E, leg. Subai & Szekeres
Figs 7–11. Lindholmiola lens. Fig. 7: NMBE 515616, Greece, Ilía (= Elis), Alfiós canal between Olim-
bía 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.

13
Revision of the genus Lindholmiol a Hesse, 1931

14 Peter Subai & Eike Neub ert
20.05.1997, S 15683/4; N border of Asvestohóri (= E of Thessaloníki), on dry lime-
stone rocks, 40 0 m alt., UTM FL 70, 40.6 466°N 23.0236°E, leg. Subai & Szekeres
8.05.1997, S 15688/1; between Rizá and Paleohóra (= 12.5 km W of Arnéa), 500 m alt.,
UTM GK 08, 40.50°N 23.44°E, leg. Gittenberger & Uit de Weerd 22.5.199 9, NNM/1
(juv.); Kassándra Peninsula, limestone rocks near Loutrá, near sea, 20–25 m alt., UTM
GK 22, 39.9235°N 23.5915°E, leg. Subai 05.05.1995, S 15681/66; Sithonía Peninsula,
mountain at S border of Porto Koufós, on limestone rocks in quarry, 30 m alt., UTM GK
42, 39.9576°N 23.9263°E, leg. Subai 06.05.1995, S 15631/1; Sithonía Peninsula,
mountain 1.5 km S of Porto Koufós, 10 m alt., UTM GK 42, 39.9494°N 23.9348°E, leg.
Subai 05.05.1995, S 15629/110. — Thessalía, E border of Theópetra (= 6 km SE of
Kalambáka), surrounding of church, UTM EJ 59, 39.67°N 21.68°E, leg. Fauer 21.8.1982,
Maa/7, S 12218/5; 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 3781/145, 3784/14; leg. Neuteboom 25.7.1979, NNM/2; leg. Git-
tenberger & Uit de Weerd 25.5.1999, NNM/16 + (alk.); 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 3785/5; N
border of Morfovoúni, on E exposed limestone rocks, 840 m alt., UTM EJ 65,
39.3567°N 21.7464°E, leg. Subai & Szekeres 12.05.1997, S 15928/2 + 1 (juv.); road
border in Petrotó, 100 m alt., UTM EJ 87, 39.58°N 21.95°E, leg. Gittenberger & Uit de
Weerd 25.5.1999, NNM/4; W border of Petrotó, on low limestone rocks, 120 m alt.,
UTM EJ 88, 39.5840°N 21.9584°E, leg. Subai, 18.05.1995, S 15691/1 (juv.); 10 km E of
Kefalóvrisso, 250 m alt., UTM EK 81, 39.92°N 21.98°E, leg. Gittenberger & Uit de
Weerd 25.5.1999, NNM/9; between Elassóna and Deskáti (= 9.5 km W of Elassóna),
250 m alt., UTM EK 91, 39.88°N 22.08°E, leg. Gittenberger & Uit de Weerd 25.5.1999,
NNM/2; 17 km W of Lárissa, bifurcation to Temkat, UTM FJ 08, 39.61°N 22.20°E, leg.
Erőss 9.8.1993, S, 18258/1; between Tírnavos and Mesohóri, 8 km along road SE of
Tírnavos, 125 m alt., UTM FJ 09, 39.71°N 22.24°E, leg. Gittenberger & Uit de Weerd
25.5.1999, NNM/19 + (alk.); road border 1 km W of Damássi, 150 m alt., UTM FJ 09,
39.71°N 22.16°E, leg. Gittenberger & Uit de Weerd 25.5.1999, NNM/1; between Tírna-
vos and Damássi, crossing to Tríkala, on low limestone rocks, 120 m alt., UTM FJ 09,
39.6991°N 22.2134°E, leg. Pintér, E. & P. Subai 28.07.1976, HNHM 85641/1, S
3782/16; 1 km E of Fársala, UTM FJ 15, 39.28°N 22.40°E, leg. Kiss & Pintér 8.7.1986,
HNHM 29905/6; 37.5 km SSE of Lárissa, on limestone rocks 3.5 roadkm. bevor Fársala,
200 m N of road Domokós–Lárissa, 425 m alt., UTM FJ 15, 39.28°N 22.32°E, leg. Gitten-
berger & Uit de Weerd 21.5.200 0, NNM/1; 2 km E of Fársala (= direction Ambelía),
UTM FJ 25, 39.30°N 22.41°E, leg. Neuteboom 21.7.1979, NNM/2; Míra, N slope, on
rocks, UTM FJ 25, 39.45°N 22.54°E, leg. Neuteboom 21.7.1979, NNM/1; Míra, path to
Sofó, UTM FJ 36, 39.45°N 22.53°E, leg. Neuteboom 21.7.1979, NNM/12; 6 km SE of
Almirós, W border of junction, 25 m alt., UTM FJ 53, 39.14°N 22.82°E, leg. Gittenberger
& Uit de Weerd 29.5.1999, NNM/24; 3 km S of Aerinó, W rim of highway, 150 m alt.,
UTM FJ 55, 39.30°N 22.74°E, leg. Gittenberger & Uit de Weerd 29.5.1999, NNM/4; Néa
Ionía (= Vólos, Volo), UTM FJ 65, 39.38°N 22.92°E, HNHM 39177/3, SMF 101969/5; leg.
Stussiner, 1886, SMF 101964/4; leg. C. R. Boettger, 1906, SMF 101978/2; Néa Ionía,
near of beach, UTM FJ 65, 39.34°N 22.92°E, leg. Schütt, 19.5.1962, SMF 279326/7;
road border 500 m E of Ahíllio (= Akhilion), 25 m alt., UTM FJ 71, 39.01°N 22.97°E, leg.

15
Revision of the genus Lindholmiol a Hesse, 1931
Gittenberger & Uit de Weerd 29.5.1999, NNM/12; surroundings of Agía Kiriakí Village
(= 32 km SE of Néa Ionía), 50 –100 m alt., UTM FJ 72, 39.09°N 23.07°E, leg. Gittenberg-
er & Uit de Weerd 28.5.1999, NNM/6; 1 km E of Agriá (= 5 km ESE of Néa Ionía), 125 m
alt., UTM FJ 75, 39.33°N 23.04°E, leg. Gittenberger & Uit de Weerd 26.5.1999, NNM/9;
SE border of Néa Ionia (= Volos) W of the freight harbor, on rocks, among weeds, UTM
FJ 75, 39.3492°N 22.9768°E, leg. Subai 30.04.1991, S 15926/11; Makrinítsa at Pílio
[Mountains], UTM FJ 76, 39.40 °N 22.98°E, leg. Stussiner, 1887, SMF 284934/3,
Figs 12–14. 12: Genital organs of Lindholmiola species. Fig. 12 : Lindholmiola lens, a: situs, Greece,
Ilía (= Elis) between Olimbía and Kréstena, b: lumina of genital organs, Greece, Ahaïa, 3 Km S of
Ptéri ; Fig. 13: Lindholmiola spectabilis, a: situs, b: lumina of genital organs, Greece, Makedonía,
Falakró Mts., Ágios Pnevma plateau; Fig. 14 : Lindholmiola gir va, a: situs, b: lumina of genital organs,
Greece, Makedonía, Taxiárhis.

16 Peter Subai & Eike Neub ert
284955/1; 500 m SSW of Makrinítsa, 275 m alt., UTM FJ 76, 39.39°N 22.97°E, leg. Git-
tenberger & Uit de Weerd 27.5.1999, NNM/33; Pílio (= Pelion) [Mountains], UTM n.
det., SMF 101870/20; foot of Pílio [Mountains] (= Pílios), UTM n. det., leg. Stussiner,
1888, SMF 284933/3; Pílio, 10 min. below peak, UTM FJ 76, 39.43°N 23.04°E, leg.
Stussiner, 1888, SMF 284931/4; Pílio, 1 hour below peak, UTM FJ 76, 39.42°N 23.05°E,
leg. Stussiner, 1888, SMF 284932/2; Survias monastery, N side of Pílio, UTM FJ 76,
39.4620°N 22.9847°E, leg. Stussiner, 1888, SMF 284935/2; 4 km above Portaria (=
direction Hánia), karst slope, 840 m alt., UTM FJ 76, 39.3704°N 23.0290°E, leg. Subai
& Szekeres 9.05.1997, S 15929/2 + 1 (juv.) + 1 (alk.); near Hánia (= ENE of Néa Ionia =
Volos), in forest beside the Hotel "Hani Zisi", in leaf litter, 1150 m alt., UTM FJ 76,
39.3949°N 23.0619°E, leg. Subai & Szekeres 9.05.1997, S 15913/2 (juv.) + 11 (alk.);
road border at Kóttes (= 30.5 km SE of Néa Ionía), 70 m alt., UTM FJ 83, 39.11°N
23.09°E, leg. Gittenberger & Uit de Weerd 28.5.1999, NNM/7; 4 km WSW of Marathéa
(= 32 km SE of Néa Ionía), 15 m alt., UTM FJ 83, 39.11°N 23.12°E, leg. Gittenberger &
Uit de Weerd 28.5.1999, NNM/35 + 7 + (alk.); Olimbos (= Olympos) [Mountain] , UTM n.
det., leg. Maltzan, 1883, SMF 284954/1; leg. Mahunka 15.9.1984, HNHM 23886/14 +
8 (juv.); Témbi (= Tempe valley, approx. 30 km NE of Lárissa), on limestone rocks,
UTM FK 31, 39.87°N 22.58°E, leg. Stussiner, 1884, SMF 284939/4; Témbi (= Tempe
valley, approx. 30 km NE of Lárissa), on limestone rocks and from the "Daphne"
spring, UTM FK 31, 39.8796°N 22.5870°E, leg. Pintér, E. & P. Subai 28.07.1976, HNHM
39188/12, S 3783/14 + 1 (juv.); leg. Neuteboom, 19.7.1979, NNM/2 + 3; leg. Maassen
7.1980, Maa/3 + 26; leg. Subai 24.7.1990, S 15927/5; small valley along old road 500
m SW of Omólion (= Moli), 50 m alt., UTM FK 31, 39.89°N 22.63°E, leg. Gittenberger &
Uit de Weerd 26.5.1999, NNM/1; Ossa Mountains, Profítis Ilías peak region, on marble
rocks, 1550 m alt., UTM FK 40, 39.7948°N 22.6710°E, leg. Subai 23.07.1990, S
15930/4 + 1 ( juv.); Ossa Mountains, Bugari Valley, UTM n. det., leg. Stussiner, leg.,
SMF 284938/2; Thessaloníki (= Saloniki), UTM n. det., leg. Haussknecht, 1885, SMF
284941/1; Langadás near Thessaloníki, UTM FL 71, 40.74°N 23.06°E, SMF 207803/9.
— Fthiótida, plane between Brálos and Parnassós [Mountains], UTM FH 28, 38.69°N
22.45°E, leg. Pfeiffer, 1926, SMF 101876/6; Brálos–Loutrá Thermopilón highway, 8 km
from bifurcation Moni Damasta direction Loutrá Thermopilón, on rocks, 110 m alt.,
UTM FH 29, 38.8008°N 22.4973°E, leg. Subai 10.04.1986, S 15795/1; entrance of
gorge at SE border of Amfíklia, in the vicinity of a small chapel, on limestone rocks,
500 m alt., UTM FH 37, 38.6306°N 22.5963°E, leg. Subai 10.04.1986, S 15696/2;
Loutrá Thermopilón, monument, UTM FH 39, 38.79°N 22.53°E, leg. Kiss & Pintér
9.7.1986, HNHM 29917/1; gorge W of Tithoréa and in the big cave, 850 m alt., UTM FH
47, 38.5702°N 22.6602°E, leg. Subai 11.04.1986, S 15694/4; SW border of Atalánti, on
limestone rocks, 190 m alt., UTM FH 77, 38.6534°N 22.9849°E, leg. Subai 11.04.1986,
S 15695/3; Atalánti (= Atalandi), UTM FH 78, 38.65°N 22.99°E, SMF 284950/4; leg.
Thiesse, 1890, SMF 284897/6, 284937/6; leg. Clessin, 1879, SMF 101859/1; Ipáti, on
limestone rocks beside the waterfall, 380 m alt., UTM FJ 00, 38.8710°N 22.2440°E, leg.
Subai 16.05.1991, S 15711/2; 1.5 km NE of Lamía (= highway direction Lárissa), stone
mine beside Xerias river, on limestone rocks, 615 m alt., UTM FJ 20, 38.9 673°N
22.3874°E, leg. Subai 10.04.1986, S 15697/6; 9 km from Glífa direction Pelasgía, on
limestone rocks, 100 m alt., UTM FJ 61, 38.9309°N 22.9229°E, leg. Subai 30.04.1991, S
15706/19. — Etoloakarnanía, Veloútsa Mountain SE slope above Astakós, on lime-

17
Revision of the genus Lindholmiol a Hesse, 1931
stone rocks, approx., 185 m alt., UTM EH 06, 38.5370°N 21.0692°E, leg. Subai,
19.05.1991, S 15715/3; Messolóngi, UTM EH 34, 38.37°N 21.43°E, leg. Krüper, 1901,
SMF 101866/4; above Kefalóvrisso, on limestone rocks, 30 m alt., UTM EH 35,
38.4598°N 21.3670°E, leg. Fauer & Subai 30.09.1989, S 15736/18; ruins of Ancient
Kalydon (= 2 km from Evinos bridge direction Messolóngi), under stones, UTM EH 44,
38.3725°N 21.5332°E, leg. Fauer & Subai 01.10.1989, S 15732/42; Krionéri, on lime-
stone rocks on Varássova Mountain, SW slope, 50 –220 m alt., UTM EH 54, 38.3477°N
21.5973°E, leg. Fauer & Subai 01.10.1989, S 15731/5; in Riza village, on limestone
rocks and mountain slope W of Riza, beside the old road to Messolóngi, 70 m alt.,
UTM EH 64, 38.3541°N 21.6978°E, leg. Fauer & Subai 02.10.1989, S 15735/13; Náfpa-
ktos (= Lepanto), UTM EH 74, 38.39°N 21.82°E, SMF 285762/7; leg. Krüper SMF
101895/23, 101898/6; leg. Thiesse, 1890, SMF 284956/6; on limestone rocks in Náf-
paktos, 20 m alt., UTM EH 74, 38.3929°N 21.8259°E, leg. Pintér, E. & P. Subai
31.07.1976, HNHM 39184/2 + 3 (juv.), S 3799/3. — Fokída, mountain W of Glifáda (=
opposite of Trizónia Island), on limestone rocks, 100 m alt., UTM EH 94, 38.3883°N
22.0711°E, leg. Subai 14.04.1986, S 15720/5; leg. Kiss & Pintér 23.7.1986, HNHM
29900/8 + 6 (juv.); leg. Riedel & Subai 15.4.1988, S 15739/36; border of Ágios Spirído
Village (= E of Glifáda), on dry limestone rocks, 30 m alt., UTM EH 94, 38.3679°N
22.1169°E, leg. Fauer & Subai 01.10.1989, S 15734/16; 6 km from Erátini direction Náf-
paktos, on rocks at sea shore, 40 m alt., UTM FH 04, 38.3316°N 22.1907°E, leg. Subai
14.04.1986, S 15723/5; 4 km S of Malandrino (= S of Lidoríki), on limestone rocks,
550 m alt., UTM FH 05, 38.4178°N 22.2498°E, leg. Subai 13.4.1986, S 15730/7;
approx. 200 m from the crossing Lidoríki–Erátini–Amfissa direction Amfissa, on lime-
stone rocks, 640 m alt., UTM FH 15, 38.4052°N 22.2550°E, leg. Fauer & Subai
01.10.1989, S 15733/8; 9 km from Mórnou dam direction Lévka, W side of a peninsula
on S banks of the dam reser voir, on limestone rocks, 550 m alt., UTM FH 06,
38.5342°N 22.1634°E, leg. Subai 13.04.1986, S 15729/3; 3 km from Penteória direction
Erátini, on limestone rocks, 750 m alt., UTM FH 15, 38.4155°N 22.2967°E, leg. Subai
14.04.1986, S 15722/8; Lidorikíou Mountains, 28 km from Amfissa direction Lidoríki
(= 19 km S of Lidoríki), 600 m alt., UTM FH 15, 38.3992°N 22.2696°E, leg. Subai
13.04.1986, S 15728/2; 7.5 km from Itéa direction Galaxídi, UTM FH 25, 38.42°N
22.39°E, leg. Neuteboom 22.7.1979, NNM/8 + 1 (juv.); 2 km S of Ámfissa (= direction
Vouníhora), on limestone rocks, 300 m alt., UTM FH 26, 38.5087°N 22.3821°E, leg.
Subai 16.05.1991, S 15718/16; Eptálofos (= Agóriani), Parnassós Mountains, UTM FH
27, 38.59°N 22.49°E, SMF 28 4867/3; leg. Krüper, 1888, SMF 101984/4; leg. Krüper,
1899 and, 1901, SMF 101923/8; leg. Schlüter, 1888, SMF 284864/1; 2.5 km of Graviá
direction Amfissa, on limestone rocks, 540 m alt., UTM FH 27, 38.6607°N 22.4253°E,
leg. Subai 12.04.1986, S 15737/4; E-exponated rocks at S border of Marioláda, 450 m
alt., UTM FH 27, 38.65°N 22.46°E, leg. Gittenberger & Uit de Weerd, 19.5.2000,
NNM/18; 2 km SE von Marioláda, an Kalkfelsen, 780 m alt., UTM FH 27, leg. Gitten-
berger & Uit de Weerd, 19.5.2000, NNM/5; E border of Desfína, UTM FH 35, 38.41°N
22.53°E, leg. Hoenselaar 27.5.1996, Maa/2; NE of Delfí, ruins of ancient Delphi, 650 m
alt., UTM FH 36, 38.4824°N 22.4981°E, SMF 207815/9; leg. Müller, SMF 285174/1; leg.
Krüper, SMF 101896/7; leg. Pfeiffer, 1926, SMF 101892/3, 101893/10, 101894/1; leg.
Schütt 4.6.1969, S 3798/3; leg. Neuteboom 22.7.1979, NNM/7 + 3 (juv.); leg. Drimmer
23.5.1982, HNHM 33703/5; leg. Podani 30.7.1984, HNHM 21467/3; leg. Neuteboom

18 Peter Subai & Eike Neub ert
26.10.1984, NNM/3; leg. Perjesi 29.4.1985, HNHM 23902/1; leg. Kiss & Pintér
23.7.1986, HNHM 29895/18 + 2 (juv.); leg. Sín 6.7.1987, HNHM 44341/5 + 3 (juv.); leg.
Erőss 6.4.1989, S, 18259/3; leg. Subai 16.5.1991, S 15719/35; leg. Hoenselaar
26–27.5.1996, Maa/1 + 2; 2.5 km E of Amfíklia, UTM FH 37, 38.64°N 22.62°E, leg. Neu-
teboom 25.10.1984, NNM/1 + 2 (juv.); Polídrosos, UTM FH 37, 38.63°N 22.53°E, leg.
Neuteboom 25.10.1984, NNM/1; Parnassós Mountains (= Parnass), UTM n. det., SMF
101858/7, 101882/1; leg. Krüper, 1883, SMF 101872/3; leg. Schmidt, 1935, SMF
101878/3; Parnassós Mountains, tree-line 1.7 km of ski-lift, UTM FH 46, 38.55°N
22.57°E, leg. Neuteboom 23.7.1979, NNM/2 + 3 (alk.). — Viotía, Tríodos, bifurcation
after Dístomo, UTM FH 45, 38.44°N 22.68°E, leg. Neuteboom 23.7.1979, NNM/25 km
from the crossing Dístomo–Delfí–Livadiá direction Livadiá, on limestone rocks, 310 m
alt., UTM FH 45, 38.4627 °N 22.7161°E, leg. Subai 14.04.1986, S 15794/1; Elikón
Mountains, 6 km from Kiriáki direction Dístomo and 4 km southward on a small path,
upper border of a gorge, on limestone rocks, 100 m alt., UTM FH 54, 38.3625°N
22.7208°E, leg. Subai 14.04.1986, S 15721/1; 3 km from Paralía Sarandi direction
Pródromos, on limestone rocks and in debris among rocks, 220 m alt., UTM FH 63,
38.2572°N 22.8756°E, leg. Subai 15.04.1986, S 15725/14; 4.3 km from Paralía Saran-
di direction Pródromos, on limestone rocks and in an olive tree plantation, 225 m alt.,
UTM FH 63, 38.2578°N 22.8826°E, leg. Subai 15.04.1986, S 15724/5; rock gorge at SE
border of Livadiá, on limestone rocks and in organic flotsam of creek, 220 m alt., UTM
FH 65, 38.4329°N 22.8920°E, leg. Subai 15.04.1986, S 15796/13; Lafísti (= Lofistion,
E of Livadiá), on limestone rocks, 310 m alt., UTM FH 65, 38.4281°N 22.9189°E, leg.
Subai 11.04.1986, S 15799/3; Herónia, ancient theatre 6 km NW of Livadiá, 250 m alt.,
UTM FH 66, 38.49°N 22.84°E, leg. Gittenberger & Uit de Weerd, 19.5.2000, NNM/1; 2
km E of Alíartos, on limestone rocks, approx. 80 m beside highway, 115 m alt., UTM FH
84, 38.4351°N 23.1997°E, leg. Riedel & Subai 09.04.1988, S 15761/1 + 3 (juv.); 13 km
W of Thíva, 9.9 roadkm. of Alíartos additional 2 km on a secondary road, 250 m alt.,
UTM FH 94, 38.33°N 23.14°E, leg. Gittenberger & Uit de Weerd 20.5.2000, NNM/5; on
limestone rocks beside the motor way, at the bifurcation to Akréfnio (= 5.5 km SE of
the crossing near Kástro), 120 m alt., UTM FH 95, 38.4500°N 23.2092°E, leg. Subai &
Szekeres 11.05.1997, S 15813/6; 7 km N of Thíva, on rocks 1 roadkm. along a second-
ary road to Movríki, UTM GH 05, 38.38°N 23.32°E, leg. Gittenberger & Uit de Weerd
20.5.2000, NNM/2 + 1 (juv.); Skoponéri Village, about 50 m S of the tavern, on low
limestone rocks, approx. 10 m alt., UTM GH 06, 38.5032°N 23.3549°E, leg. Subai &
Szekeres 11.05.1997, S 15690/47 + 2 (juv.) + 9 (alk.); above Skoponéri, E border of the
village, on limestone rocks, 270 m alt., UTM GH 06, 38.5013°N 23.3675°E, leg. Subai &
Szekeres 11.05.1997, S 15689/8 + 1 (juv.); mountain slope 10 km from Halkída (=
Chalkis) direction Thiva (= Thebe), on limestone rocks, 270 m alt., UTM GH 25,
38.4282°N 23.5309°E, leg. Riedel & Subai 09.04.1988, S 15698/7. — Atikí (= Attika),
UTM n. det., SMF 284881/7, 284893/2, 284894/1, 284980/3; leg. Krüper, 1883, SMF
101861/8; approx. 8.5 km from Paralía Shínou direction Káto Alepohóri, on limestone
rocks, approx. 10 m alt., UTM FH 81, 38.0514°N 23.0571°E, leg. Subai 14.05.1991, S
15712/7 + 1 (juv.); Aigósthena (= Porto Germeno), on rocks below the castle ruins, 45
m alt., UTM FH 92, 38.1483°N 23.2321°E, leg. Subai 16.05.1991, S 15693/2 (frag-
ment); 11 km from the highway Vília–Aigósthena direction Kithairón peak, on lime-
stone rocks, 1300 m alt., UTM FH 92, 38.1835°N 23.2583°E, leg. Subai 16.05.1991, S

19
Revision of the genus Lindholmiol a Hesse, 1931
15915/7 + 1 (juv.); Vouliagméni (= opposite of Fléves Island), on limestone rocks
beside the highway, 20 m alt., UTM GG 48, 37.8086°N 23.7841°E, leg. Subai,
19.07.1979, S 15701/51; 3.4 km SE of Varkíza (= coast road direction Agía Marína), on
limestone rocks, 30 m alt., UTM GG 48, 37.8127°N 23.8314°E, leg. Subai 07.05.1991, S
15705/1 (fragment); "Varkíza Beach" camping near Varkíza, on limestone rocks, 30 m
alt., UTM GG 48, 37.8226°N 23.8170°E, leg. Subai 20.07.1979, S 6644/10; 3 km from
Vári direction Koropí, on limestone rocks, 50 m alt., UTM GG 49, 37.8415°N 23.8215°E,
leg. Subai 20.07.1979, S 6700/2; coastal strip near Paleá Fókaia, UTM GG 57, 37.71°N
23.94°E, leg. Neuteboom 20.10.1984, NNM/1; 3 km N of Legrená, UTM GG 67, 37.69 °N
24.00 °E, leg. Neuteboom 20.10.198 4, NNM/3; coast near Megara, UTM GH 00,
37.97°N 23.32°E, leg. Pfeiffer, 1926, SMF 101844/1; Sborder of Megara, UTM GH 00,
37.97°N 23.35°E, leg. Kiss & Pintér, 19.7.1986, HNHM 29918/2 + 1 (juv.); mountain
slope at W border of Mégara, on limestone rocks, 65 m alt., 37.9962°N 23.3240°E,
UTM GH 00, leg. Subai 07.07.1979, S 6515/2 + 2 (juv.); mountain slope beside Ágios
Ierothéou monastery (= W of Mégara), 300 m alt., UTM GH 01, 38.093N, 23.2677°E,
leg. Subai 07.07.1979, S 15702/11 + 1 ( juv.); 3.5 km from the highway Vília–Aigósthe-
na direction Kithairón peak, rocks in coniferous forest, 850 m alt., UTM GH 02,
38.1706°N 23.2728°E, leg. Subai 16.05.1991, S 15692/2 (fragment); Elefsína–Thíva
country road, mountain behind Mándra, UTM GH 11, 38.08°N 23.48°E, leg. Pfeiffer,
1926, SMF 101875/4; Pérama, on rocks at coastal strip direction Piráeus, UTM GH 20,
37.96°N 23.60°E, leg. Neuteboom 4.9.1982, NNM/1; Elefsína, dry mountain slope
beside a quarry, 130 m alt., UTM GH 21, 38.0508°N 23.4950 °E, leg. Subai 07.07.1979,
S 6559/1; Athína (= Athen), UTM n. det., SMF 284878/9, 284879/2, 284946/2; leg.
Krüper, 1883, SMF 101863/5, 101972/11; leg. Heldreich, SMF 284882/3; Piráeus,
UTM GH 30, 37.94°N 23.63°E, leg. Krüper, 1883, SMF 101983/3; Athína, Paleo Falíro (=
Phaleron, part of Athína), UTM GH 30, 37.92°N 23.70°E, leg. Krüper, SMF 101975/39;
Athína, Akropolis, UTM GH 30, 37.97°N 23.72°E, leg. Pfeiffer, 1926, SMF 101843/24;
Párnis Mountain (= Parnés) Chasia, UTM n. det., leg. Krüper, 1883, SMF 101971/1; W
side of Párnitha Mountain (= N of Athína), river valley along the circular path on top of
the mountain, on limestone rocks, 100 0 m alt., UTM GH 32, 38.1631°N 23.7020°E, leg.
Subai 21.04.1988, S 15916/13; SW border of Avlóna, on limestone rocks, 300 m alt.,
UTM GH 33, 38.2454°N 23.6855°E, leg. Subai 16.04.1986, S 15801/2; leg. Riedel &
Subai 9.4.1988, S 15804/15; leg. Gittenberger & Uit de Weerd 20.5.2000, NNM/13;
Likavittós (= Lykabettos) Mountain in Athína, on limestone rocks, UTM GH 4 0,
37.98°N 23.74°E, SMF 101874/2, 285169/24, 285170/2; leg. Krüper, 18 83, SMF
101860/1; leg. Heldreich, SMF 284892/3; leg. Coneménos, 1889, SMF 284886/9; leg.
Krüper, 1888, SMF 101985/9; leg. Pintér 21.8.1976, HNHM 39181/2 + 1 (juv.); leg.
Perjesi 22.4.1985, HNHM 23890/1; top of Likavittós Mountain in Athína, on limestone
rocks, 250 m alt., UTM GH 40, 37.9826°N 23.7436°E, leg. Subai, 19.7.1979, S 6249/12;
leg. Subai 22.04.1988, S 15704/2; Hyméttos Mountain, UTM GH 40, 37.96°N 23.81°E,
leg. Schmidt, SMF 285168/2; leg. Köhler, SMF 285167/2; leg. Knobbe, SMF 284885/2;
leg. Brenske, 1883, SMF 101963/2, 284888/2; leg. Krüper, 1883, SMF 101862/6; leg.
Krüper, 1899, SMF 101967/10; leg. Pfeiffer, 1926, SMF 101954/1; N side of mount
Hyméttos, on limestone rocks, approx. 800 m alt., UTM GH 40, 37.9604°N 23.8162°E,
leg. Subai 4.4.1986, S 15713/5 + 1 (juv.) 15714/100 + 3 (juv.); leg. Subai 21.4.1988, S
15710/3; Kifissiá (= Kephissia), UTM GH 41, 38.07°N 23.82°E, leg. Pfeiffer, 1926, SMF

20 Peter Subai & Eike Neub ert
101886/3; mountain SW of Anatolí (= W of Néa Mákri), on limestone rocks, approx.
200 m alt., UTM GH 51, 38.0895°N 23.9468°E, leg. Subai 02.05.1991, S 15808/14;
Pendéli (= Pentelikon) [Mountains], UTM n. det., SMF 207814/2; leg. Krüper, SMF
101974/40; leg. Pfeiffer, 1926, SMF 101888/1; Pendéli monastery, UTM GH 51,
38.05°N 23.86°E, leg. Pfeiffer, 1926, SMF 101887/1; approx. 4 km NE of Pendéli,
Pendéli (= Pentelikon) Mountains W side, on crystalline rocks, 700 m alt., UTM GH 51,
38.0653°N 23.9 081°E, leg. Subai 02.05.1991, S 15809/12; Soúnio, UTM KB 37,
37.65°N 24.02°E, leg. Pfeiffer, 1926, SMF 101885/29 ; 4 km from Lavrio direction
Markópoulo, on limestone rocks, 125 m alt., UTM KB 38, 37.7523°N 24.0260°E, leg.
Subai 05.05.1991, S 15703/15; road Athína–Soúnio, before Lavrio, UTM KB 48,
37.74°N 24.05°E, KB 38, leg. Pfeiffer, 1926, SMF 101884/5; road Athína–Marathónas,
UTM n. det., leg. Pfeiffer, 1926, SMF 101890/9; road Athína–Soúnio, UTM n. det., leg.
Pfeiffer, 1926, SMF 101883/7. — Égina Island, UTM n. det., SMF 207811/2; Mesagrós,
Aphaia holy place, UTM GG 28, 37.75°N 23.53°E, leg. Erőss 5.4.1989, S, 18261/10. —
Méthana Island, Méthana Town, UTM GG 25, 37.58°N 23.39°E, leg. Haussknecht, 1885,
SMF 284887/2. — Kíthira Island (= Cerigo), "Keramoto", UTM n. det., leg. Krüper,
1893, SMF 101869/8, 284913/2. — Kefaloniá Island, UTM n. det., HNHM 39171/1,
39175/2, 39179/2, SMF 101857/6, 101881/4, 84910/9, 284911/1; leg. Müller, SMF
285162/2; Argostóli Peninsula, UTM DH 52, 38.18°N 20.48°E, leg. Visser 1.11.1985,
NNM/1 (alk.). — Zakinthos Island (= Zante), UTM n. det., SMF 284884/5; above Chili-
omenon, UTM n. det., HNHM 39236/3 + 1 (juv.). — Ilía (= Elis), theatre 15 km E of Gas-
toúni, UTM EG 39, 37.89°N 21.37°E, leg. Hoenselaar 17.4.1996, Maa/5; Kaíafás (= NW
of Zaháro), on limestone rocks and from the thermal well, UTM EG 55, 37.5193°N
21.6022°E, leg. Subai 07.04.1986, S 15798/11; ruins of Arhea Olympia, 35 m alt., UTM
EG 56, 37.6384°N 21.6307°E, HNHM, 18649/1; SMF 279327/4, 284900/10, 285173/6;
leg. Schlüter, SMF 284901/15; leg. Pfeiffer, 1926, SMF 101851/7; leg. Kühnel 4.1933,
HNHM 9784/2; leg. Hansen, 1937, HNHM 39187/2; leg. Jaeckel, 1958, SMF
207816/10; leg. Seidl 3.10.1965, NNM/4; leg. Guyat & Dawber 8.1973, HNHM 85645/1,
S 3793/12; leg. Pintér & Subai 2.8.1976, S 3494/61; leg. Sin 4–5.7.1985, HNHM
24029/17 + 3 (juv.), 24069/25 + 4 (juv.); leg. Kiss & Pintér 23.7.1986, HNHM 29897/9
+ 3 (juv.); leg. Hoenselaar, 19.4.1996, Maa/7; between Arhea Olympia and Kréstena,
beside the channel of Alfiós River, in grass and under stones, UTM EG 56, 37.6279°N
21.5923°E, leg. Subai 10.07.1979, S 15882/36 + 17 (alk.); Giannitsohóri, Camping
"Apollo", UTM EG 63, 37.39°N 21.69°E, leg. Hoenselaar 21–23.4.1996, Maa/2 + 3; river
gorge approx. 1.5 km E of Néa Figália, on limestone rocks, approx. 550 m alt., UTM EG
64, 37.4469°N 21.7890°E, leg. Riedel & Subai, 18.04.1988, S 15762/1; road Tholon–
Andrítsena, border of Lépreo, UTM EG 64, 37.43°N 21.72°E, leg. Hoenselaar 22.4.1996,
Maa/2; Koúmanis (= Kumani), UTM EG 68, 37.79°N 21.74°E, SMF 225648/3; river
gorge between the two bifurcations of the Lámbia–Tripótama highway to Orini, on
limestone rocks, 630 m alt., UTM EG 79, 37.8608°N 21.8612°E, leg. Riedel & Subai
17.04.1988, S 15760/9; leg. Hadjicharalambous & Subai 12.5.1991, S 15756/1. — Ahaïa,
4.1 km from the crossing Kalávrita–Trípoli–Tripótama direction Tripótama, on lime-
stone rocks, 1120 m alt., UTM EG 99, 37.8992°N 22.0087°E, leg. Hadjicharalambous &
Subai 14.05.1991, S 15716/5; dry meadow 2 km S of Klitoria (= direction Trípoli), 50 0
m alt., UTM EG 99, 37.8808°N 22.1355°E, leg. Subai 09.07.1979, S 15738/3; Araxos, on
limestone rocks beside the stone mine at banks of Lake Kalogera, 20 m alt., UTM EH

21
Revision of the genus Lindholmiol a Hesse, 1931
32, 38.1642°N 21.4011°E, leg. Subai 07.04.1986, S 15727/9; Skollis Mountain W slope
above Sandoméri, on limestone rocks, in rock debris, 550 m alt., UTM EH 40,
37.9899°N 21.5781°E, leg. Riedel & Subai 16.04.1988, S 15766/4; W slope of Skollis
Mountain, NE of Sandoméri, on limestone rocks, 530 m alt., UTM EH 50, 37.9929°N
21.5778°E, leg. Subai 07.04.1986, S 15797/9; Erímanthos Mountains above Kaléntzi,
on limestone rocks, 1060 m alt., UTM EH 60, 37.9447°N 21.7650°E, leg. Hadjicharalam-
bous & Subai 13.05.1991, S 15921/4; Pátra (= Patras), UTM EH 63, 38.24°N 21.74°E,
SMF 284890/2, 284908/5, 284953/4; leg. Coneménos, SMF 101854/5; leg. Brenske,
1883, SMF 101968/6; leg. Krüper, 1895, SMF 2849 60/6; leg. Krüper, 1899, SMF
101952/11; leg. C. R. Boettger, 1905, SMF 101979/5; Psathópirgos, UTM EH 74,
38.32°N 21.87°E, SMF 225650/19 + 20; river valley near Psathopírgos, among dry
plants and on rocks (no limestone !), 230 m alt., UTM EH 74, 38.3183°N 21.8823°E,
leg. Subai 07.04.1986, S 15726/5; in a dry river bed S of Flámpoura village (= Metohi),
900 m alt., UTM EH 81, 38.0286°N 21.9965°E, leg. Pintér, E. & P. Subai 01.08.1976, S
3791/8, HNHM 39170/5; bifurcation of the highway Kalávrita–Diakoftó to Rogí, Ker-
piní, on limestone rocks, 700 m alt., UTM EH 91, 38.0534°N 22.1384°E, leg. Hadji-
charalambous & Subai 13.05.1991, S 15717/3; river gorge 3 km S from Ptéri, on lime-
stone rocks and under stones, 1150 m alt., UTM EH 92, 38.1418°N 22.0710°E, leg.
Riedel & Subai 15.04.1988, S 15920/3 + 1 ( juv.) + 2 (alk.); leg. Fauer & Subai
3.10.1989, S 15919/9; leg. Hadjicharalambous & Subai 13.05.1991, S 15780/1; in and
around the cave N of Kastría, on limestone rocks, 840 m alt., UTM FH 00, 37.9603°N
22.1404°E, UTM FH 00, leg. Hadjicharalambous & Subai 14.05.1991, S 15778/2; on
rocks and on walls of Mega Spileo monastery (= N of Kalávrita), 950 m alt., UTM FH 01,
38.0884°N 22.1746°E, leg. Pintér, E. & P. Subai 01.08.1976, S 3792/3; Subai 9.7.1979,
S 6577/1; in "Lemon Beach" camping near Paralia Akráta, under stones, UTM FH 12,
38.1653°N 22.3408°E, leg. Subai 09.07.1979, S/2 (alk.); Nezerá, UTM n. det., leg.
Coneménos, SMF 225649/2. — Korinthía, Killini Mountains, in spruce forest above
Goúra, on limestone rocks, 1100 m alt., UTM FG, 19, 37.9347°N 22.3488°E, leg. Subai
08.07.1979, S 6428/1 + 2 ( juv.); Killini Mountains, 7.5 km from Goúra direction
Kastanéa, on limestone rocks, 800 m alt., UTM FG, 19, 37.8866°N 22.3408°E, leg.
Subai 08.07.1979, S 6573/2; Oligirtos Mountains, 1 km from Kartéri direction Goúra,
on limestone rocks and in cave crevices, 850 m alt., UTM FG 29, 37.8510°N 22.3996°E,
leg. Subai 08.07.1979, S 15750/2; Killini Mountains, mountain slope 6 km from Kaliáni
direction Kiáto, on conglomerate rocks, 710 m alt., UTM FG 39, 37.9236°N 22.5262°E,
leg. Subai 08.07.1979, S 6661/2; 1 km S of Neméa, on limestone rocks, 370 m alt.,
UTM FG 48, 37.8024°N 22.6670°E, leg. Hadjicharalambous & Subai 07.05.1991, S
15781/2; ancient Neméa (= near Arhéa Neméa), UTM FG 58, 38.82°N 22.66°E, leg.
Hoenselaar 22.5.1996, Maa/5; 8.5 km from Ágios Vassílios direction Árgos, on lime-
stone rocks, 290 m alt., UTM FG 58, 37.7779°N 22.7339°E, leg. Subai, 18.07.1979, S
6561/2 + 1(juv.); ruins of Arhea Kórinthos, 80 m alt., UTM FG 69, 37.9059°N 22.8792°E,
SMF 284951/6; leg. Brenske, 1883, SMF 284920/1; leg. Krüper, SMF 101853/5; leg.
Pfeiffer, 1926, SMF 101845/6; leg. Pintér & Subai 6.08.1976, HNHM 39172/4, S
3795/4; leg. Nosek 3.10.1976, HNHM 21461/2; leg. Hoenselaar 23.5.1996, Maa/2;
road to Akrokorinthos, UTM FG 69, 37.89°N 22.87°E, leg. Pfeiffer, 1926, SMF 101846/1
+ 4 ( juv.); leg. Neuteboom 29.8.1982, NNM/1; ruins of the old castle of Akrokorinthos,
400 m alt., UTM FG 69, 37.8900°N 22.8690°E, SMF 285172/2; leg. Schlüter 27.4.1912,

22 Peter Subai & Eike Neub ert
SMF 285765/8; Tomlin, 1904, SMF 284889/1; leg. Subai, 18.7.1979, S 15744/57 + 1
(alk.); leg. Kiss & Pintér 21.8.1988, HNHM 44600/24 +, 18 (juv.); leg. Neuteboom
29.8.1982, NNM/11 + 3 (juv.); highway Epidauros–Kórinthos, 8 km from the bifurca-
tion to Sofikó direction Epidauros, 230 m alt., UTM FG 87, 37.7334°N 23.1010°E, leg.
Pintér, E. & P. Subai 06.08.1976, HNHM 39182/2 + 1 (juv.), S 3796/2; Kórfos, UTM FG
88, 37.76°N 23.12°E, leg. Kiss & Pintér 16.8.1988, HNHM 85644/7; 2 km S of Kórfos,
UTM FG 88, 37.73°N 23.12°E, leg. Kiss & Pintér, 19.7.1986, HNHM 29909/1 (juv.); N
exposed lime hill slope between Ano Loutró and Kariá, 550 m alt., UTM FH 31,
38.0842°N 22.5403°E, leg. Fauer & Subai 03.10.1989, S 15740/1; ruins of Heraion and
on limestone rocks beside Lake Heraiou, 60 m alt., UTM FH 61, 38.0349°N 22.8708°E,
leg. Subai, 18.07.1979, S 6352/13 + 3 (alk.); Loutráki, UTM FH 70, 37.98°N 22.97°E,
leg. Krüper, SMF 101973/10; leg. Butot & Subai, 19.2.1981, S 17540/12 + 3 (alk.); leg.
Kiss & Pintér, 19.7.1986, HNHM 29901/18 + 7 (juv.); leg. Erőss 22.7.1992, S, 18257/1;
dry mountain slope at NW border of Loutráki, on limestone rocks, 80 m alt., UTM FH
70, 37.9836°N 22.9715°E, leg. Subai 07.07.1979, S 15751/15; W border of Perahóra,
UTM FH 71, 38.02°N 22.94°E, leg. Kiss & Pintér 16.8.1988, HNHM 44685/15 + 6 (juv.);
1 km S of Perahóra, UTM FH 71, 38.01°N 22.94°E, leg. Kiss & Pintér 16.8.1988, HNHM
44589/6 + 1 (juv.); approx. 1.5 km from Paralía Shínou direction Káto Alepohóri, on
limestone rocks, approx. 100 m alt., UTM FH 81, 38.0570°N 23.0055°E, leg. Subai
14.05.1991, S 15914/10. — Arkadía, 1 km from Stavrodrómio direction Arhea Olympia
(= highway Arhea Olympia–Trípoli), 630 m alt., UTM EG 87, 37.6989°N 21.9392°E, leg.
Pintér, E. & P. Subai 02.08.1976, S 3775/10; 2 km from Langádia direction Arhea Olym-
pia (= highway Arhea Olympia–Trípoli), 850 m alt., UTM EG 87, 37.6815°N 22.0147°E,
leg. Pintér, E. & P. Subai 02.08.1976, S 3777/1; 3 km from Kalonério direction Arhea
Olympia (= highway Arhea Olympia–Trípoli), 1090 m alt., UTM EG 97, 37.6727°N
22.0534°E, leg. Pintér, E. & P. Subai 02.08.1976, S 3776/2; lime mountain N side, 6 km
from Trópea direction Dáfni, approx. 500 m alt., UTM EG 87, 37.7573°N 21.9626°E, leg.
Riedel & Subai 17.04.1988, S 15759/1; 2 km from Paradísia direction Megalópoli
(= highway Megalópoli–Kalamáta), 475 m alt., UTM EG 93, 37.3319°N 22.0743°E, leg.
Pintér, E. & P. Subai 03.08.1976, S 3779/1; 400 from the crossing Dimitsána–Andrítsena–
Megalópoli direction Dimitsána, on limestone rocks along border of road, 355 m alt.,
UTM EG 94, 37.4810°N 22.0492°E, leg. Subai 05.04.1986, S 15784/1; 12.5 km S from
Dimitsána (= crossing Dimitsána–Trípoli–Megalópoli), on limestone rocks along border
of road, 1130 m alt., UTM EG 95, 37.5453°N 22.0894°E, leg. Subai 05.04.1986, S
15786/1 (juv.); S border of Magoúliana, UTM EG 97, 37.67°N 22.12°E, leg. Pintér &
Varga 21.7.1992, HNHM 44846/1; on rocks below Moni Ambeláki (= Taigetos Moun-
tains NE border), 650 m alt., UTM FG 02, 37.2692°N 22.2181°E, leg. Hadjicharalam-
bous & Subai 08.05.1991, S 15773/2; 5 km from Vitína direction Arhea Olympia
(= highway Arhea Olympia–Trípoli), 985 m alt., UTM FG 06, 37.6515°N 22.1685°E, leg.
Pintér, E. & P. Subai 02.08.1976, S 3778/1; leg. Subai 5.4.1986, S 15800/1; leg. Hadji-
charalambous & Subai 8.5.1991, S 15775/1; 3.7 km from Perthóri direction Moni
Epáno Hrépas, on limestone rocks, 1000 m alt., UTM FG 15, 37.5417°N 22.3347°E, leg.
Hadjicharalambous & Subai 08.05.1991, S 15774/3; Levídi, UTM FG 17, 37.68°N
22.29°E, SMF 207810/3; 4.7 km from Levídi direction Kandíla, on limestone rocks,
720 m alt., UTM FG 17, 37.7214°N 22.3276°E, leg. Hadjicharalambous & Subai
07.05.1991, S 15783/10; 4 km from Manthiréa direction Spárti, on limestone rocks,

23
Revision of the genus Lindholmiol a Hesse, 1931
860 m alt., UTM FG 23, 37.3813°N 22.4034°E, leg. Pintér, E. & P. Subai 05.08.1976, S
3780/1; 1.5 km SSE of Kandíla, near Kandílas monastery, 800 m alt., UTM FG 28,
37.7635°N 22.3745°E, leg. Gittenberger 17.8.2000, NNM/5; Parnon Mountains, approx.
5.5 km SSE of Ágios Pétros (= approx. 2.5 km from road junction to Kastanitsa), on
limestone rocks and among rock pebbles, 970 m alt., UTM FG 32, 37.3114°N
22.5775°E, leg. Riedel & Subai 20.04.1988, S 15769/2; Ágios Ioánnis, UTM FG 43,
37.35°N 22.63°E, leg. Neuteboom 4.9.1982, NNM/1; 1 km from Ágios Ioánnis direction
Ástros, on limestone rocks, 750 m alt., UTM FG 43, 37.3539°N 22.6401°E, leg. Subai
16.07.1979, S 6425/9; between Elónis monastery and Kosmās, small caves near Ágios
Dimitri, UTM FG 50, 37.13°N 22.76°E, leg. Neuteboom 30.8.1982, NNM/2 (juv.); SW
border of Ágios Andreas, on limestone rocks, 90 m alt., UTM FG 53, 37.3437°N
22.7690°E, leg. Subai 16.07.1979, S 6424/9; 1 km E of Ágios Andréas, UTM FG 53,
37.34°N 22.78°E, leg. Hoenselaar 14.5.1996, Maa/1; 2 km SE of Ágios Andréas, 25 m
alt., UTM FG 53, 37.33°N 22.79°E, leg. Neuteboom 30.8.1982, NNM/1 + 1 (juv.); 3 km
SE of Ágios Andréas, UTM FG 53, 37.32°N 22.79°E, leg. Kiss & Pintér, 18.8.1988,
HNHM 44594/2 + 1 (juv.); 5 km SE of Ágios Andréas, opposite of Camping "Arkadia",
UTM FG 53, 37.33°N 22.80 °E, leg. Hoenselaar 12.5.1996, Maa/1; leg. Neuteboom
30.8.1982, NNM/1 + 2 (juv.); 3 km SE of Ástros, UTM FG 53, 37.38°N 22.77°E, leg. Kiss
& Pintér, 18.8.1988, HNHM 44593/39 + 7 (juv.); 7 km from Ástros direction Árgos, on
limestone rocks, 75 m alt., UTM FG 54, 37.4663°N 22.7421°E, leg. Subai 16.07.1979, S
15753/14; leg. Neuteboom 4.9.1982, NNM/3 + 3 (juv.); on rocks of coastal road Kivéri–
Ástros, 7.4 roadkm. N of bifurcation to Parálio Ástros, 25 m alt., UTM FG 54, 37.49°N
22.72°E, leg. Gittenberger 25.7.2000, NNM 71842/2; 1 km S von Kivéri, an Kalkfelsen,
UTM FG 55, Subai 17.7.1979, S 15755/13 + 2 (juv.); leg. Kiss & Pintér 17.8.1988, HNHM
44592/1 + 3 (juv.); Pirgoúdi, UTM FG 60, leg. Kiss & Pintér, 18.8.1988, HNHM 85639/4;
on rocks near Livádi (= 5 km NE of Leonídio), 25 m alt., UTM FG 61, 37.19°N 22.89°E,
leg. Gittenberger 20.7.2000, NNM 71861/1; on rocks along the road, 1.7 km NE of
Tsitália, UTM FG 61, 37.13°N 22.87°E, leg. Gittenberger 21.7.2000, NNM 71858/3; on
conglomerate rocks at W border of Leonídio, 100 m alt., UTM FG 61, 37.1694°N
22.8529°E, leg. Subai 16.07.1979, S 6690/39 ; leg. Butot & Subai, 19.2.1981, S
15806/25 + 4 ( juv.); 1 km W of Leonídio, UTM FG 61, 37.16°N 22.83°E, leg. Kiss &
Pintér, 18.8.1988, HNHM 44595/14 + 4 ( juv.); Pláka, UTM FG 61, 37.14°N 22.89°E, leg.
Kiss & Pintér, 18.8.1988, HNHM 85640/1; gorge 4 km WSW of Leonídio, 285 m alt.,
UTM FG 61, 37.15°N 22.79°E, leg. Neuteboom 30.8.1982, NNM/6; leg. Kiss & Pintér,
18.8.1988, HNHM 44596/1; 5.5 km from Ágios Andréas direction Tirós, UTM FG 62,
37.25°N 22.85°E, leg. Neuteboom 30.8.1982, NNM/1; 5 km from Ágios Andreas direc-
tion Tiros, on limestone rocks, 25 m alt., UTM FG 63, 37.3242°N 22.8161°E, leg. Subai
16.07.1979, S 6426/57. — Argolída, mountain pass 7 km from Kandíla direction Psári,
Olígirtos Mountains, on limestone rocks, 1400–1450 m alt., UTM FG 28, 37.7944°N
22.3979°E, leg. Hadjicharalambous & Subai 07.05.1991, S 15782/9; 3 km NW of Ahl-
adókambos, UTM FG 35, 37.54°N 22.53°E, leg. Kiss & Pintér 20.7.1986, HNHM 29907/2
(juv.); 8 km from Ahladókambos direction Trípoli, on limestone rocks, 720 m alt., UTM
FG 35, 37.5356°N 22.5136°E, leg. Pintér, E. & P. Subai 05.08.1976, S 3772/1; mountain
slope 12.5 km from Agiorgítika direction Árgos (= environment of Ahladókambos), on
limestone rocks, 800 m alt., UTM FG 35, 37.5385°N 22.5741°E, leg. Subai 05.04.1986,
S 15787/7; on rocks 1.3 roadkm. E of Andrítsa, 120 m alt., UTM FG 44, 37.47°N

24 Peter Subai & Eike Neub ert
22.62°E, leg. Gittenberger 24.7.2000, NNM 71848/2; Trípoli–Nafplio Landstrasse,
highway Trípoli–Nafplio, 4 km from the bifurcation to Spiliotákis direction Trípoli, on
limestone rocks, 600 m alt., UTM FG 45, 37.5232°N 22.6404°E, leg. Pintér, E. & P.
Subai 05.08.1976, HNHM 39176/4, S 3774/5; mountain slope 7 km from Ahladókam-
bos direction Árgos, on limestone rocks, 650 m alt., UTM FG 45, 37.5181°N 22.6173°E,
leg. Subai 05.04.1986, S 15788/3; leg. Riedel & Subai 20.4.1988, S 15768/1; in
Kefalari, on limestone rocks and from a rich spring, 30–35 m alt., UTM FG 46,
37.5959°N 22.6882°E, leg. Subai, 18.07.1979, S 15743/7; on rocks 12 roadkm. E of
Andrítsa (= 3 km W of Kivéri), 55 m alt., UTM FG 55, 37.50°N 22.68°E, leg. Gittenberger
24.7.2000, NNM 71853/1; Nafplio (= Nauplia, Nauplion), UTM FG 55, 37.56°N 22.81°E,
SMF 101867/6, 101880/1, 207813/2, 284883/6, 284898/3, 284899/3, 284904/2; leg.
Subai, 18.7.1979, S 15742/19 ; leg. Sin 6–7.7.1985, HNHM 24 037/11 + 2 (juv.),
24042/6; leg. Kiss & Pintér, 19.7.1986, HNHM 29891/122 + 46 (juv.); on ruins of the
old castle of Tiryns, 15 m alt., UTM FG 56, 37.5990 °N 22.8001°E, leg. Pintér, E. & P.
Subai 06.08.1976, HNHM 39180/12 + 1 (juv.), S 3769/20; leg. Neuteboom 30.8.1982,
NNM/5; leg. Mészáros 1.5.1986, HNHM 44342/2; leg. Kiss & Pintér, 19.7.1986, HNHM
29913/8 + 4 (juv.); Árgos, on ruins of the old castle, 260 m alt., UTM FG 56, 37.6384°N
22.7150°E, leg. Pintér, E. & P. Subai 05.08.1976, S 3773/15; Subai, 18.7.1979, S
6699/97 + 1 (alk.); Mykenai, UTM FG 57, 37.72°N 22.75°E, SMF 284961/8, 285171/6;
leg. Pfeiffer, 1926, SMF 101847/4; leg. Erőss 21.7.1992, S, 18260/1; ruins of the old
castle of Mykene and in valley beside it, 220 m alt., UTM FG 57, 37.7291°N 22.7577°E,
leg. Pintér, E. & P. Subai 06.08.1976, HNHM 39168/5, S 3770/5; leg. Nosek 29.9.1976,
HNHM 23962/1; leg. Sín 8.7.1985, HNHM 24046/5 + 1 (juv.); leg. Kiss & Pintér
20.7.1986, HNHM 29912/16 + 4 (juv.); leg. Subai 20.4.1988, S 15767/24; Toló, UTM FG
65, 37.52°N 22.85°E, leg. Kiss & Pintér 17.8.1988, HNHM 44591/58 + 13 (juv.); Toló,
Camping "Kastráki", UTM FG 65, 37.52°N 22.85°E, 37.52°N 22.85°E, leg. Hoenselaar
15.5.1996, Maa/9; surroundings of Drépano, UTM FG 65, 37.53°N 22.89 °E, leg.
Mészáros 30.4.1986, HNHM 44340/41 + 30 (juv.); ancient Asine (= near Asíni, SE of
Nafplio), UTM FG 65, 37.52°N 22.87°E, leg. Hoenselaar 15.5.1996, Maa/2; from road
Pirgiótika–Arkadiko towards Ágios Dimítrios, UTM FG 66, 37.58°N 22.93°E, leg. Neute-
boom 4.9.1982, NNM/3; mountain slope 2 km from Agionóri direction Límnes, on
limestone rocks, 700 m alt., UTM FG 67, 37.7439°N 22.8739°E, leg. Subai 04.04.1986,
S 15792/22; mountain slope 3 km from Agionóri direction Límnes, on limestone rocks,
730 m alt., UTM FG 67, 37.7388°N 22.8772°E, leg. Subai 04.04.1986, S 15793/1 + 1
(juv.); Néa Epídavros, bifurcation to Ligoúrio, UTM FG 87, 37.68°N 23.09°E, leg. Neute-
boom 4.9.1982, NNM/8; 1 km N of Néa Epídavros, UTM FG 87, 37.68°N 23.13°E, leg.
Kiss & Pintér 16.8.1988, HNHM 85642/19; 1 km N of Ermióni, UTM FG 94, 37.39°N
23.24°E, leg. Kiss & Pintér 17.8.1988, HNHM 44590/3 + 2 ( juv.); steps of the amphithe-
atre in Epidauros, UTM FG 86, 37.6334°N 23.1599 °E, leg. Pintér, E. & P. Subai
06.08.1976, HNHM 39173/4, S 3771/6; 1 km N of Trahiá, UTM FG 96, 37.58°N 23.15°E,
leg. Kiss & Pintér 16.8.1988, HNHM 85638/12; 1 km W von Eleónas, UTM n. det., leg.
Kiss & Pintér 24.7.1986, HNHM 29898/12 + 3 (juv.); Ília bei Kastrí, UTM n. det., leg.
Philippson 8.9.1887, SMF 284896/7 + 1 (juv.). — Messenía, ruins of the old Navarino
palace (= Paleókástro, S of Petrohóri), under stones, on limestone rocks, 120 m alt.,
UTM EF 58, 36.9582°N 21.6568°E, leg. Riedel & Subai, 18.04.1988, S 15764/6; on
walls of the old castle in Pílos, 30 m alt., UTM EF 68, 36.9118°N 21.6909°E, leg. Subai

25
Revision of the genus Lindholmiol a Hesse, 1931
06.04.1986, S 15790/8; E border of Pílos, on limestone rocks, 50 m alt., UTM EF 68,
36.9204°N 21.7052°E, leg. Subai 11.07.1979, S 15748/3; 6 km from Pílos direction
Kalamáta, on limestone rocks, 70 m alt., UTM EF 68, 36.9373°N 21.7429°E, leg. Subai
06.04.1986, S15791/2; Camping "Navarino Beach" NE of Pílos, UTM EF 68, 36.93°N
21.71°E, leg. Hoenselaar 26.4.1996, Maa/2; cave niches 8 km from Methóni direction
Pílos, on limestone rocks, UTM EF 68, 36.8877°N 21.7044°E, leg. Subai 11.07.1979, S
15749/6; 8 km from Handrinós direction Pílos (= highway Pílos–Messíni), on lime-
stone rocks, 50 m alt., UTM EF 68, 36.9373°N 21.7423°E, leg. Subai 12.07.1979, S
6572/3 + 5 (alk.); Néstor’s Palace between Hóra and Korifási, tomb of Néstor, UTM EF
69, 37.04°N 21.70°E, leg. Hoenselaar 27.4.1996, Maa/2; mountain SW of Hrisokellariá,
on limestone rocks, 400 m alt., UTM EF 77, 36.7864°N 21.8801°E, leg. Hadjicharalam-
bous & Subai 11.05.1991, S 15777/10; Kastánia near Petalidi, UTM EF 89, 36.95°N
21.88°E, leg. Philippson 26.10.1887, SMF 284917/1, 284918/3; Kalamáta, UTM EF 99,
EG 90, 37.02°N 22.11°E, leg. Philippson 14.10.1887, SMF 284895/1; cave niches 1 km
from Gargaliáni direction Kiparissía, on limestone rocks, 20 0 m alt., UTM EG 50,
37.0667°N 21.6199°E, leg. Subai 10.07.1979, S 6666/8; Filiatrá, UTM EG 51, 37.15°N
21.58°E, leg. Thiesse, 1890, SMF 284903/17; 2.9 km from Filiatrá direction Pláti, on
limestone rocks, 140 m alt., UTM EG 51, 37.1657°N 21.6212°E, leg. Hadjicharalambous
& Subai 12.05.1991, S 15757/11; in Touloupa Hani Village (= E of Hóra and Metamór-
fosi), on limestone rocks, 470 m alt., UTM EG 6 0, 37.0621°N 21.7804°E, leg. Riedel &
Subai, 18.04.1988, S 15763/5; E border of Rodiá (= SE of Kiparissía), on limestone
rocks and along river bank, 420 m alt., UTM EG 61, 37.2106°N 21.7336°E, leg. Riedel &
Subai, 18.04.1988, S 15765/2; 1.2 km E of Rodiá, on low limestone rocks, 390 m alt.,
UTM EG 61, 37.2129°N 21.7400°E, leg. Hadjicharalambous & Subai 12.05.1991, S
15758/8; ruins of the old castle in Kiparissía, 100 m alt., UTM EG 62, 37.2506°N
21.6783°E, leg. Subai 10.07.1979, S 6411/42; on dry limestone rocks 13 km N of
Kiparissía (= highway Pírgos–Pílos), 50 m alt., UTM EG 63, 37.3473°N 21.7015°E, leg.
Subai 10.07.1979, S 15881/8; Ithómi (= antic Messene), on ruins of the town walls,
350 m alt., UTM EG 81, 37.1784°N 21.9184°E, leg. Pfeiffer, 1926, SMF 101850/7; leg.
Pintér & Subai 4.8.1976, HNHM 39190/4, S 3788/1; leg. Subai 12.07.1979, S 6410/1;
leg. Kiss & Pintér 22.7.1986, HNHM 29908/5 + 2 (juv.); leg. Hoenselaar 24.4.1996,
Maa/2; 2 km W of the crossing near Lámbena (= N of Messíni), 230 m alt., UTM EG 81,
37.1485°N 21.9418°E, leg. Pintér, E. & P. Subai 04.08.1976, S 3787/10; mountain slope
6.6 km from Lámbena direction Mavrongáti, on limestone rocks, 280 m alt., UTM EG
81, 37.1517°N 21.9299°E, leg. Subai 05.04.1986, S 15785/2; 5.5 km from Ithómi direc-
tion Lámbena, on limestone rocks along border of road, 275 m alt., UTM EG 81,
37.1511°N 21.9299°E, leg. Subai 12.07.1979, S 15754/3; 3 km N of Bassai, UTM EG 84,
37.45°N 21.92°E, leg. Kiss & Pintér 20.8.1988, HNHM 44599/2 + 1 (juv Málta village (=
SE of Kalamáta), on medieval castle ruins, 370 m alt., UTM FF 08, 36.9313°N
22.1915°E, leg. Pintér, E. & P. Subai 04.08.1976, HNHM 39178/9, S 3789/7; 4 km of
Kalamáta, near Camping "Maria’s See and Sun", UTM FF 09, 37.02°N 22.02°E, leg.
Hoenselaar 28.4.1996, Maa/4; above Mikra Mandínia (= Taigetos Mountains SE of
Kalamáta), 120 m alt., UTM FF 09, 36.9845°N 22.1619°E, leg. Pintér, E. & P. Subai
04.08.1976, HNHM 39181/4, S 3790/5; N of Kámbos (= Taigetos Mountains SE of
Kalamáta), on limestone rocks, 210 m alt., UTM FF 09, 36.9556°N 22.1913°E, leg.
Pintér, E. & P. Subai 04.08.1976, HNHM 39183/1; limestone rocks in valley near Langá-

26 Peter Subai & Eike Neub ert
da, 430 m alt., UTM FF 17, 36.7801°N 22.3383°E, leg. Butot & Subai 16.02.1981, S
17531/13; leg. Neuteboom 31.8.1982, NNM/4; Roman stone highway near Plátsa, on
limestone rocks, 370 m alt., UTM FF 17, 36.8074°N 22.3205°E, leg. Butot & Subai
16.02.1981, S 17532/11; border of Plátsa direction Nomítsis, UTM FF 17, 36.80°N
22.31°E, leg. Neuteboom 31.8.1982, NNM/1; 15 km from Pigí direction Kalamáta, on
limestone rocks, 25 m alt., UTM FF, 18, 36.8592°N 22.2564°E, leg. Subai 13.07.1979, S
6427/3; S border of Kardamili, on limestone rocks, UTM FF, 18, 36.8849 °N 22.2356°E,
leg. Butot & Subai 16.02.1981, S 17530/1; leg. Neuteboom 31.8.1982, NNM/5; in valley
W of Kéndro, on limestone rocks and from a spring at river bank, 575 m alt., UTM FF,
18, 36.9504°N 22.2502°E, leg. Subai 06.04.1986, S 15789/1; gorge 3.8 km E of Kéndro
(= Chóra Gaitses), approx. 720 m alt., UTM FF, 19, 36.9749°N 22.2596°E, leg. Hadji-
charalambous & Subai 10.05.1991, S 15770/4; highway Kalamáta–Spárti, start of
Artemíssia gorge (= Taigetos Mountains NE of Kalamáta), 710 m alt., UTM FG 00,
37.0298°N 22.1865°E, leg. Pintér, E. & P. Subai 05.08.1976, HNHM 39185/15 + 2 ( juv.),
S 3786/16; Artemissía gorge 9.4 km from Kalamáta direction Artemissía, on limestone
rocks, 310 m alt., UTM FG 00, 37.0840°N 22.1575°E, leg. Hadjicharalambous & Subai
11.05.1991, S 15776/41 + 4 (juv.); Dimióvis monastery (= Taigetos Mountains NE of
Kalamáta), on limestone rocks, 820 m alt., UTM FG 00, 37.0412°N 22.2001°E, leg.
Pintér, E. & P. Subai 03.08.1976, HNHM 39189/4; on limestone rocks at border of Eleohóri
Village (= Janitsa), 545 m alt., UTM FG 00, 37.0408°N 22.1817°E, leg. Hadjicharalam-
bous & Subai 10.05.1991, S 15771/10; Taigetos Mountains, at the governmental bor-
der of Messinía to Lakonía, on marble rocks, 88 0 m alt., UTM FG 10, 37.0745°N
22.2955°E, leg. Pintér, E. & P. Subai 05.08.1976, HNHM 39169/2; leg. Neuteboom
31.8.1982, NNM/1; Kalamáta–Areópoli highway, chapel St. Nicholas Kabinari, UTM n.
det., leg. Hoenselaar 28.4.1996, Maa/1. — Lakonía, on limestone rocks at W border of
Geroliménas, 25 m alt., UTM FF 23, 36.4842°N 22.3993°E, leg. Butot & Subai
14.02.1981, S 17527/8 + 5 (juv.); leg. Kiss & Pintér 21.7.1986, HNHM 29904/1 + 1
(juv.); Geroliménas, road to Areópoli, UTM FF 23, 36.48°N 22.39°E, leg. Neuteboom
2.9.1982, NNM/2 + 1 (juv.); Geroliménas, road to Koúnos, UTM FF 23, 36.49°N 22.39°E,
leg. Neuteboom 2.9.1982, NNM/1 + 1 (juv.); W of Pírrihos (= between Kótronas and
Areópoli), UTM FF 25, 36.65°N 22.43°E, leg. Neuteboom 2.9.1982, NNM/2 (juv.); Pir-
gos (= S of Areópoli), on limestone rocks beside the "Diros" cave, 5–10 m alt., UTM FF
25, 36.6382°N 22.3792°E, leg. Subai 13.07.1979, S 15812/18; leg. Butot & Subai
14.2.1981, S 158 07/71 + 4 (juv.) + 1 (alk.); leg. Kiss & Pintér 21.7.1986, HNHM
29916/6 + 3 (juv.); 2 km W of Vahós, UTM FF 26, 36.68°N 22.40°E, leg. Kiss & Pintér
21.7.1986, HNHM 29899/3; on limestone rocks near Liméni (= N of Areópoli), 40 m
alt., UTM FF 26, 36.6787°N 22.3778°E, leg. Butot & Subai 16.02.1981, S 17529/21; S
border of Néo Ítilo, on limestone rocks, 30 m alt., UTM FF 26, 36.6877°N 22.3862°E,
leg. Hadjicharalambous & Subai 10.05.1991, S 15772/2; mountain slope 11 km from
Petrina direction Árna, on low limestone rocks, 760 m alt., UTM FF 28, 36.8785°N
22.4254°E, leg. Butot & Subai 15.02.1981, S 17528/3; between Ágios Dimítrios and
Kokála, UTM FF 33, 36.51°N 22.47°E, leg. Neuteboom 2.9.1982, NNM/3 + 2 (juv.);
mountain slope at border of Githio direction Spárti, on limestone rocks, 30 m alt., UTM
FF 36, 36.7594°N 22.5562°E, leg. Butot & Subai, 18.02.1981, S 17537/2; 8.5 km from
Githio direction Areópoli, in forest S of the highway, on limestone rocks, 150 m alt.,
UTM FF 36, 36.7398°N 22.4936°E, leg. Hadjicharalambous & Subai 10.05.1991, S

27
Revision of the genus Lindholmiol a Hesse, 1931
15880/3; gorge 10 km from harbor of Githio direction Areópoli, on limestone rocks, 40
m alt., UTM FF 36, 36.7299°N 22.5055°E, leg. Butot & Subai, 18.02.1981, S 17536/7 +
3 (juv.); gorge 15 km from port of Githio direction Areópoli, on limestone rocks, 100 m
alt., UTM FF 36, 36.7023°N 22.4725°E, leg. Butot & Subai leg.14.02.1981, S 17526/4;
gorge 12 km from Githio direction Areópoli, on limestone rocks and from a spring, 30
m alt., UTM FF 36, 36.7178°N 22.4998°E, leg. Subai 13.07.1979, S 15752/1; approx. 14
km from Githio direction Areópoli, on limestone rocks beside the highway, 100 m alt.,
UTM FF 36, 36.7017°N 22.4713°E, leg. Hadjicharalambous & Subai 10.05.1991, S
15810/1 + 1 (juv.); 1 km S of Goritsá, UTM FF 39, 37.01°N 22.56°E, leg. Kiss & Pintér,
19.8.1988, HNHM 44597/36 + 7 (juv.); Spárti–Geráki country road, 1 km E of Kefalās,
UTM FF 39, 37.02°N 22.53°E, leg. Hoenselaar 4.5.1996, Maa/1; Stefaniá, UTM FF 47,
36.84°N 22.60°E, leg. Kiss & Pintér 20.7.1986, HNHM 29914/2 + 2 ( juv.); 6.5 km from
Skála direction Githio, on limestone rocks beside a bridge and from the creek, UTM FF
47, 36.8048°N 22.6179°E, leg. Subai 14.07.1979, S 6578/4; 13 km from Skála direction
Githio, on limestone rocks, 50 m alt., UTM FF 47, 36.7959°N 22.5957°E, leg. Subai
14.07.1979, S 6660/2; 3 km from Skála direction Spárti, on limestone rocks beside
highway, 100 m alt., UTM FF 47, 36.8594°N 22.6 058°E, leg. Subai 14.07.1979, S
15747/76; 4 km SE of Agii Anárgiri, UTM FF 49, 37.00°N 22.66°E, leg. Kiss & Pintér,
19.8.1988, HNHM 44598/1; Ortsrand von Geráki in Richtung Spárti, unter Steinen,
UTM FF 59, leg. Butot & Subai, 18.02.1981, S 17538/1; 5 km from the crossing Geráki–
Spárti, Skála–Kosmás direction Kosmás, on limestone rocks, 800 m alt., UTM FF 59,
37.0607°N 22.7071°E, leg. Butot & Subai, 18.02.1981, S 17539/3 + 2 (juv.); 4 km N
from the bifurcation to Mólai, on limestone rocks 500 m E of the highway, 250 m alt.,
UTM FF 67, 36.8456°N 22.8448°E, leg. Subai 14.07.1979, S 15745/12; 4 km N of Mólai,
UTM FF 67, 36.84°N 22.83°E, leg. Neuteboom 1.9.1982, NNM/4 + 1 (juv.); Apídia, road
direction Spárti–Monemvassía, UTM FF 68 , 36.88°N 22.79°E, leg. Neuteboom
1.9.1982, NNM/2 (juv.); rock gorge 10.5 km from the bifurcation to Mólai direction
Skála, 250 m alt., UTM FF 68, 36.8635°N 22.8154°E, leg. Subai 14.07.1979, S 6802/1;
highway Skála–Mólai, on limestone rocks near bifurcation to Niáta, 230 m alt., UTM FF
68, 36.8683°N 22.8145°E, leg. Butot & Subai 17.02.1981, S 17534/11 + 2 (juv.); 700 m
N of Nómia (= road direction Kardenitsa), UTM FF 75, 36.65°N 23.01°E, leg. Neute-
boom 2.9.1982, NNM/1 + 2 (juv.); Elika, before junction to Arhángelos, UTM FF 75,
36.62°N 22.89°E, leg. Neuteboom 1.9.1982, NNM/5 + 2 (juv.); 9 km from Lira direction
Nómia, on limestone rocks, 85 m alt., UTM FF 75, 36.6521°N 23.0139°E, leg. Butot &
Subai 17.02.1981, S 17533/7 + 1 (juv.); 9 km from Monemvassía direction Skála, on
limestone rocks along border of road, 100 m alt., UTM FF 76, 36.7301°N 22.9882°E,
leg. Subai 14.07.1979, S 15746/8; Sikía–Monemvassía highway after junction to
Veliés, UTM FF 76, 36.72°N 22.97°E, leg. Neuteboom 1.9.1982, NNM/3 + 3 (juv.);
Veliés–Angelóna road, 3 km before Sikía, UTM FF 76, 36.74°N 22.95°E, leg. Neute-
boom 1.9.1982, NNM/4 + 1 (juv.); Talanda, before junction to Ágios Nikólaos, UTM FF
76, 36.67°N 22.94°E, leg. Neuteboom 1.9.1982, NNM/3; Ágios Apóstoli (= 16 km SE of
Pandanassa), 36.55°N 23.02°E, UTM FF 84, leg. Neuteboom 1.9.1982, NNM/7 + 3
(juv.); mountain near Dermatiánika (= Kabos, N of Neapolis), on limestone rocks and
on house ruins, 70 m alt., UTM FF 84, 36.5451°N 23.0533°E, leg. Butot & Subai
17.02.1981, S 15805/53 + 4 (juv.) + 12 (alk.); 1 km N of Monemvassía, UTM FF 86,
36.68°N 23.03°E, leg. Kiss & Pintér 21.7.1986, HNHM 29906/4; in the old town of Mon-

28 Peter Subai & Eike Neub ert
emvassía, on wall remains and under stones, 50 m alt., UTM FF 86, 36.688 8°N
23.0572°E, leg. Subai 14.07.1979, S 6579/11; leg. Butot & Subai 17.2.1981, S 17525/6
+ 1 (juv.) + 3 (alk.); leg. Neuteboom 1.9.1982, NNM/10 + 3 (juv.); leg. Kiss & Pintér
20.7.1986, HNHM 29893/50 + 4 (juv.); 2 km from Monemvassía direction Nómia, on
rocks at the sea shore beside the highway, UTM FF 86, 36.6710°N 23.0306°E, leg.
Butot & Subai 17.02.1981, S 17535/6 + 1 (alk.); leg. Kiss & Pintér 21.7.1986, HNHM
29892/28 + 14 ( juv.); well of Trípi Village, UTM FG 10, 37.09°N 22.35°E, leg. Neute-
boom 31.8.1982, NNM/1 (juv.); Mistra, byzantine ruin town W of Spárti, 410 m alt.,
UTM FG 20, 37.0739°N 22.3691°E, leg. Pintér, E. & P. Subai 05.08.1976, HNHM
39167/6, S 3797/7; leg. Neuteboom 3.9.1982, NNM/1 (juv.); leg. Kiss & Pintér
22.7.1986, HNHM 29910/1 + 1 (fragment); Spárti, UTM FG 20, 37.07°N 22.43°E, leg.
Pfeiffer, 1926, SMF 101849/1; 4 km from Spárti direction Hrísafa, UTM FG 20, 37.12°N
22.49°E, leg. Neuteboom 3.9.1982, NNM/2; 2 km S of Kástro, UTM n. det., leg. Kiss &
Pintér 9.7.1986, HNHM 29902/3. — Adelfí Island, NW part of the Island, 20 m alt., UTM
GJ 53, 39.11°N 23.97°E, leg. Uit de Weerd 5.6.1999, NNM/2. — Adelfópoulo Island, S
part of central area, 20 m alt., 39.12°N 23.98°E, UTM GJ 53, leg. Uit de Weerd 5 6.1999,
NNM/5. — Alónissos Island, Patitíri–Miléa road, 1.2–1.6 km NE of harbour Patitíri (=
approx. 2.5–2.8 km NE of Alónissos town), 210 m alt., 39.15°N 23.87°E, UTM GJ 43, leg.
Uit de Weerd 4.6.1999, NNM/1 + 3; 150 m from road Patitíri–Miléa towards Leftogiálo
(= 4.5 km NE of Alónissos town), 160 m alt., UTM GJ 44, 39.16°N 23.88°E, leg. Uit de
Weerd 4.6.1999, NNM/2; Ormos Tsortsi (= 4.2 km NE of Alónissos town), 4 m alt., UTM
GJ 53, 39.16°N 23.88°E, leg. Uit de Weerd 6.6.1999, NNM/2; Patitíri–Gerakis road, 2.1
km NE from turning point to Lef tos Giakos (= 6.5 km NE of Alónissos town), UTM GJ 54,
39.17°N 23.90°E, leg. Uit de Weerd 6.6.1999, NNM/1. — Gioúra Island, UTM KD 56,
39.39°N 24.16°E, leg. Storch 7.7.1978, SMF 284930/1; Gioúra Island, E coast, UTM KD
56, 39.38°N 24.18°E, leg. Storch 6.7.1978, SMF 284929/1. — Kirá Panagiá Island, UTM
KD 45, 39.32°N 24.07°E, leg. Storch 1.7.1978, SMF 284928/3. — Pipéri Island, UTM KD
65, 39.35°N 24.32°E, leg. Storch 4.7.1978, SMF 284927/1. — Skándzoura Island, UTM
KD 52, 39.08°N 24.11°E, leg. Storch 9.7.1978, SMF 284925/9; Skándzoura monastery,
UTM KD 52, 39.08°N 24.11°E, leg. Storch 10.7.1978, SMF 284926/3. — Skíathos Island,
UTM n. det., SMF 284945/2; Skíathos town, "Odos Filokleus Georgiadou" 200 m NE of
harbour, 20 m alt., UTM GJ 13, 39.16°N 23.49°E, leg. Gittenberger & Uit de Weerd
29.5.1999, NNM/3; Evangelistrías monastery (= 4 km NNW of Skíathos town), 80 m
alt., UTM GJ 14, 39.1904°N 23.4806°E, leg. Gittenberger & Uit de Weerd 30.5.1999,
NNM/2 + 1 (juv.); 400 m N of Evangelistrías monastery, 150 m alt., UTM GJ 14, 39.19°N
23.48°E, leg. Gittenberger & Uit de Weerd 30.5.1999, NNM/4; 1 km N of Evangelistrías
monastery, 230 m alt., 39.19°N 23.48°E, UTM GJ 14, leg. Gittenberger & Uit de Weerd
29.5.1999, NNM/4. — Skópelos Island, UTM n. det., leg. Banzhaf, 1929, SMF 284876/1
(juv.); Kafasi, 1 km ESE of Metamórfosis monastery (= 3 km ESE of Skópelos town),
200 m N of junction, 300 m alt., UTM GJ 33, 39.11°N 23.75°E, leg. Uit de Weerd
3.6.1999, NNM/1 (juv.); Koprisies Mountain, 400–1300 m from peak (along the road),
320–370 m alt., UTM GJ 33, GPS n. det., leg. Uit de Weerd 1.6.1999, NNM/1 + 2 + 24;
500 m NNE of church Ágios Konstantínos (= 1.2 km NW of Skópelos town), S slope,
UTM GJ 33, 39.13°N 23.71°E, leg. Uit de Weerd 2.6.1999, NNM/1. — Évvia Island (=
Euböa), N border of Ágios Geórgios, in valley, on dry limestone rocks, 50 m alt., UTM
FJ 60, 38.8410°N 22.8920°E, leg. Subai & Szekeres 10.05.1997, S 15687/2; spruce for-

29
Revision of the genus Lindholmiol a Hesse, 1931
est 5 km from Gregolimano direction Ágios Geórgios, approx. 200 m N of the highway,
on dry limestone rocks, 50 m alt., UTM FJ 60, 38.8401°N 22.9054°E, leg. Subai & Szek-
eres 10.05.1997, S 15699/14 + 3 (juv.) + 1 (alk.); Loutrá Edipsoú, near church, UTM FJ
70, 38.85°N 23.04°E, leg. Neuteboom 25.10.1984, NNM/1 (juv.); Límni, UTM GH 09,
38.76°N 23.32°E, leg. Krüper, 19 03, SMF 101970/3; Mantoúdi, UTM GH, 19, 38.79°N
23.48°E, leg. Ottermann, 1912, SMF 284860/22; S border of Mantoúdi, on limestone
rocks at river side, 20 m alt., UTM GH, 19, 38.7942°N 23.4770°E, leg. Subai 01.05.1991,
S 15708/2; leg. Gittenberger & Uit de Weerd 10.6.1999, NNM/2; Halkída (= Chalcis),
UTM GH 26, 38.46°N 23.58°E, SMF 284943/3, 284944/4, 285166/2; leg. Fitz-Gerard,
1881, SMF 101986/3; Kiparíssi (= 7 km N of Psahná), UTM GH 27, 38.62°N 23.59°E,
leg. Neuteboom 23.10.1984, NNM/1 + 4 (juv.); 1.5 km S of Néa Pagóntas, UTM GH 28,
38.65°N 23.56°E, leg. Neuteboom 24.10.1984, NNM/1; approx. 1–2 km N of Néa Artáki,
approx. 200 m E of the highway, on low limestone rocks, UTM GH 26, 38.5404°N
23.6344°E, leg. Riedel & Subai 09.04.1988, S 15803/4 + 2 (alk.); Dírfis Mountains (=
Monte Delphi), UTM GH 47, 38.62°N 23.84°E, SMF 101879/1, 284921/5, 284922/4,
284923/3, 285764/8; Stení, UTM GH 47, 38.58°N 23.83°E, SMF 101856/4, 101871/15,
284924/3; above Áno Steni, on limestone rocks, 520 m alt., 520 m alt., UTM GH 47,
38.5847°N 23.8349°E, leg. Subai & Szekeres 10.05.1997, S 15917/1 + 3 (juv.); Dírfi
Mountains, N of the highway Steni Dirfios–Strópones, 200 m NE of the E.O.S. refuge
hut, on limestone rocks, 1150 m alt., UTM GH 47, 38.6142°N 23.8597°E, leg. Subai &
Szekeres 10.05.1997, S 15918/1; 9 km from Stení Dírfios towards Strópones, UTM GH
47, 38.60°N 23.87°E, leg. Neuteboom 23.10.1984, NNM/2 + 3 (juv.); 3 km from
Strópones towards Stení Dírfios, UTM GH 57, 38.60°N 23.88°E, leg. Neuteboom
23.10.1984, NNM/1 (juv.); small rock gorge at W border of Strópones (= Dírfis Moun-
tains), on limestone rocks, 360 m alt., UTM GH 57, 38.6205°N 23.8853°E, leg. Subai
01.05.1991, S 15709/6; S of Vassiliká, UTM GJ 01, 38.96°N 23.35°E, leg. Neuteboom
24.10.1984, NNM/2 + 1 (juv.); 1.7 km from Elliniká direction Vassiliká, on limestone
rocks beside the quarry, 35 m alt., UTM GJ 01, 38.9979°N 23.3488°E, leg. Subai
30.04.1991, S 15707/2; on rocks E of Elliniká, UTM GJ 02, 39.00°N 23.34°E, leg. Neute-
boom 24.10.1984, NNM/1; W border of Alivéri, on limestone rocks and in rock debris,
350 m alt., UTM KC 45, 38.4022°N 24.0138°E, leg. Subai 16.04.1986, S 15883/35 + 1
(alk.); leg. Neuteboom 22.10.1984, NNM/1; S of Kriezá, UTM KC 45, 38.38°N 24.14°E,
leg. Neuteboom 21.10.1984, NNM/1; SE border of Koutoumoulas, under stones, 125
m alt., UTM KC 45, 38.3787°N 24.1105°E, leg. Subai 16.04.1986, S 15802/2; cave nich-
es W of Vrissi, foot of Mavrovoúni Mountains, on limestone rocks, 450 m alt., UTM KC
46, 38.5549°N 24.0268°E, leg. Subai 16.04.1986, S 15779/2; road between Orológi
and Monódrio, UTM KC 46, 38.53°N 24.08°E, N leg. euteboom 22.10.1984, NNM/2; 1
km S of Kaliáni, UTM n. det., leg. Neuteboom 22.10.1984, NNM/2; Konistraes, UTM KC
47, 38.54°N 24.06°E, SMF 101899/4, 284957/4, 284958/2, 284959/2; Kími, UTM KC
48, 38.63°N 24.10°E,SMF 101873/2; 8.1910, SMF 285164/4, 285165/40; leg. Holtz
4.1926, SMF 207818/7; between Néa Stíra and Stíra, UTM KC 52, 38.16°N 24.22°E, leg.
Pintér 22.8.1976, HNHM 39166/14 + 3 (juv.); Kapsála (= 3 km of Stíra), UTM KC 52,
38.14°N 24.24°E, leg. Neuteboom 21.10.1984, NNM/5; "Malithianon", "Malithianu",
"Malithiedin", UTM n. det., leg. Krüper, SMF 101855/13, 284152/9, 285162/2; leg.
Krüper, 1890, SMF 284940/4, 284949/7; Fleva, UTM n. det., leg. Krüper 7.1928, SMF
101962/5; "Achmet Aga", UTM n. det., leg. Godet, SMF 284942/4; between Lépoura

30 Peter Subai & Eike Neub ert
and Ag. Nikolaou monastery, UTM n. det., leg. Krüper, SMF 101868/20. — Kea Island
(= Keos), approx. 1 km E of Otziás, on limestone rocks on top of a slate mountain,
180–200 m alt., UTM KB 67, 37.6774°N 24.3625°E, leg. Subai 06.05.1991, S 15700/34.
— Páros Island, UTM n. det., leg. Krüper, 1883, SMF 101864/4. — Lipsí Island (= Lipsos,
S of Sámos), surrounding Lipsí [Village], UTM MB 72, 37.29°N 26.77°E, leg. Becker &
Wedekind 9.1984, S 15741/1. — Sámos Island, UTM n. det., SMF 101981/3; leg. Par-
avicini, 1933, SMF 284912/2; between Marathókambos and Ágios Theódori, UTM MB
77, 37.73°N 26.71°E, leg. Neuteboom 8.5.1978, NNM/1 (juv.); Balos (= coast above
Kouméika), UTM MB 77, 37.69°N 26.74°E, leg. Neuteboom 10.5.1978, NNM/1; Ireon,
Hera Temple, UTM MB 86, 37.66°N 26.87°E, leg. Neuteboom 7.5.1978, NNM/1 (juv.); 4
km from Spatharéi direction Pagóndas, UTM MB 86, 37.66°N 26.79°E, leg. Neuteboom
10.5.1978, NNM/6; leg. Maassen 5.1993, Maa/7; 2 km N of Mavratzéi, UTM MB 87,
37.73°N 26.66°E, leg. Maassen 5.1993, Maa/1; Mavratzéi, UTM MB 87, 37.71°N
26.86°E, leg. Maassen 5.1993, Maa/13; 2 km SE of Míli, UTM MB 87, 37.67°N 26.87°E,
leg. Maassen 5.1993, Maa/8; 1.5 km SE of Stravrinídes, UTM MB 88, 37.78°N 26.82°E,
leg. Maassen 5.1993, Maa/1; 3 km E of Avlákia (= W of Kokkári), UTM MB 88, 37.79°N
26.86°E, leg. Maassen 5.1993, Maa/2; river valley 0.5 km from Platanakia direction
Manolates, UTM MB 88, GPS n. det., leg. Maassen 5.1993, Maa/9; 0.5 km W of Vourli-
otes, along path, UTM MB 88, 37.78°N 26.83°E, leg. Maassen 5.1993, Maa/3; wells N
of Vourliotes, UTM MB 88, 37.78°N 26.84°E, leg. Maassen 5.1993, Maa/2; Sámos,
UTM MB 97, 37.76°N 26.97°E, leg. Neuteboom 5.5.1978, NNM/11 (alk.); Pithagório,
near Hotel "Dorissa Bay", under stones, UTM MB 97, 37.69°N 26.92°E, leg. Neute-
boom 7.5.1978, NNM/9 + 1 (juv.); between Pithagório and Vathí, before pass, under
stones, UTM MB 97, 37.72°N 26.96°E, leg. Neuteboom 7.5.1978, NNM/9 + 3 (juv.);
Vathí (= between Sámos and Pithagório), UTM MB 97, 37.74°N 26.98°E, leg. Neute-
boom 7–9.5.1978, NNM/6 + 3 (juv.); Thios Mountain, 1 km NE of Sámos, 200 m alt.,
UTM MB 97, 37.75°N 26.98°E, leg. Maassen 5.1993, Maa/1; Platanaki (= 1.5 km NW of
Sámos), UTM MB 97, GPS n. det., leg. Maassen 5.1993, Maa/15, S 15937/3; 3 km S of
Kokkári (= direction Mitilinií ), 200 m alt., UTM MB 97, 37.75°N 26.90°E, leg. Maassen
5.1993, Maa/6; 1 km N of Mitilinií (= direction Kokkári), UTM MB 97, 37.74°N 26.89°E,
leg. Maassen 5.1993, Maa/2; 0.5 km from Pithagório direction Spilianis, UTM MB 97,
37.69°N 26.93°E, leg. Maassen 5.1993, Maa/3; above Spilianis monastery (= N of Pith-
agório), under stones, UTM MB 97, 37.69°N 26.93°E, leg. Maassen 5.1993, Maa/12 + 1
(juv.) S 15938/3; 1 km W of Kokkári, UTM MB 98, 37.78°N 26.86°E, leg. Maassen
5.1993, Maa/1; 0.5 km S of Kokkári (= direction Mitilinií), 50 m alt., UTM MB 98,
37.77°N 26.89°E, leg. Maassen 5.1993, Maa/3; 1 km S of Kokkári (= direction Mitilinií),
150 m alt., UTM MB 98, 37.76°N 26.89 °E, leg. Maassen 5.1993, Maa/1; Kótsikas mon-
astery (= NE of Sámos), UTM MB 98, 37.78°N 26.96°E, leg. Maassen 5.1993, Maa/4;
Ágios Paraskeví (= NE of Sámos), along beach, UTM MB 98, 37.78°N 26.99°E, leg.
Maassen 5.1993, Maa/1 + 1 (juv.); 0.5 km S of Ágios Paraskeví, Thios Mountain, 150
m alt., 37.77°N 26.99°E, UTM MB 98, leg. Maassen 5.1993, Maa/3; 1 km S of Ágios
Paraskeví, Thios Mountain, 300 m alt., UTM MB 98, 37.76°N 26.99°E, leg. Maassen
5.1993, Maa/2; 3 km W of Ágios Paraskeví, UTM MB 98, 37.78°N 26.95°E, leg. Maas-
sen 5.1993, Maa/16; Thios Mountain (= NE of Sámos), Profítis Ilías peak, 430 m alt.,
37.76°N 26.99°E, UTM MB 98, leg. Maassen 5.1993, Maa/3; 0.5 km N of Profítis Ilías
peak, 350 m alt., 37.77°N 26.99°E, UTM MB 98, leg. Maassen 5.1993, Maa/27 + 1

31
Revision of the genus Lindholmiol a Hesse, 1931
(juv.) S 15939/3; W border of Kokkári, UTM MB 98, 37.78°N 26.87°E, leg. Neuteboom
8.5.1978, NNM/1; 1 km W of Agia Zóni and walls of monastery, UTM NB 07, 37.75°N
26.98°E, leg. Neuteboom 9.5.1978, NNM/13 + 37 (alk.); approx. 2–3 km E of Sámos,
on rocks along sea shore, UTM NB 07, 37.76°N 27.02°E, leg. Maassen 5.1993, Maa/13;
coast 1 km W of Psilí Ámmos (= E of Pithagório), UTM NB 07, 37.70°N 26.99°E, leg.
Maassen 5.1993, Maa/12; 1.5 km N of Psilí Ámmos, along path, UTM NB 07, 37.72°N
27.01°E, leg. Maassen 5.1993, Maa/9. — Kálimnos Island (= Calino), UTM n. det., SMF
284915/18, 284916/2; Kálimnos town (= Pothia), UTM MA 98, 36.95°N 26.98°E, leg.
Neuteboom 24.10.1979, NNM/10; gorge behind harbor of Kálimnos, UTM MA 98,
36.95°N 26.99°E, 1934, SMF 101877/6; Kástro Hrisohórias, UTM MA 98, 36.96°N
26.96°E, leg. Neuteboom 24.10.1979, NNM/6 + 1 (juv.) + 15 (alk.); behind harbor of
Vathis (= Vati), UTM NA 09, 36.97°N 27.02°E, leg. K. L. Pfeiffer, 1934, SMF 101976/3.
— Psérimos Island, UTM NA, 18, SMF 284914/2; leg. K. L. Pfeiffer, 1934, SMF
207817/2; leg. K. L. Pfeif fer, 1940, SMF 285175/4; leg. Neuteboom 20.10.1979,
NNM/18; on rocks near harbour, UTM NA, 18, 36.93°N 27.13°E, leg. K. L. Pfeiffer, 1934,
SMF 101977/30. — Kós Island, UTM n. det., SMF 207812/2. — Sphanteria Island near
Tilos Island, UTM n. det., leg. Philippson 2.4.1887, SMF 284891/1.
Turkey, Çanakkale County, Gökçeada Island, UTM n. det., leg. Kleefisch 10.9.1985,
(ex. coll. Schütt Nr. 1180) S 20789/1; Çanakkale, Kilidürbahir, UTM ME 44, 40.17°N
26.37°E, leg. Schütt 28.4.2002, (ex. coll. Schütt Nr. 1751) S 20790/29. — Manisa
County, Cornana, UTM n. det., HNHM 39174/2; Sochia, UTM n. det., SMF 101865/4;
Manisa (= ancient Magnesia), UTM NC 37, 38.60°N 27.37°E, SMF 284909/2. — Aydın
County, Dılek Yarımadası Mıllı Parkı, Kanyon 7 km direction Güzelçamlı, UTM NB
17, 37.68°N 27.16°E, leg. Schütt 23.4.2000, (ex. coll. Schütt Nr. 1686 ) S 20793/15 +
2 (juv.); ruins of Milet (= Miletos), UTM NB 25, 37.52°N 27.27°E, leg. Puts, 1983,
Maa/1; Samsun Mountains, Priene near Güllübahçe, above ancient ruins, UTM NB
26, 37.65°N 27.29°E, leg. Schlickum, 19.9.1968, SMF 279324/3; leg. Neuteboom
24.4.1989, NNM/1 + 1 (juv.); leg. Erőss 28.7.1992, coll Erőss /12 + 2 (juv.); leg. Schütt
24.4.200 0, (ex coll. Schütt Nr. 1687) S 20794/22; Kuşadası, UTM NB 29, 37.85°N
27.26°E, leg. Neuteboom 6.5.1978, NNM/2 ( juv.). — Izmir County, Efes (= Ephesos),
among ruins, UTM NB 29, 37.93°N 27.34°E, leg. Kaysser, 1904, SMF 284902/1; leg.
Schlickum 15.9.1968, SMF 279323/1; leg. Papp 10.10.1972, HNHM 33704/1; leg. Seidl
7.5.1975, HNHM 39186/2, 49552/2; leg. Neuteboom 6.5.1978, NNM/11 + 3 ( juv.); leg.
Erőss 30.7.1992, E&F/31; leg. Bánki 26.7.1995, HNHM 41708/3 + 3 (juv.); surround-
ings hous of death of Saint Maria on Ala Dag Mountain (= 5 km S of Efes), UTM NB
29, 37.91°N 27.33°E, leg. Seidl 7.5.1975, Maa/5; Pamucak, Orman Kampi, UTM NB
29, 37.93°N 27.28°E, leg. Bilgin, (ex. coll. Schütt) S 20791/1; surroundings of Izmir (=
Smyrna), UTM n. det., leg. Basoglu 2.12.1956, SMF 166916/4; Izmir–Bornova, UTM NC
15, 38.47°N 27.24°E, SMF 101956/5; halfway between Izmir and Manısa, UTM NC 26,
38.54°N 27.30°E, leg. Schütt 4.9.2004, (ex coll. Schütt Nr., 1842) S 20792/1.
Remarks: Lindholmiola lens is very variable in shell shape and size, a fact
which caused authors to describe several varietal taxa, which here are all
synonymised under L. lens. The nominotypical subspecies is known from the
Pelopónnisos and the Ionian islands of Kefallinia and Zakinthos and is charac-

32 Peter Subai & Eike Neub ert
terized by a blunt keel, a narrow and only slightly perspective umbilicus, the
moderately enlarged peristome, the medium shell size and narrow whorls.
The mountains in the north of the Pelopónnisos are inhabited by a form which
differs by shells with slightly broader whorls, an even more blunt keel, and
a larger perspective umbilicus (= elia). However, this form also lives on the
Island Évvia and close to Neohóri (= NE of Aridéa) in Makedonía. This form
is intermediate between the nominotypical subspecies and another, quite
remarkable form (= callojuncta). This very distinct form has a shell with a
sharp keel, broad whorls, a wide perspective umbilicus, and a surface sculp-
ture of small riblets. Its shells are of medium size (sometimes scalarid shells
occur), and the peristomial insertions are connected by a more conspicuous
parietal callus. The peristome of this form is wider than in any other form of
L. lens. The form callojuncta lives around the Bay of Korinthos, but also in
Argolída and in Arkadía.
Fig. 15. Distribution of Lindholmiola lens and L. spectabilis.

33
Revision of the genus Lindholmiol a Hesse, 1931
The eastern part of continental Greece, Atikí, the Island Évvia, Thessalía
and the Halkidikí Peninsula is inhabited by a small to medium-sized form (=
lentiformis). It can be characterised by its moderately broad whorls, the blunt
keel, and the narrow to slightly perspective umbilicus. A similar but unnamed
form inhabits the Dodecanese Islands and the western coast of Turkey. It has
a somewhat sharper keel and a wider umbilicus if compared to lentiformis.
The form piligera is intermediate between the forms elia and lentiformis. It
lives in Viotía, Fthiótida and on the Island Évvia. It differs by a more tightly
coiled shell, the whorls which are more rounded (surface and subsurface),
and the only slightly perspective umbilicus.
Distribution (Fig. 15): Lindholmiola lens inhabits quite a large area. It is
known from the Ionian Islands of Kefallinia and Zakinthos, the Pelopónnisos,
the islands of the Aegean Sea to the western part of Turkey. In the north, it
is recorded from the Bay of Korinthos, and the eastern part of continental
Greece to the western part of the Halkidikí Peninsula.
Lindholmiola barbata (A. Férussac, 1821) Figs 1, 16–18, 19, 45
1821 Helix barbata A. Férussac, Tableaux systématiques des animaux mol-
lusques classés en familles naturelles: 41 (Folio) or 37 (Quarto), Nr. 152
[partim].
1832 Helix barbata α, – A. Férussac, (in A. Férussac & Deshayes), Histoire natu-
relle générale et particulière des mollusques terrestres et fluviatiles (Atlas),
22–27: T. 66, F. 3 [Lectotype, design. Gittenberger & Groh, 1986] + Expl. pl.: ij.
1837 Helix (Caracollina) barbata, – Beck, Index molluscorum praesentis aevi
musei principis augustissimi Christiani Frederici: 28.
1848 Helix corcyrensis ?, – L. Pfeiffer, Monographia heliceorum viventium, 1:
415, Nr. 1080 [partim, nec Syn. pro H. barbata Férussac; non H. corcyrensis
Rossmaessler, 1838].
1859 Helix barbata, – Mousson, Vierteljahrsschrift der Naturforschenden Gesell-
schaft in Zürich, 4: 258.
1859 Helix corcyrensis, – L. Pfeiffer, Monographia heliceorum viventium, 4: 312,
Nr. 2004 [partim, nec Syn. pro H. barbata α Férussac, 1832; non H. corcy-
rensis Rossmaessler, 1838].
1868 Helix barbata, – L. Pfeiffer, Monographia heliceorum viventium, 5: 415.
1876 Helix barbata, – L. Pfeiffer, Monographia heliceorum viventium, 7: 467.
1879 Helix (Trigonostoma) lens var. barbata, – Westerlund & Blanc, Aperçu sur
la faune malacologique de la Grèce inclus l’Epire et la Thessalie: 35.
1887 Helix (Caracolina) barbata, – Tryon, Manual of Conchology, 3: 118.

34 Peter Subai & Eike Neub ert
1889 Helix (Gonostoma) lens var. barbata, – Martens, Archiv für Naturgeschichte,
55 (1): 186, 229.
1889 Helix (Caracolina) barbata, – Westerlund, Fauna der in der paläarctischen
Region lebenden Binnenconchylien, 2: 20 [partim].
1894 Helix barbata, – O. Boettger, Nachrichtsblatt der Deutschen Malakozoo-
logischen Gesellschaft, 26: 2.
1894 Helix (Caracollina) barbata, – Pilsbry, Manual of Conchology, 9: 228.
1904 Helix (Gonostoma) barbata, – Sturany, Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 36: 108.
1918 Caracollina barbata, – Hesse, Nachrichtsblatt der Deutschen Malakozoo-
logischen Gesellschaft, 50: 109.
1978 Helix (Gonostoma) barbata, – Seidl, Mitteilungen der Zoologischen Gesell-
schaft Braunau, 3 (5/7): 160.
1986 Lindholmiola barbata, – Gittenberger & Groh, Archiv für Molluskenkunde,
116 (4/6): 222, Abb. 1 b (= 3).
1990 Lindholmiola barbata, – Fechter & Falkner, Steinbachs Naturführer: 222.
1991 Lindholmiola barbata, – Maassen, De Kreukel, 27 (1/2): 9, Taf. 1.
Diagnosis: depressed, brown to yellowish shell; covered with yellowish hairs;
last whorl with a blunt peripheral keel; umbilicus cylindrical, partly covered.
Description of shell: shell depressed, basic colour brown to yellowish; pro-
toconch 1–1¼ smooth whorls; initial teleoconch whorls with a very fine radial
sculpture, riblets slightly wavy (under strong magnification), inconspicuous
on the last whorl; teleoconch surface (except the initial whorls) covered by
fine granules and with small hair knuckles; hairs yellowish, 0.3–0.8 mm long
(shorter in the umbilicus), permanently fixed to the shell; 5¼–6¾ regularly
increasing whorls, depressed above, rounded below, with a blunt peripheral
keel; suture of medium depth; umbilicus almost cylindrical, slightly increasing
in the last whorl, diameter 1.25–2 mm, partly covered by the columellar reflec-
tion of the peristome; aperture oblique, slightly curved in lateral view; in fron-
tal view subquadrate, peristomial insertions with a gap of 2.8–5.8 mm; pari-
etal callus lacking; peristome sharp, laterally broadened and slightly reflected
basally; columellar reflection of peristome broad, covering ¼ of the umbilicus
(in adult shells).
Measurements: H: 4.2–6.25; D: 9.7–14.4; aH: 3.75–5.8 (2–3.6); aW: 5.1–8.3.
Details of body (n = 5, Crete, Bay of Soúda (= E of Haniá)): head and dor-
sum grey to greyish brownish, flank and foot sole laterally grey to beige; man-
tle collar irregularly greyish pigmented; parallel to the ureter, small elongated
greyish spots present (sometimes fused to a stripe); secondary ureter opens
0.5 mm from the respiratory pore.

35
Revision of the genus Lindholmiol a Hesse, 1931
Morphology of the genital organs: penis short with a slightly broadened
distal part; glandular tissue covering almost half of the penial tube; proxi-
mal penial lumen with broad and shallow pilasters; distal penial lumen with
two raised, parallel pilasters; several shorter longitudinal folds parallel to
or between the major distal pilasters; vagina of moderate length; glandula
reaches 2/3 of the length of the bursa copulatrix, with a short stem and a
strong sculpture of crossing furrows; pedunculus of bursa copulatrix and
the slightly broadened vesicle of same length; interior of the terminal genital
organs with four to five well pronounced pilasters starting at the genital pore
and fusing to a thickened area in the centre of the atrium; this area connects
to a single long pilaster stretching into the vaginal lumen.
Figs 16–18. Lindholmiola barbata. Fig. 16: NMBE 515636, Island Kríti, Nomos Iráklio, valley E from
the airport Iráklio (small rounded form), D = 9.66 mm; Fig. 17: NMBE 515635, Island Kríti, Nomos
Haniá, canyon ca. 4 Km N of Thériso (large flat form), D = 13.78 mm; Fig. 18 : NMBE 515581, Rethim-
non, Zousidi, 09.04.1993, leg. N. Chalwatzis, D = 12.49 mm. — All photos Neubert/Bochud, × 3.

36 Peter Subai & Eike Neub ert
Differential diagnosis: Lindholmiola corcyrensis is conchologically quite
close, but differs by its more tightly coiled whorls, its coarser growth ribs on
the surface, its deeper suture, and the umbilicus, which is only slightly cov-
ered by the columellar callus. Its aperture is more strongly bent in lateral view,
and the peristome is broader. In L. corcyrensis, the penial lumen is filled by
v-shaped, obliquely arranged short folds.
Forms of L. lens with rounded whorls are also similar to L. barbata. How-
ever, shells of L. lens of the same size differ by having an additional whorl,
coarser growth ribs, a deeper suture, and an almost completely open umbili-
cus. Their aperture is more bent in frontal view. In L. lens, the short folds in
the proximal penial lumen are usually missing, and the pilasters in the distal
lumen are weaker.
Specimens examined (all Greece, Island of Crete):
Nomos Haniá: Plátanos, UTM GE 32, leg. Butot & Subai 10.2.1981, S 17518/8+1(juv.);
leg. Kiss & Pintér 13.8.1988, HNHM 44682/1(juv.); Falásarna, N of the excavations, UTM
GE 33, leg. Neuteboom 5.4.1984, NNM/2+2(juv.); coast 2.5 km S of Falásarna, UTM
GE 33, leg. Butot & Subai 10.2.1981, S 17519/6; Kotsoumatádos (= S of Topólia) UTM
GE 42, leg. Maassen 4.1987, Maa/1; N boundary of Kakópetros, UTM GE 42, leg. Drim-
mer, Pintér & Varga 25.7.1994, HNHM 43541/1+1; Rodopós (= SW of Afráta) UTM GE 43,
leg. Drimmer, Pintér & Varga 24.7.1994, HNHM 43540/1+2(juv., Bruchst.); Giflos river
drift close to Kaloudianá and under stones, UTM GE 43, leg. Butot & Subai 10.2.1981, S
14099/1+6(juv.); 1 km N of Soúgia, UTM GE 50, leg. Drimmer, Pintér & Varga 25.7.1994,
HNHM 43542/1(juv.); 1 km N of Prasés, UTM GE 51, leg. Kiss & Pintér 14.8.1988, HNHM
44683/3+1(juv.); 1 km S of Tavronítis (= S of Kolimvári) UTM GE 53, leg. Maassen 4.1987,
Maa/3; N boundary of Kolimvári, UTM GE 53, leg. Maassen 4.1987, Maa/6; 1 km N of
Kolimvári, UTM GE 53, leg. Drimmer, Pintér & Varga 24.7.1994, HNHM 43539/1+1(juv.,
Bruchst.); 2 km N of Kolimvári, UTM GE 53, leg. Kiss & Pintér 13.8.1988, HNHM 44681/1;
between Kolimvári and Afráta, 700 m on a path to W, UTM GE 53, leg. Butot & Subai
11.2.1981, S 17516/3 (juv.); 1 km N of Afráta, UTM GE 54, leg. Pintér & Varga 25.7.1992,
HNHM 45252/1 (juv.); N boundary of Omalós, UTM GE 61, leg. Drimmer, Pintér & Varga
25.7.1994, HNHM 43543/1; 3 km S of Chiliaró, UTM GE 62, leg. Drimmer, Pintér & Varga
25.7.1994, HNHM 43544/1 (juv.); Mesklá, at the bridge at the affluent of two creeks,
UTM GE 62, leg. Butot & Subai 8.2.1981, S 17522/3+2(juv.)+1(juv. alk.); Mesklá, at both
churches, UTM GE 62, leg. Butot & Subai 8.2.1981, S 17521/4+3 (juv.)+2(alk.); Thériso,
old church, UTM GE 72, leg. Butot & Subai 11.2.1981, S 17515/1; 1 km N of Thériso, on
limestone rocks, UTM GE 72, leg. Maassen 10.1992, Maa/2; 2 km N of Thériso, on lime-
stone rocks, UTM GE 72, leg. Maassen 10.1992, Maa/2; canyon ca4 km N of Thériso, on
limestone rocks, UTM GE 72, leg. Butot & Subai 11.2.1981, S 14100/69+12 (juv.)+1 (alk.);
leg. Maassen 10.1992, Maa/4; canyon 5.7 km of Thériso in direction to Haniá, UTM GE 72,
leg. Butot & Subai 11.2.1981, S 17514/10; canyon 7.6 km of Thériso in direction to Haniá,
UTM GE 72, leg. Butot & Subai 11.2.1981, S 17517/5+1 (juv.); 7 km E of Anópoli, UTM
KU 39, leg. Maassen 4.1990, Maa/1; canyon 1.2 km of Sfakia in direction to Anópoli, on
limestone rocks, UTM KU 39, leg. Butot & Subai 9.2.1981, S 14102/3+5 (juv.); Haniá (=

37
Revision of the genus Lindholmiol a Hesse, 1931
Chania, Kanea) UTM KV 23, SMF 101959/1, 207801/1; Haniá, on the walls of the castle,
UTM KV 23, leg. Butot & Subai 8.2.1981, S 17523/14+2 (juv.)+27 (alk.); canyon "Farangi
N., Venizélou" UTM KV 23, leg. Drimmer, Pintér & Varga 27.7.1994, HNHM 43548/10+6
(juv.), 43549/2+2 (juv.), 43550/30 (adult+juv.); boundary of Anópoli, in direction to Sfa-
kia, UTM KV 30, leg. Butot & Subai 9.2.1981, S 17520/1; Chorafia (= SE of Haniá) UTM
KV 32, leg. Drimmer, Pintér & Varga 28.7.1994, HNHM 43551/2+1 (juv.); Haniá–Stérnes
road, at the crossing to Soúda, UTM KV 33, leg. Butot & Subai 12.2.1981, S 17512/3+6
(alk.); peninsula Akrotíri, Aróni, UTM KV 33, leg. Kiss & Pintér 16.7.1986, HNHM 29879/6;
peninsula Akrotíri, Kalavás, UTM KV 33, leg. Drimmer, Pintér & Varga 27.7.1994, HNHM
43547/2; peninsula Akrotíri, 3 km NW from the airport, UTM KV 33, leg. Drimmer, Pintér
& Varga 27.7.1994, HNHM 43545/2+1 (juv.); Bay of Soúda, 100–150 m of the rivermouth,
S from the English cemetery, UTM KV 33, leg. Butot & Subai 12.2.1981, S 14104/6+3
(juv.)+61(alk.)+3 (juv. alk.); Asígonia (= Gonia), UTM KV 40, leg. Drimmer, Pintér & Varga
20.7.1994, HNHM 43524/1; 2 km S of Vrises (= in direction to Sfakia) UTM KV 41, leg.
Maassen 4.1990, Maa/1; 10.4 km of Askýfou in direction to Vrises, on limestone rocks
UTM KV 41, leg. Butot & Subai 9.2.1981, S 14101/19+6 (juv.); between Kalamítsi and
Amigdáli, UTM KV 41, leg. Drimmer, Pintér & Varga 23.7.1994, HNHM 43534/1+1; 2 km W
of Georgioúpoli, UTM KV 41, leg. Maassen 10.1992, Maa/8; 3 km W of Georgioúpoli, UTM
KV 41, leg. Maassen 4.1990, Maa/23; 2 km E of Neo Horió, UTM KV 42, leg. Maassen
4.1990, Maa/10 ; at the N boundary of Kefalás, UTM KV 42, leg. Butot & Subai 12.2.1981,
S 17513/4; leg. Drimmer, Pintér & Varga 23.7.1994, HNHM 43536/1; hill slope SE of Kalá-
mi, UTM KV 42, leg. Drimmer, Pintér & Varga 23.7.1994, HNHM 43538/1+1 (juv.); Xirostér-
ni, UTM KV 42, leg. Drimmer, Pintér & Varga 23.7.1994, HNHM 43535/1; N boundary of
Drápanos, UTM KV 42, leg. Drimmer, Pintér & Varga 23.7.1994, HNHM 43537/3; penin-
sula Akrotíri, 2 km N of Agia Triáda, UTM KV 42, leg. Drimmer, Pintér & Varga 27.7.1994,
HNHM 43546/1; peninsula Akrotíri, 1 km N of Moni Gouvernétou, UTM KV 44, leg. Maas-
sen 4.1987, Maa/2; between Georgioúpoli and Exópoli, UTM KV 51, leg. Drimmer, Pintér
& Varga 23.7.1994, HNHM 43533/5+7 ( juv.); W boundary of Geourgioúpoli, UTM KV 51,
leg. Kiss & Pintér 17.7.1986, HNHM 29876/26+18 (juv.); rock face along the motorway
close to Georgioúpoli, UTM KV 51, leg. Maassen 4.1987, Maa/26; NW of Kournás, UTM KV
51, leg. Drimmer, Pintér & Varga 20.7.1994, HNHM 43528/3+1 (juv.); on the shore of the
Kourna Lake (= W of Episkopí) under stones, UTM KV 51, leg. Butot & Subai 12.2.1981, S
14103/32+18 (juv.)+6 (alk.)+2 (juv. alk.); leg. Maassen 4.1987, Maa/4.
Nomos Réthimno: 1 km N of Assomatos (= SW of Spíli) UTM KU 69, leg. Maassen
10.1992, Maa/2; hill slope in the village of Agía Galíni, on rocks, UTM KU 88, leg. Pintér
& Subai 10.8.1976, S 3828/1; 4 km S of Ágios Ioánnis, UTM KU 99, leg. Maassen 10.1992,
Maa/1 (juv.); road crossing close to Ágios Konstantínos, UTM KV 60, leg. Drimmer, Pintér
& Varga 20.7.1994, HNHM 43527/3+1 (juv.); Atsipopoulo (= W of Réthimno) UTM KV 61,
leg. Drimmer, Pintér & Varga 20.7.1994, HNHM 43523/1; between Kato Valsamónero
and Ágios Andréas, UTM KV 61, leg. Drimmer, Pintér & Varga 20.7.1994, HNHM 43525/1;
road crossing close to Kaloníktis, UTM KV 61, leg. Drimmer, Pintér & Varga 20.7.1994,
HNHM 43526/1; 2 km W of Ágios Andréas, UTM KV 61, leg. Kiss & Pintér 17.7.1986, HNHM
29877/20+15 (juv.+); E of Geráni (= W of Réthimno) at the cave, UTM KV 61, leg. Maas-
sen 4.1987, Maa/7; river Pétres (= 9 km W of Réthimno) UTM KV 61, leg. Butot & Subai
11.2.1981, S 17511/1; leg. Maassen 4.1990, Maa/6; 1 km S of Réthimno, UTM KV 71, leg.
Kiss & Pintér 15.7.1986, HNHM 29878/1 (juv.); 3 km S of Réthimno, UTM KV 71, leg. Drim-

38 Peter Subai & Eike Neub ert
mer, Pintér & Varga 20.7.1994, HNHM 43522/2+2 (juv.); canyon 5 km SE of Réthimno, at
the bridge, UTM KV 71, leg. Maassen 10.1992, Maa/1; 2 km S of Prasiés, UTM KV 71, leg.
Maassen 10.1992, Maa/1; S boundary of Xeró Horió, UTM KV 71, leg. Maassen 4.1990,
Maa/12; 2 km S of Xeró Horió, UTM KV 71, leg. Maassen 4.1990, Maa/1 (juv.); 2 km E
of Apóstoli, UTM KV 80, leg. Maassen 10.1992, Maa/3; S boundary of Kalógeros, UTM
KV 80, leg. Kiss & Pintér 14.8.1988, HNHM 44684/1; Monastir Arkadí, on walls, UTM
KV 81, leg. Maassen 10.1992, Maa/2; 1 km N from the Monastir Arkadí, UTM KV 81, leg.
Maassen 10.1992, Maa/3; in the village of Eléftherna, UTM KV 81, leg. Drimmer, Pintér &
Varga 22.7.1994, HNHM 43530/4; 0.5 km N of Amnátos, UTM KV 81, leg. Drimmer, Pintér
& Varga 20.7.1994, HNHM 43529/1; 1 km N of Pérama, UTM KV 91, leg. Maassen 10.1992,
Maa/3; leg. Drimmer, Pintér & Varga 22.7.1994, HNHM 43532/2; rocks at the N boundary
of of Orthés, UTM KV 91, leg. Maassen 10.1992, Maa/1; SE boundary of Orthés, UTM KV
91, leg. Drimmer, Pintér & Varga 22.7.1994, HNHM 43531/2 (juv.); Bali, UTM KV 92, leg.
Maassen 1.1993, Maa/1; 3 km W of Anógia, UTM LV 00, leg. Maassen 4.1990, Maa/1;
at the boundary of Aímonas, UTM LV 01, leg. Drimmer, Pintér & Varga 11.7.1994, HNHM
43521/2+1 ( juv.); 1 km SE of Sises (= in direction to Achláda) UTM LV 11, leg. Drimmer,
Pintér & Varga 11.7.1994, HNHM 43520/2+4 .
Nomos Iráklio: 1 km W of Panassós (= S of Ágia Varvára) UTM LU, 18, leg. Maassen
10.1992, Maa/1; Houdétsi, UTM LU 39, leg. Maassen 10.1991, Maa/1; 2 km of Astiráki
in direction to Tílisos, UTM LV 10, leg. Maassen 10.1992, Maa/4; Park in Fódele, UTM LV
11, leg. Maassen 10.1992, Maa/2; 1 km N of Fódele, UTM LV 11, leg. Maassen 10.1992,
Maa/1; Rodiá, UTM LV 21, leg. Maassen 10.1991, Maa/2; "Tris Ekklesias" close to Rodiá,
UTM LV 21, leg. Drimmer, Pintér & Varga 11.7.1994, HNHM 43519/1+4 (juv.); E of Irák-
lio, an of the crossing the road to Prasás, UTM LV 31, leg. Maassen 4.1990, Maa/4; leg.
Maassen 1.1993, Maa/4; valley E from the airport of Iráklio, on limestone rocks, under
stones, UTM LV 31, leg. Butot & Subai 13.2.1981, S 14098/14+16 (juv.)+5 (alk.)+1 (juv.,
alk.); 4 km N of Kastéli, UTM LV 50, leg. Maassen 4.1990, Maa/2.
Nomos Lassíthi: Vidiáni, UTM LU 59, leg. Pintér & Varga 3.7.1993, HNHM 45291/1 (juv.).
Fig. 19. Distribution of Lindholmiola barbata.

39
Revision of the genus Lindholmiol a Hesse, 1931
Remarks: Concerning both, size and shell shape, L. barbata does not vary
considerably. In this species, no local forms are known.
Distribution (Fig. 19): This species is endemic to the Island of Crete, where
it occupies the Lassíthi-plane towards the central and western part of the
island.
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 contor ta 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 typi-
cus: "Corfu". Type material: lost [(partim, not the variety minor = girva Fri-
valdszky, 1835) (homonym of Helix contorta Linnaeus, 1758)].
1839 Helix corcyrensis, – Deshayes in Férussac & Deshayes, 1820–1851: His-
toire naturelle générale et particulière des mollusques terrestres et fluvia-
tiles, 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.

40 Peter Subai & Eike Neub ert
1859 Helix corcyrensis var. octogyrata Mousson, Vierteljahrsschrift der Natur-
forschenden Gesellschaft in Zürich, 4: 257.
1859 Helix corcyrensis var. cefalonica Mousson, Vierteljahrsschrift der Natur-
forschenden Gesellschaft in Zürich, 4: 29.
1859 Helix corcyrensis var. canalifera, – Mousson, Vierteljahrsschrift der Natur-
forschenden Gesellschaft in Zürich, 4: 17, 257.
1859 Helix corcyrensis var. girva, – Mousson, Vierteljahrsschrift der Naturfor-
schenden 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 Thessa-
lie: 36.
1879 Helix (Trigonostoma) corcyrensis var. octogyrata, – Westerlund & Blanc,
Aperçu sur la faune malacologique de la Grèce inclus l’Epire et la Thessa-
lie: 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 Con-
chology, 3: 118.
1887 Helix (Caracolina) corcyrensis var. octogyrata, – Tryon, Manual of Con-
chology, 3: 118.

41
Revision of the genus Lindholmiol a Hesse, 1931
1887 Helix (Caracolina ) corcyrensis var. canalifera, – Tryon, Manual of Con-
chology, 3: 118.
1889 Helix (Caracolina) corcyrensis, – Westerlund, Fauna der in der paläarcti-
schen 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 Deut-
schen Malakozoologischen Gesellschaft, 23 (5/6): 83.
1894 Helix (Gonostoma) corcyrensis, – Sturany, Annalen des Kaiserlich-König-
lichen 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 Con-
chology, 9: 288.
1894 Helix (Caracollina) corcyrensis var. octogyrata, – Pilsbry, Manual of Con-
chology, 9: 288.
1894 Helix (Caracollina) corcyrensis var. canalifera, – Pilsbry, Manual of Con-
chology, 9: 288.
1905 Gonostoma corcyrensis, – Sturany, in Paganetti-Hummler: Verhandlun-
gen 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 Malako-
zoologischen Gesellschaft, 50 : 109.
1918 Caracollina corcyrensis octogyrata, – Hesse, Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 50: 109.
1918 Caracollina corcyrensis cephalonica, – Hesse, Nachrichtsblatt der Deut-
schen 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.

42 Peter Subai & Eike Neub ert
1930 Caracollina contorta contorta, – Käufel, Sitzungsberichte der Akademie
der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse,
139 (3/4): 183.
1930 Caracollina contor tata cephalonica, – Käufel, Sitzungsberichte der Aka-
demie 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-Naturwis-
senschaftliche Klasse, Abt. I, 171 (6/7): 253.
1962 Lindholmiola corcyrensis cephalonica, – Klemm, Sitzungsberichte der
Österreichischen Akademie der Wissenschaften, Mathematisch-Naturwis-
senschaftliche 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 Naturforschen-
den 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 Abhand-
lungen aus dem Staatlichen Museum für Tierkunde Dresden, 14 (12): 106.
1990 Lindholmiola corcyrensis, – Reischütz & Sattmann, Annalen des Natur-
historischen Museums in Wien, 91 (B): 258.
1990 Lindholmiola girva, – Fechter & Falkner, Steinbachs Naturführer: 222,
Abb. 6 (partim) (non girva Frivaldszky, 1835).

43
Revision of the genus Lindholmiol a Hesse, 1931
1996 Lindholmiola girva, – Welter-Schultes, Schriften zur Malakozoologie, 9:
23–31 (non gir va Frivaldszky, 1835).
1996 Lindholmiola girva, – Dhora & Welter-Schultes, Schriften zur Malakozo-
ologie, 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 col-
our light to chestnut brown; first ½ to ¾ protoconch whorl smooth, then
with a few fine radial stripes; the following teleoconch whorls with increas-
ingly 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 cov-
ered 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, peduncu-
lus and vesicle of the bursa copulatrix of the same length; atrial lumen filled

44 Peter Subai & Eike Neub ert
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
Figs 20–2 3. 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., hik-
ing path of Pápigo, 1–2 Km N of the shelter (large umbilicus, flat shell), D = 11.23 mm; Fig. 23: syn-
type Helix (Trigonostoma) corcyrensis var. cefalonica, ZMZ 508737, Island Kefaloniá, "au x environs
d'Argostoli, ...à Frangata et Grisata", D = 12.0 mm. — All photos Neuber t/Bochud, × 3.

45
Revision of the genus Lindholmiol a Hesse, 1931
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
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.

46 Peter Subai & Eike Neub ert
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 aper-
ture is narrower and sometimes shows a tooth-like swelling. As in L. pirinensis,
L. gir va 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 weak-
er labial callus. The penial lumen of L. barbata shows two parallel elongated
pilasters and thus differs profoundly from the v-shaped pilasters of L. corcy-
rensis.
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, 42.4 4°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

47
Revision of the genus Lindholmiol a Hesse, 1931
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
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 corc yrensis, 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.

48 Peter Subai & Eike Neub ert
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 (= direc-
tion 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. 30 0 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,

49
Revision of the genus Lindholmiol a Hesse, 1931
in forest 10 km from the crossing Carina–Sv. Naum– Ohrid direction Carina, on lime-
stone 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 Trpe-
jca, 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 11.5.200 0, 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 Pan-
agia 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 & Szek-

50 Peter Subai & Eike Neub ert
eres 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 40 0 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.); Auf-
fahrtweg zum Pantokrátor, 700 m alt., UTM CJ 99, 39.7499°N, 19.8428°E, leg. Neute-
boom 3.10.1991, NNM/1; Áno Korakiana, UTM CJ 99, 39.70°N, 19.79°E, leg. Neute-
boom 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 lime-
stone 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 high-
way, 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, 39.77 34°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, byzanti-
nische Festung, UTM DJ 07, 39.4770°N, 19.88 48°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. Gittenberg-
er 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

51
Revision of the genus Lindholmiol a Hesse, 1931
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 [Moun-
tain], 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 (= Lef-
kas), 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 lime-

52 Peter Subai & Eike Neub ert
stone 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, 4 0 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 Igou-
menítsa–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–Igou-
menítsa, 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 bifur-
cation 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 Mórf i, 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,

53
Revision of the genus Lindholmiol a Hesse, 1931
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. Gitten-
berger & Uit de Weerd 15.5.200 0, 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) ceme-
tery, 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
16.7.199 0, 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

54 Peter Subai & Eike Neub ert
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, 1650–170 0 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, 40.040 8°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 20.90 69°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. — Etoloakarn-
anía, 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.

55
Revision of the genus Lindholmiol a Hesse, 1931
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 direc-
tion Sardínia, 6 0 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 consid-
ered a nomen oblitum. However, Rossmässler (1838: 40, Taf. 39, Fig. 538) vali-
dated 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 Ross-
mässler 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.

56 Peter Subai & Eike Neub ert
Distribution (Fig. 29): Lindholmiola corcyrensis inhabits the south-western
Balkan Peninsula. There are two isolated localities known, the Pelješac Penin-
sula 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 distri-
bution area follows the eastern rim of the Pindos Mountains.
Fig. 29. Distribution of Lindholmiola corcyrensis and L. girva.

57
Revision of the genus Lindholmiol a Hesse, 1931
Lindholmiola girva (Frivaldszky, 1835) Figs 14, 29, 30–32, 33–34
1835 Helix girva Frivaldszky, A Magyar Tudós Társaság évkönyvei, 2: 274, Taf. 7
Fig. 11.
1838 Helix contorta var. minor Rossmässler, Iconographie der Land- & Süss-
wasser-Mollusken mit vorzüglicher Berücksichtigung der europäischen noch
nicht abgebildeten Arten, (1) 2 (1/2): 40, Taf. 39 Fig. 538 b.
1848 Helix corcyrensis var. ß L. Pfeiffer, Monographia heliceorum viventium 1:
415.
1859 Helix corcyrensis var. girva, – Mousson, Vierteljahrsschrift der Naturfor-
schenden Gesellschaft in Zürich, 4: 257, 259.
1865 Helix corcyrensis, – L. Pfeiffer, Malakozoologische Blätter, 12: 103 [non
corcyrensis Rossmässler, 1838] .
1868 Helix corcyrensis var. minor – L. Pfeiffer, Monographia heliceorum viven-
tium 5: 415.
1887 Helix (Caracolina) corcyrensis var. girva, – Tryon, Manual of Conchology,
3: 118.
1889 Helix (Caracolina) corcyrensis var. girva, – Westerlund, Fauna, 2: 20.
1894 Helix (Caracollina) corcyrensis var. girva, – Pilsbry, Manual of Conchology,
9: 288.
1894 Helix (Gonostoma) corcyrensis var. girva, – Sturany, Annalen des Kaiser-
lich-Königlichen Naturhistorischen Hofmuseums, 9: 371.
1915 Helicodonta (Caracollina) contorta girva, – Sturany & Wagner, Denkschrif-
ten der Kaiserlichen Akademie der Wissenschaften in Wien, Mathematisch-
Naturwissenschaftliche Klasse, 91: 32, Taf. 2 Fig. 8 a–c.
1918 Caracollina corcyrensis girva, – Hesse, Nachrichtsblatt der Deutschen
Malakozoologischen Gesellschaft, 50: 109.
1927 Caracollina contorta girva, – A. J. Wagner. Annales zoologici Musei Polonici
Historiae Naturalis, 6 (4): 364, Taf., 19 Fig. 169.
1929 Caracollina corcyrensis forma gir va, – Hesse, Senckenbergiana, 11
(1/2): 96.
1936 Lindholmiola corcyrensis, – Fuchs & Käufel, Archiv für Naturgeschichte,
5 (4): 651, Fig. 78 (partim, not those of Podgorica) (non corcyrensis Ross-
mässler, 1838).
1960 Lindholmiola contorta, – Urbanski, Bulletin de l'Institut Zoologique de
l'Académie Bulgare des Sciences, 9: 92.
1971 Lindholmiola corcyrensis, – Schileyko, Nauchnykh Dokladi Vysshey Shkoly,
Biologicheskie nauki., (Biol. Wiss. Zool.), 12: 12, Fig. 2 B (p.11), Fig. 4, Fig. 5d
[non corcyrensis Rossmässler, 1838].

58 Peter Subai & Eike Neub ert
1972 Lindholmiola (Lindholmiola) corcyrensis, – Hudec, Annales zoologici Musei
Polonici Historiae Naturalis, 29 (10): 337, Abb. 9, Taf. 3 Fig. 7 [non corcy-
rensis Rossmä ssler, 1838].
1973 Lindholmiola (Lindholmiola) corcyrensis girva, – Hudec & Vašátko, Acta
scientiarum naturalium Academiae Scientiarum Bohemoslovacae, 7 (9): 13.
1983 Lindholmiola corcyrensis, – Reischütz, Annalen des Naturhistorischen
Museums in Wien, 85 (B): 135.
1978 Lindholmiola corcyrensis, – Shileyko, Fauna UDSSR, 3 (6): 116, 118, Fig.
44–45 [non corcyrensis Rossmässler, 1838].
1988 Lindholmiola corcyrensis corcyrensis, – Reischütz, Annalen des Natur-
historischen Museums in Wien, 90 (B): 347 [non corcyrensis Rossmässler,
1838].
1988 Lindholmiola corcyrensis girva, – Reischütz, Annalen des Naturhistorischen
Museums in Wien, 90 (B): 347.
Diagnosis: shell depressed, shell surface covered by granulation, whorls
slightly to strongly shouldered, penial lumen with two equally sized pilasters.
Description of shell: shell depressed, spire only slightly raised, shell col-
our light to chestnut brown; initial half protoconch whorl smooth, then with a
dense pattern of few fine radial stripes; the following teleoconch whorls with
increasingly stronger stripes, the radial sculpture considerably pronounced
on the second and third teleoconch whorls, but often decreasing on the last
whorls; radial threads may irregularly vary in height, while on the subsurface
of the shell, stripes are usually weak and blurred; the complete shell surface
covered by a fine granulation giving the shell a dull appearance; hair warts
are missing in general; besides a small, hair-free zone below the suture, shell
completely covered by permanent hairs, hairs in the umbilicus usually short-
er than those on the rest of the shell; hairs brownish-yellowish, hair length
at the shell’s shoulder ca 0.4–0.9 mm; 5¾–7½ regularly increasing, slightly
to strongly shouldered whorls, never keeled; last whorl slightly descending
before the aperture, then abruptly curving downwards, the down-bent part in
a brighter to white colour; suture deep; umbilicus almost cylindrical to slight-
ly perspective, only slightly increasing at the beginning, and the last whorl
remaining circular, reaching a diameter of 1.4–2.5 mm; aperture very oblique
and strongly bent in lateral view, and ear-like in frontal view, upper part flat;
apertural insertion with a distance of 2.3–3.9 mm, connected by a very thin,
almost indiscernible callus; peristomial rim sharp, laterally with a strong swell-
ing, basally reinforced, forming a narrow lip (sometimes even with a small
tooth); umbilical shield small, covering less than ca 1/10 of the umbilicus.
Measurements: H: 4.2–6.1; D: 8.1–11.7; aH: 3.3–5(1.6–3.4); aW: 3.75–5.8.

59
Revision of the genus Lindholmiol a Hesse, 1931
Details of body (according to 14 animals from 10.5 km N of Vorinó/3, Kéla/2,
Taxiárhis/6, and 2.5 km of Paleá Kavála towards Korifés/3): head and dorsum
light brown to grey, lateral side of body brighter; sole cream with brownish
lateral rims; mantle with a pattern of dispersed, irregularly arranged, dark-grey
spots covering the last 1–1½ whorls, often with a broad brown area on the last
half of the last whorl; ureter opens 0.5–1.5 mm from the pneumostome.
Morphology of the genital organs (Fig. 14): penis long, slightly enlarged,
penial glandular tissue covers ca 1/3–1/6 of the total penis length, penial
lumen narrow, with two equally sized pilasters, starting at the distal penial
wall and stretching to at least half of the penis length, a third usually shorter
and sometimes completely missing pilaster, and sometimes with short per-
pendicular folds between the pilasters; vagina short, glandula usually of the
same length as bursa copulatrix with a scarcely folded part, pedunculus and
vesicle of bursa copulatrix not observably differentiated; atrial lumen with
a large, broad and flat, serrated pilaster leading towards the vaginal lumen,
usually with another narrow fold in parallel.
Figs 30–32 . Lindholmiola girva. Fig. 30: syntype Helix girva SMF 7047, "Albanien", D = 9.22 mm;
Fig. 31: NMBE 515656, Greece, Makedonía, N boundary of Taxiárhis (small rounded form), D = 8.66 mm;
Fig. 32: NMBE 515655, Greece, Makedonía, 400 m S of Kehrókambos (broad form, wide umbilicus),
D = 9.75 mm. — All photos Neubert/Bochud, × 3.

60 Peter Subai & Eike Neub ert
Differential diagnosis: Large specimens of L. girva are quite similar to small
shells of L. lens (= lentiformis sensu auct.), which can be found in Viotía,
Fthiótida and on the Island Évvia. This form of L. lens, however, differs from L.
girva by having a blunt peripheral keel, a flatter spire, and continuous acute
radial riblets; in lateral view, its aperture is less strongly curved, its upper
part not narrowed by the labial callus, as it is found in L. girva. In the geni-
tal morphology, L. lens has a relatively shorter penis, but a longer part of it is
covered by the glandular tissue, and the pilasters in its penial lumen are usu-
ally shorter.
On average, shells of L. pirinensis are larger than those of L. girva. In this
species, the shell surface is covered with finer and more regularly arranged
radial riblets, its hairs are longer but are lost more easily; in lateral view, its
aperture is less strongly curved and lipped. In the genital morphology, L. piri-
nensis differs in the form of one of the penial pilasters, which is recurved and
points back towards the distal penial wall.
Lindholmiola girva has often been confused with L. corcyrensis, which is
larger on average. In this species, the teleoconch whorls are more rounded
with a blunt periphery. The radial riblets are more irregularly arranged, and
the last whorl is broader. The umbilicus is oval in both species, but in L. corcy-
rensis, it is more eccentric. Both species can easily be separated by the form
of their penial pilasters, which consist of small v-shaped folds in L. corcyrensis
instead of the longitudinal pilasters in L. girva.
Type specimens: "Albanien", UTM n. det., syntype girva: SMF 7047/1; "...a "Balkány"
hegyeinek aljasabb domboldalaiban" (see remarks).
Additional specimens examined:
Moldavia, Kodry (= Codrii, Chodrii) reservat (= 30 km WNW of Chişinau), UTM PN
01, 47.08°N 28.42°E, leg. Mészáros 14–20.7.1986, HNHM 23876/1.
Romania, Craiova, UTM GQ 20, 44.30°N 23.80°E, leg. Knorr, 1940, SMF 279320/2;
Slobozia (= Slobosia), UTM MJ 15, 43.85°N 25.91°E, leg. Wohlberedt, 1903, SMF
101903/3; Bucuresti–Băneasa, UTM MJ 27, 44.49°N 26.08°E, leg. Grossu, 18.5.1964,
SMF 227417/5; Comana (= near Bucureşti), UTM MJ 39, 44.16°N 26.14°E, leg. Wohl-
beredt, 1903, SMF 101902/3; Kuznetzov 11.1985, Maa/2; Bucureşti (= Bukarest),
UTM n. det., SMF 101904/3, 284877/4; Fundeni (= Fundiaeni), UTM MK 41, 44.38°N
26.35°E, leg. Goldfuß, 1884, SMF 284862/1; Căldărusani monastery (= Caldarouchani),
UTM MK 44, 44.6761°N 26.2665°E, leg. Licherdopol, 1901, SMF 101925/4; Adamclisi (=
Adamklissi), UTM NJ 78, 44.08°N 27.94°E, leg. Martens 2.7.1918, SMF 284835/1; Man-
galia, UTM PJ 25, 43.83°N 28.58°E, leg. Montandon, 1902, SMF 101924/4; forest S of
Babadag, UTM PK 37, 44.88°N 28.71°E, 7.6.1938, SMF 207889/2.
Fyrom, Katlanovo, UTM EM 53, 41.89°N 21.68°E, leg. Maassen 7.1980, Maa/3;
leg. Kiss & Pintér 4.7.1985, HNHM 30857/5 (+ 1 pirinensis); Demir Kapija, UTM

61
Revision of the genus Lindholmiol a Hesse, 1931
FL 08, 41.40°N 22.23°E, leg. Brankovics 2.5.1973, HNHM 49541/5; leg. Kiss &
Pintér 4.7.1986, HNHM 29884/1 (juv.); Demir Kapija, banks of Vardar River, UTM FL
08, 41.40°N 22.25°E, leg. Mikuska & Sipos 28–30.9.1975, HNHM 49538/3 + 1 (juv.),
49539/9 + 3 (juv.); gorge 3.5 km from Demir Kapija direction Gevgelija, banks of
Vardar River, on limestone rocks, UTM FL 08, 41.40°N 22.30°E, leg. Maassen 7.1980,
Maa/24 + 2 (juv.); leg. Subai 28.7.1990, S 15637/7; Stari Dojran, church ruins, UTM FL 46,
41.17°N 22.72°E, leg. Neuteboom 15.7.1976, NNM/19.
Bulgaria, Kostinbrod near Sofija, UTM FN 84, 42.81°N 23.22°E, leg. Mészáros
9.1981, HNHM 13871/1 (juv.) (+ 1 pirinensis); [Gara] Lakatnik, UTM FN 97, 43.08°N
23.38°E, leg. Pintér 5.8.1981, HNHM 33702/3 + 2 (juv.); 2.5 km from [Gara] Lakatnik
towards Milanovo, UTM FN 97, 43.10°N 23.39°E, leg. Kiss & Pintér 15.7.1984, HNHM
37469/1 (juv.); between Bačkovo and Dobrostan [Mountain], UTM LG 24, 41.95°N
24.86°E, leg. Pintér 17.7.1970, HNHM 33700/1 (juv.); Asenovgrad, UTM LG 25,
42.00°N 24.86°E, leg. Pintér 25.7.1967, SMF, 198754/1; Karlovo, UTM LH 22, 42.65°N
24.81°E, leg. Pintér 28.7.1967, HNHM 33697/1 + 1 (juv.); Šipka Pass, UTM LH 63,
42.73°N 25.34°E, leg. Pintér 13.7.1973, HNHM 49536/1; Ropotamo Bank, Arkutino
Camping, sand dune, UTM NG 67, 42.32°N 27.73°E, leg. Szigethy 4.7.1973, HNHM
49550/1; Madara bei Šumen, UTM NH 09, 43.27°N 27.10°E, leg. Kroupa 6.8.1978,
SMF 254021/14; N border of Beloslav (= W of Varna), UTM NH 58, 43.20°N 27.70°E,
leg. Pintér, 19.7.1968, HNHM 33698/61 + 13 (juv.); Kamcija (= S of Varna), UTM NH
76, 43.02°N 27.88°E, leg. Pintér 16.7.1968, HNHM 33695/8 + 5 (juv.); Galata (= S of
Varna), UTM NH 77, 43.16°N 27.93°E, 1936, SMF 101927/2; Varna, UTM NH 78, SMF
284872/2; leg. Urbanski 8.1936, SMF 158897/8; Varna, coast, UTM NH 78, 43.20°N
27.92°E, leg. Balogh 20.6.1956, HNHM 33701/2; Aladža monastery (= N of Varna),
UTM NH 89, 43.27°N 28.01°E, leg. Podani 9.8.1980, HNHM 11641/1.
Greece, Makedonía, in dry valley at NW border of Kéla, on limestone rocks, 820 m
alt., UTM EL 51, 40.3482°N 21.7730°E, leg. Subai & Szekeres, 19.05.1997, S 15677/3
+ 3 (alk.); 2 km S of Vévi (= 17.5 km ESE of Flórina), 900 m alt., UTM EL 51, 40.74°N
21.61°E, leg. Gittenberger & Uit de Weerd 10.5.2000, NNM/1; rocks 11 roadkm. of
Kéla (= 24 km W of Édessa), 650 m alt., UTM EL 61, 40.81°N 21.78°E, leg. Gittenberg-
er & Uit de Weerd 11.5.2000, NNM/6; 1.5 km S of Árnissa, on dr y limestone rocks,
520 m alt., UTM EL 71, 40.7775°N 21.8352°E, leg. Subai 09.05.1995, S 15648/15 + 1
(alk.); in valley behind Loutráki village, on marble rocks, 350 m alt., UTM EL 73,
40.9769°N 21.9454°E, leg. Subai & Szekeres, 19.05.1997, S15676/25; 8 km N of Édes-
sa (= Édessa–Ápsalos country road), 3.8 roadkm. N of Sotíra, on rocks at S rim oft he
road, 180 m alt., UTM EL 82, 40.88°N 22.05°E, leg. Gittenberger & Uit de Weerd
10.5.2000, NNM/2; gorge 2 km N of Neohóri (= NE of Aridéa), on limestone rocks,
250 m alt., UTM EL 94, 41.0475°N 22.0916°E, leg. Subai 09.05.1995, S 15649/1; valley
10.5 km N of Vorinó (= direction Pefkotó) and 1 km walking path to limestone rocks,
850 m alt., UTM EL 94, 41.0846°N 22.0849°E, leg. Subai & Szekeres, 19.05.1997, S
15675/21 + 3 (alk.); above Nótia, on dry limestone rocks, 650 m alt., UTM FL 05,
41.1072°N 22.2115°E, leg. Subai 09.05.1995, S 15650/6; on limestone rocks 200 m N
of Nótia, 600 m alt., UTM FL 05, 41.10°N 22.20°E, leg. Gittenberger & Uit de Weerd
10.5.2000, NNM/3; gorge at N border of Gríva, on limestone rocks, 480 m alt., UTM FL
13, 40.9574°N 22.4055°E, leg. Subai 09.05.1995, S 15647/12; 500 m W of Gríva (=
WSW of Axioúpoli), on limestone rocks, 540 m alt., UTM FL 13, 40.9562°N 22.4009°E,

62 Peter Subai & Eike Neub ert
leg. Subai & Szekeres 20.05.1997, S 15683/4 + 1 (alk.); bifurcation of highway Rendi-
na–Asproválta to Stavrós, on crystalline rocks, 15 m alt., UTM GL 20, 40.6728°N
23.6633°E, leg. Subai 28.05.1997, S 15660/2; mountain slope N of Podohóri, on lime-
stone rocks, 450 m alt., UTM KF 42, 40.8473°N 24.0249°E, leg. Subai 27.05.1997, S
15672/15 + 3 (juv.); Mesorópi (= W of Kavála), UTM KF 52, 40.8 6°N 24.08°E, leg.
Maassen 7.1980, Maa/1; Kokkinohóri, UTM KF 52, 40.81°N 24.00°E, leg. Neuteboom
11.7.1976, NNM/2 (juv., alk.); on rocks E of Platantopos (= "Platanohóri" ?), UTM KF
52, 40.84°N 24.06°E, leg. Neuteboom 11.7.1976, NNM/6; E border of Mousthéni, on
low limestone rocks, 175 m alt., UTM KF 52, 40.8566°N 24.1137°E, leg. Subai
06.05.1995, S 15658/7 + 3 (alk.); on marble rocks 1 km from Platanótopos direction
Elef theroupoli (= 4.7 km NE of Podohóri), 280 m alt., UTM KF 52, 40.8524°N
24.0717°E, leg. Subai 27.05.1997, S 15671/3; Simvolo Mountains, 1.2 km from Sidiro-
hóri direction Foliá (= W of Karavangélis), on limestone rocks, 250 m alt., UTM KF 52,
40.8380°N 24.1375°E, leg. Subai 27.05.1997, S 15670/2; 1 km NE of Akrovoúni, 180
m alt., UTM KF 52, 40.90°N 24.21°E, leg. Gittenberger & Uit de Weerd, 18.5.1999,
NNM/3; small gorge at S border of Nikíssiani, 350 m alt., UTM KF 53, 40.9467°N
24.1439°E, leg. Fauer & Subai 22.09.1989, S 15656/1; beech forest 7.5 km from
Akrovouni direction Pangéo peak, in leaf litter and under stones, 480 m alt., UTM KF
53, 40.9237°N 24.2178°E, leg. Subai 27.05.1997, S 15669/4 + 1 (juv.) + 2 (alk.); river
valley 8 km from Akrovouni direction Pangéo peak, in forest, on limestone rocks, 600
m alt., UTM KF 53, 40.9210°N 24.2060°E, leg. Fauer & Subai 23.09.1989, S 15641/6;
oak forest 11.6 km from Akrovouni direction Pangéo peak, on limestone rocks, 90 0 m
alt., UTM KF 53, 40.9171°N 24.1906°E, leg. Fauer & Subai 23.09.1989, S 15642/2;
river valley 12.3 km from Akrovouni direction Pangéo peak, on limestone rocks, 950
m alt., UTM KF 53, 40.9222°N 24.1876°E, leg. Fauer & Subai 23.09.1989, S 15643/2;
17.8 km from Akrovouni direction Pangéo peak, on NW exposed limestone rocks,
1280 m alt., UTM KF 53, 40.9147°N 24.1465°E, leg. Fauer & Subai 23.09.1989, S
15644/2; 22.7 km from Akrovouni direction Pangéo peak, on limestone rocks, 1600
m alt., UTM KF 53, 40.9079°N 24.1176°E, leg. Fauer & Subai 23.09.1989, S 15645/1;
25.2 km from Akrovouni direction Pangéo peak, meadow with limestone rocks below
the peak, 1780 m alt., UTM KF 53, 40.9113°N 24.0973°E, leg. Fauer & Subai
23.09.1989, S 1564 6/2; 2.5 km of Pangéo peak, 1650 m alt., UTM KF 53, 40.91°N
24.10°E, leg. Gittenberger & Uit de Weerd, 18.5.1999, NNM/1; 4.5 km of Pangéo peak,
1400 m alt., UTM KF 53, 40.90°N 24.12°E, leg. Gittenberger & Uit de Weerd,
18.5.1999, NNM/2; mountain slope 4.5 km W of the bifurcation to Loutrá Eleftheró (=
48 km W of Kavála), on limestone rocks, 30 m alt., UTM KF 51, 40.7211°N 24.0601°E,
leg. Fauer & Subai 21.09.1989, S 15659/22; N border of Taxiárhis, under crystalline
stones, 500 m alt., UTM KF 66, 41.2328°N 24.1888°E, leg. Subai 22.05.1997, S
15679/5 + 8 (alk.); Batis, UTM KF 73, 40.91°N 24.37°E, leg. Neuteboom 12.7.1976,
NNM/1; ruin field of Philippi near Krinídes, UTM KF 74, 41.01°N 24.28°E, leg. Neute-
boom 12.7.1976, NNM/1 (juv.); leg. Maassen 7.1980, Maa/9 + 2 + 3 (juv.); leg. Kiss &
Pintér 7.7.1986, HNHM 29882/2; 5 km from Limnia direction Zigós, on low limestone
rock s and under stones, 730 m alt., KF 74, 41.0589°N 24.3816°E, leg. Subai
26.05.1997, S 15667/1; 1 km W of Peristéria (= direction Adriani), on limestone rocks,
approx. 4 00 m alt., UTM KF 75, 41.1177°N 24.3837°E, leg. Subai 23.05.1997, S
15673/2; 1.3 km from Agorá direction Peristéria, on limestone rocks, approx. 380 m

63
Revision of the genus Lindholmiol a Hesse, 1931
alt., UTM KF 75, 41.1165°N 24.3244°E, leg. Subai 23.05.1997, S 15674/1; 1 km E of
Nikifóros, UTM KF 76, 40.17°N 24.33°E, leg. Kiss & Pintér 5.7.1986, HNHM 29886/1;
in Makriplágio Village, beech forest and border of forest with stone walls, under
stones and in leaf litter, on crystalline rocks, 880 m alt., UTM KF 76, 41.2267°N
24.2713°E, leg. Subai 22.05.1997, S 15680/11; beech forest 3.2 km N of Makriplágio
(along the highway), under stones, in leaf litter, crystalline rocks, 900 m alt., UTM KF
76, 41.2350°N 24.2945°E, leg. Subai 22.05.1997, S 15678/1 + 1 (juv) + 1 (alk.); Kavá-
la, citadel, UTM KF 83, 40.93°N 24.41°E, leg. Maassen 7.1980, Maa/7 + 1 (juv.); 2 km
E of Kavála, UTM KF 83, 40.95°N 24.45°E, leg. Maassen 7.1980, Maa/1; 2.5 km from
Paleá Kavála direction Korifés, on low limestone rocks and under stones, 500 m alt.,
UTM KF 84, 41.0170°N 24.4388°E, leg. Subai 07.05.1995, S 15651/15 + 3 (alk.); 3 km
W of Paleá Kavála, on limestone rocks above the river, 200 m alt., UTM KF 84,
41.0009 °N 24.3934°E, leg. Subai 07.05.1995, S 15654/1; 4.2 km from Paleá Kavála
direction Halkeró, S exposed mountain slope, on limestone rocks, 440 m alt., UTM KF
84, 40.9994°N 24.4273°E, leg. Subai 26.05.1997, S 15664/1; on limestone rocks at
SE border of Likóstomo, 740 m alt., UTM KF 85, 41.1078°N 24.4424°E, leg. Subai
26.05.1997, S 15668/7; 4 km from Platamónas direction Lekáni, 720 m alt., UTM KF
95, 41.1149°N 24.5478°E, leg. Subai 07.05.1995, S 15655/4 + 3 (alk.); 800 m W of
Dipótamos, on limestone rocks beside highway, 690 m alt., UTM KF 95, 41.1196°N
24.5762°E, leg. Subai 26.05.1997, S 15663/8; on low marble rocks at Disvato Village,
under stones and in leaf litter, 450 m alt., UTM KF 95, 41.0681°N 24.5713°E, leg.
Subai 26.05.1997, S 15665/5 + 1 (juv.); stone hedge in valley of two mountains 1 km
from Ágios Kosmás direction Makrihóri, 380 m alt., UTM LF 04, 41.0783°N 24.6536°E,
leg. Subai 26.05.1997, S 15662/1; 1 km N of Zarkadia (= 1.3 km along the road), UTM
LF 04, 41.02°N 24.63°E, leg. Gittenberger 29.5.1999, NNM/2; limestone mountain
approx. 2.5 km NNE of Zarkadía (along the road), 450 m alt., UTM LF 04, 41.0288°N
24.6417°E, leg. Kiss & Pintér 7.7.1986, HNHM 29885/6 + 2 ( juv.); leg. Fauer & Subai
22.9.1989, S 15657/2; Subai 7.5.1995, S 15653/4; 400 m S of Kehrókambos, under
(lime) stones, 390 m alt., UTM LF 05, 41.1514°N 24.6454°E, leg. Subai 07.05.1995, S
15652/3; small river gorge 4.1 km from Kehrókambos direction Makrihóri, stone
hedge, under stones, 4 85 m alt., UTM LF 05, 41.1292 °N 24.6448°E, leg. Subai
26.05.1997, S 15666/1; 7.2 km from Kehrókambos direction Makrihóri, at crossing to
Skopós, under stones, 550 m alt., UTM LF 05, 41.1043°N 24.6392 °E, leg. Subai
26.05.1997, S 15661/1. — Thráki, gorge at N border of Xánthi, UTM LF 25, 41.14°N
24.88°E, leg. Maassen 7.1980, Maa/2; surroundings of the cave of Marónia (= NW of
Marónia), UTM LF 73, 40.8900°N 25.5560°E, leg. Subai 25.07.1990, S 15639/13;
mountain at sea shore 6.8 km W of Díkella, on limestone rocks, approx. 30 m alt.,
UTM LF 82, 40.8631°N 25.6399°E, leg. Subai 26.07.1990, S 15640/1. — Thássos
Island, N of Skála Marió, UTM KF 90, 40.64°N 24.51°E, leg. Kiss & Pintér 6.7.1986,
HNHM 29881/4 + 1 (juv.); narrow (partim) of a valley 2.8 km NE of Theológos, on
marble rocks, 530 m alt., UTM LF 00, 40.6696°N 24.7329°E, leg. Subai 25.07.1990, S
15932/1; Thássos Liménas, amphitheater, UTM LF 01, 40.7818°N 24.7174°E, leg. Neute-
boom 13.7.1976, NNM/4 (alk.); between Thássos Liménas and Pahí (= 5 km W of
Thássos Liménas), UTM LF 01, 40.787°N 24.66°E, leg. Neuteboom 13.7.1976, NNM/1.
Erroneous or imprecise localities: "Albanien" UTM n. id., 284847/2, 284869/1;
"Rumälien" UTM n. id., HNHM 49547/1+1 (juv.), SMF 284868/3.

64 Peter Subai & Eike Neub ert
Remarks: According to Frivaldszky (1835: 274), this species was collected
"at low hill slopes in the mountains of the Balkan". In his introduction Frivald-
szky explains that the specimens were collected by colleagues from the sur-
roundings of "Szlivno" (= Sliven, Bulgaria). Today, there is only a single speci-
men left from the collections of Frivaldszky, i.e. SMF 7047/1 (in coll. Ross-
mässler), which is here considered a syntype. This specimen is small, rounded,
and has a weak denticle on the upper part of the lip. The label states "Alban-
ien", but according to Zilch (pers. comm..), this label was written by Kobelt.
Three other specimens in SMF (SMF 284847/2 and 284869/1) are also said to
originate from Albania but stem from Stenz, and thus do not represent type
material.
Lindholmiola girva shows only a small degree of shell variability. There are
two forms, which occur in parallel in many populations and are always con-
nected by intergrading specimens. The first form has rounded small shells
with a narrow umbilicus, while the other form is more flattened with a slightly
larger umbilicus.
Distribution (Fig 29): Lindholmiola gir va can be found in quite a large area
in the northeastern Balkan Peninsula. It can be found in southern Moldavia,
southern and southeastern Rumania and it is widespread in Bulgaria and north-
ern Greece. It has also been recorded from the valley of the Vardar in Fyrom.
Lindholmiola gyria (Roth, 1839) Figs 33–34, 35, 44
1839 Helix gyria Roth, Molluscorum species, quas in itinere per Orientem facto
comites clariss. Schubert doctores M. Erdl et J. R. Roth collegerunt: 16, Taf. 1
Fig. 17–18.
1846 Helix gyria, – L. Pfeiffer, in: Martini & Chemnitz: Systematisches Conchylien-
Cabinet, 1. 12 (II): 283, Taf. 124 Fig. 31–33.
1848 Helix gyria, – L. Pfeiffer, Monographia heliceorum viventium, 1: 414.
1853 Helix gyria, – L. Pfeiffer, Monographia heliceorum viventium, 3: 261.
1868 Helix gyria, – L. Pfeiffer, Monographia heliceorum viventium, 4: 312.
1859 Helix gyria, – Mousson, Vierteljahrsschrift der Naturforschenden Gesell-
schaft in Zürich, 4: 259.
1868 Helix gyria, – L. Pfeiffer, Monographia heliceorum viventium, 5: 415.
1876 Helix gyria, – L. Pfeiffer, Monographia heliceorum viventium, 7: 466.
1887 Helix gyria, – Tryon, Manual of Conchology, 3: 177.
1889 Helix (Caracolina) gyria, – Westerlund, Fauna, 2: 20.
1894 Helix (Caracollina) gyria, – Pilsbry, in Tryon, Manual of Conchology, 9: 228.

65
Revision of the genus Lindholmiol a Hesse, 1931
1898 Helix (Gonostoma) gyria, – Kobelt, in Rossmässler, E. A.: Iconographie
der Land- & Süsswasser-Mollusken mit vorzüglicher Berücksichtigung der
europäischen noch nicht abgebildeten Arten, (2) 8 (1/4): 53, Taf. 226 Fig.
1447.
1918 Caracollina gyria, – Hesse, Nachrichtsblatt der Deutschen Malakozoolo-
gischen Gesellschaft, 50: 109.
1991 Helicodonta gyria gyria, – Maassen, Basteria 55: 123, Fig. 1–3.
Diagnosis: shell strongly depressed, protoconch and first teleoconch whorls
slightly immersed, whorls densely coiled, extremely long bursa copulatrix.
Description of shell: shell strongly depressed, protoconch and first teleo-
conch whorls slightly immersed, basic shell colour brown; initial part of pro-
toconch whorls smooth, followed by 1–1¼ whorls with a pattern of a few fine
Figs 33–34. Lindholmiola gyria. Fig. 33: syntype Helix gyria, ZSM 1781, Turkey, "Cacamo Cariae",
D = 14.9 mm; Fig. 34: NMBE 515613, Turkey, Vilayet Antalya, Phaselis, ruins close to Tekirova, D =
13.68 mm. — All photos Neuber t/Bochud, × 3.

66 Peter Subai & Eike Neub ert
radial stripes; the following teleoconch whorls with increasingly stronger
radial stripes, but almost lacking on the last whorls; surface of the last whorl
covered by a fine granulation; sometimes with hairs, hair warts can be found
in the granulated surface sculpture; 6¼–7¼, densely coiled and well round-
ed whorls, last whorl slightly descending before the aperture; suture deep;
umbilicus broad, almost cylindrical, reaching a diameter of 2.8–3.3 mm; aper-
ture oblique, slightly bent in lateral view, and strongly ear-like in frontal view;
apertural insertion with a distance of 4.2–5.3 mm, connected by a very thin,
almost indiscernible callus; peristomial rim ± sharp, laterally with a strong
swelling, basally reinforced with a small umbilical shield.
Measurements: H: 5.3–6.4; D: 12.3–15; aH: 4.4–5.5 (5.6–6.6); aW: 5.8–6.9.
Details of body (after one specimen from Phaselis): head and dorsum dark
grey, flanks and tail light grey; sole cream with a greyish rim; mantle collar
translucent with grey spots of pigment; secondary ureter opens 1.5 mm apart
from the respiratory pore.
Morphology of the genital organs (Fig. 44): penis relatively long, slightly
enlarged in its central part, penial glandular tissue covering ca 1/4 of the
total penis length, penial lumen filled by a single, broad and serrated pilas-
ter, starting at the distal penial wall and stretching to the narrow penial pore,
where it recurves and ends centrally in the penis, remaining penial lumen with
short perpendicular folds; vagina relatively long, filled with a long, narrow
pilaster connecting to one of the atrial pilasters, glandula with a short stalk
and a strongly folded part, bursa copulatrix three times the length of the glan-
dula, bursa copulatrix narrow, same length as glandula, pedunculus reaching
ca 40% of the total length of the bursa copulatrix, vesicle of bursa copulatrix
elongate, double the diameter of the pedunculus; atrial lumen with several
broad and short interdigitating pilasters.
Differential diagnosis: The shell of L. gyria with its broad last whorl and
the slightly concave spire resembles that of Helicodonta obvoluta, and thus
cannot be confused with any of the other species of the genus Lindholmiola.
The extremely long bursa copulatrix differentiates it from all its congeners.
Type specimens: Turkey, Antalya County, Kekova Island, "Cacamo Cariae", 36.17°N
29.87°E, syntype gyria: ZSM 1781.
Additional specimens examined:
Turkey, Antalya County, Kekova Island (= Cacamo Cariae, 20 km E of Kaş), UTM QA
50/60, 36.17°N 29.87°E, (Roth, 1839: 16, Maassen, 1991: 124); rocks NW of lake near
Beymelek (= 9 km as the crow flies WSW of Finike), UTM TF 31, 36.2726°N 30.0456°E,
leg. Welter-Schultes 27.9.1998, coll. Welter-Schultes/1; Karamanbeyli pass near Finike
(= Phinika Pass), UTM TF 43, 36.54°N 29.98°E, SMF 284853/1; forest in gorge of

67
Revision of the genus Lindholmiol a Hesse, 1931
Kemer creek (= W of Kemer), UTM TF 75, 36.5943°N 30.5049°E, (Maassen, 1991: 124);
Phaselis, ruins near Tekirova (= 10 km S of Kemer), UTM TF 84, 36.4704°N 30.5131°E,
(Maassen, 1991: 124); leg. Subai 26.2.1999, S 16083/17 + 7 (juv.) + 1 (alk.).
Remarks: This species is probably endangered, because even empty shells
are very rare. In February 1999, it took several hours until the only hitherto
known living specimen could be found close to Phaselis. The anatomical inves-
tigation revealed that it belongs to Lindholmiola and not to Helicodonta. In
1991 (p. 124, Fig. 4–6), Maassen described Helicodonta gyria wilhelminae from
Crete, which is considered an endemic Helicodonta species.
Distribution (Fig 35): This species is only known from a small coastal stripe
in the south-western part of the vilayet Antalya. The distribution range covers
the Island Kekova (= E of Kaş) and the surroundings of Kemer to Phaselis.
Lindholmiola pirinensis Jaeckel, 1954 Figs 26, 36–38, 39
1954 Lindholmiola contorta pirinensis Jaeckel, Mitteilungen aus dem Zoologi-
schen Museum in Berlin, 30 (1): 86, Abb. 22.
1957 Lindholmiola corcyrensis pirinensis, – Jaeckel, Klemm & Meise, Abhand-
lungen und Berichte aus dem Staatlichen Museum für Tierkunde in Dresden,
23 (2): 188.
Fig. 35. Distribution of Lindholmiola gyria.

68 Peter Subai & Eike Neub ert
1962 Lindholmiola corcyrensis pirinensis, – Klemm, Sitzungsberichte der Öster-
reichischen Akademie der Wissenschaften, Mathematisch-Naturwissenschaft-
liche Klasse, Abt. I, 171 (6/7): 253.
1968 Lindholmiola corcyrensis, – L. Pintér, Malakologische Abhandlungen aus
dem Staatlichen Museum für Tierkunde Dresden, 2 (15): 226, Abb. 4, Abb.
32–34 [non corcyrensis Rossmässler] .
1973 Lindholmiola (Lindholmiola) corcyrensis pirinensis, – Hudec & Vašátko,
Acta scientiarum naturalium Academiae Scientiarum Bohemoslovacae, 7 (9):
13, Abb. 10 right side Taf. 3 Fig. 1.
1988 Lindholmiola corcyrensis pirinensis, – Reischütz, Annalen des Naturhisto-
rischen Museums in Wien, 90 (B): 347.
Diagnosis: shell depressed, usually no granulation, whorls with a blunt
shoulder, distal penial lumen with two equally sized strong pilasters.
Description of shell: shell depressed, with a slightly raised spire, basic col-
our light to chestnut brown; initial half of the protoconch whorl smooth, then
with a very fine and dense radial sculpture; teleoconch surface with strong radi-
al riblets, becoming very weak on the penultimate and ultimate whorl, a gran-
ulation usually lacking; yellowish hairs usually eroded (length 0.47–1.4 mm,
shorter in the umbilicus), but if present covering the whole shell except a
small, hair-free zone around the shells periphery, where only randomly some
small hair knuckles can be found; shell with 5¼–6¾ regularly increasing and
well rounded whorls, with a blunt shoulder, last whorl descending shortly
before the aperture; suture of medium depth; umbilicus almost cylindrical at
its beginning, suddenly increasing in size in the last whorl, diameter 1.55–2.8
mm; aperture oblique, slightly curved in lateral view; in frontal view obliquely
oval, peristomial insertions with a gap of 2.65–5.9 mm; parietal callus very
thin, often missing; peristome sharp, strongly broadened laterally, slightly
reflected basally; oblique columellar reflection, but only covering ca 1/10 of
its diameter.
Measurements: H: 4.7–7.5; D: 9.2–16.4; aH: 4.2–7.5(2.65–5); aW: 4.5–8.
Details of body (according to 10 specimens (3 km from the crossing Vóla-
kas/6; 3.2 km N of Makriplágio/4; Hudec & Vašátko, 1973: 13, Abb. 10): head
and neck greyish to brown-greyish, flanks and tail cream; sole cream with dark-
er rims; mantle rim with a 1 mm broad stripe without pigmentation, rest of
mantle for 1½ whorl with a pattern of large grey to brown spots, subsuturally
with a broad brown stripe; secondary ureter opens 1 mm before pneumostome,
split into two 1 to 1.5 mm long canals.
Morphology of the genital organs (Fig. 26): penis relatively long, slightly to
well thickened, penial glandular tissue covers ca 1/3 of the total penis length,

69
Revision of the genus Lindholmiol a Hesse, 1931
distal penial lumen with two equally sized strong pilasters, one of them with
proximal warts and a rounded, thickened distal end, the other pilaster is in
parallel, but recurving and leading back to unite with the warty end of the
first pilaster; vagina short to relatively long, lumen filled with a long and thick
pilaster which splits distally into two branches (sometimes with additional
Figs 36 –38. Lindholmiola pirinensis. Fig. 36: holotype Lindholmiola contorta pirinensis, ZMB 101257,
Bulgaria, Banderica Valley, D = 14.2 mm; Fig. 37: NMBE 515637, Greece, Thráki, Canyon 2 km N of
Xánthi (flat form with wide umbilicus, arid areas), D = 10.8 mm; Fig. 38: NMBE 515638, Greece,
Makedonía, beech forest 3.2 km N Makriplágio (large form of wooded areas), D = 14.5 mm. — All
photos Neubert/Bochud, × 3.

70 Peter Subai & Eike Neub ert
parallel folds); glandula with a long stalk and a strongly folded part, bursa
copulatrix as long as or sometimes longer than the glandula, pedunculus con-
stituting ca half of the total length of the bursa copulatrix, vesicle of bursa
copulatrix not observably differentiated; atrial lumen with a large serrated
area, and with a central pilaster that connects to the vaginal pilaster.
Differential diagnosis: In its general habit, L. pirinensis is quite close to
L. corcyrensis, but it differs from this species by its well rounded whorls and
the deeper suture. In L. corcyrensis, the teleoconch surface sculpture is coars-
er, with stronger irregular radial stripes. The hairs are usually permanent in
L. pirinensis, but on average shorter. The descending part of the last whorl
is bright to almost whitish, and the apertural rim usually more strongly rein-
forced with a small columellar shield. The penial pilasters are v-shaped wrin-
kles but longitudinal pilasters in L. pirinensis.
On average, shells of L. girva are smaller than those of L. pirinensis, with
the first species having a coarser teleoconch surface sculpture, with stronger
irregular radial stripes. The hairs in L. pirinensis are usually permanent but
shorter than those of L. girva. In L. pirinensis, the descending part of the last
whorl is bright to almost whitish, and the apertural rim usually more strongly
reinforced. In L. gir va, the penial pilasters never recurve like in L. pirinensis.
Type specimens: Bulgaria, Pirin Mountains, Banderica Valley, 1800–2000 m alt.,
UTM GM 02, 41.76°N 23.41°E, leg. Rensch, holotype pirinensis: ZMB 101257, paratype
pirinensis: ZMB 101258/5 + 2 (juv.); ex Knipper, ex coll. Jaeckel, paratype pirinensis:
SMF 153359/2.
Additional specimens examined:
Fyrom, Katlanovo, UTM EM 53, 41.89°N 21.68°E, leg. Kiss & Pintér 4.7.1985, HNHM
30857/1 (+ 5 girva).
Bulgaria, Kostinbrod near Sofija, UTM FN 84, 42.81°N 23.22°E, leg. Mészáros
9.1981, HNHM 13871/1 (+ 1 juv. girva); [Gara] Lakatnik, UTM FN 97, 43.08°N 23.38°E,
leg. Pintér 13.7.1967, HNHM 33699/38 + 2 (juv.); leg. Erőss 24.8.1976, S, 18264/1;
between Belogradčik and Orešec, UTM FP 33, 43.65°N 22.70°E, leg. Kiss & Pintér,
18.7.1984, HNHM 37470/1; Pirin Mountains, Banderica Valley, near Banderica
hut, UTM GM 02, 41.78°N 23.44°E, leg. Clauß, Maa/2; leg. Pintér 14.7.1974, HNHM
49535/1; Plovdiv (= Philippopel), hill "Schachti Tepe", UTM n. det., leg. Matsche-
va, SMF 284839/3; surroundings of Bačkovski monastery, UTM LG 24, 41.9426°N
24.8491°E, leg. Pintér 22.7.1967, S 3839/3; leg. Pintér 10.7.1973, HNHM 49546/1 + 3
(juv.); leg. Pintér 13.8.1981, HNHM 33696/3; 2 km NW of Malko Tarnovo (= bifurcation
to Stoilovo), UTM NG 44, 42.0071°N 27.5044°E, leg. Erőss 29.7.1997, coll. Erőss/3;
Ropotamo River right bank, UTM NG 67, 42.32°N 27.74°E, leg. Szigethy 4.7.1973,
HNHM 49530/1; Zdravec im Rodopi Mountains, UTM n. det., leg. Uminski, 19.9.1957,
HNHM 40344/1.

71
Revision of the genus Lindholmiol a Hesse, 1931
Greece, Thráki, 1 km W of Neohóri, UTM LF 06, 41.22°N 24.60°E, leg. Kiss & Pintér
5.7.1986, HNHM 29883/1 (juv.); 2 km N of Xánthi (= direction to Stavroúpoli), on lime-
stone rocksW of the highway, 140 m alt., UTM LF 25, 41.1591°N 24.8806°E, leg. Subai
27.07.1990, S 15515/2; 2.1 km from the highway Xánthi–Stavroúpoli direction Ehīnos,
on marble rocks, 220 m alt., UTM LF 26, 41.2085°N 24.8612°E, leg. Subai 24.05.1997,
S 15518/4; forest border 700 m from Melivoia Village direction Kotili, crystalline rocks,
415 m alt., UTM LF 27, 41.3328°N 24.9218°E, leg. Subai 24.05.1997, S 15516/3; forest
border 1.8 km from Melivoia Village direction Kotili, in leaf litter, crystalline rocks, 450
m alt., UTM LF 27, 41.3354°N 24.9063°E, leg. Subai 24.05.1997, S 15517/6; 6.3 km from
Amaxádes direction Íasmos (= 1 km from Kopteró direction Íasmos), on limestone rocks
approx. 100–150 m N of the highway, 60 m alt., UTM LF 45, 41.1204°N 25.1406°E, leg.
Subai 27.07.1990, S 15638/1; 2.6 km from Mákri direction Komotiní (on the old high-
way), mountain with cave beside new highway, on limestone rocks, 100 m alt., UTM LF
92, 40.8758°N 25.7095°E, leg. Subai 26.07.1990, S 15635/2; gorge at NW border of Avas
(= 13 km N of Alexandroúpoli), on limestone rocks, 100 m alt., UTM MF 02, 40.9366°N
25.9129°E, leg. Subai 26.07.1990, S 15636/2. — Makedonía, 2.9 km from Ahladia direc-
tion Káto Nevrokópi, no lime stone, 690 m alt., UTM GL 48, 41.3885°N 23.9748°E, leg.
Subai 22.05.1997, S 15525/21 + 1 (alk.); Falakró Mountains, 3 km from the bifurcation
Vólakas direction skiing center, deciduous forest with limestone rocks, 1200 m alt.,
UTM KF 57, 41.2952°N 24.0091°E, leg. Subai 08.05.1995, S 15514/8 + 6 (alk.); upper
border of gorge 1 km from Mikroklissoúra direction Potami, on limestone rocks, 400 m
alt., UTM KF 58, 41.3867°N 24.0698°E, leg. Subai 22.05.1997, S 15526/9 + 3 (juv.); oak
forest 12.9 km from Sidirónero direction Livaderó, in leaf litter, crystalline rocks, 600 m
alt., UTM KF 67, 41.2740°N 24.2088°E, leg. Subai 22.05.1997, S 15524/2 + 1 (juv.) + 1
(alk.); mountain slope at NE border of Sidirónero, river valley, among herbs and under
stones, 650 m alt., UTM KF 68, 41.3686°N 24.2339°E, leg. Subai 22.05.1997, S 15522/3;
beech forest 3.2 km N of Makriplágio (along the highway), under stones, in leaf litter,
crystalline rocks, 900 m alt., UTM KF 76, 41.2350°N 24.2945°E, leg. Subai 8.5.1995, S
15513/8 + 2 (alk.); leg. Subai 22.5.1997, S 15523/6 + 4 (alk.); on low marble rocks at Dis-
vato Village, under stones and in leaf litter, 450 m alt., UTM KF 95, 41.0681°N 24.5713°E,
leg. Subai 26.05.1997, S 15521/2; mountain E of Makrihóri, beside a marble quarry, 650
m alt., UTM LF 04, 41.0692°N 24.6298°E, leg. Subai 07.05.1995, S 15511/1; small river
gorge 4.1 km from Kehrókambos direction Makrihóri, stone hedge, under stones, 485 m
alt., UTM LF 05, 41.1292°N 24.6448°E, leg. Subai 26.05.1997, S 15520/2; in forest at W
border of Stavroúpoli, Nestos river W bank, on limestone rocks, 120 m alt., UTM LF 06,
41.2031°N 24.7056°E, leg. Subai 7.5.1995, S 15512/11; leg. Subai 24.5.1997, S 15519/5
+ 1 (juv.); NE border of Parádissos, on limestone rocks, 60 m alt., UTM LF 15, 41.0843°N
24.7612°E, leg. Subai 25.05.1997, S 15634/1.
Remarks: Specimens from more humid forested areas tend to have larger
shells with a higher spire than those from more arid habitats, where the shells
tend to be flatter and have a larger umbilicus. Next to populations with a wide
range in shell size, there are also populations which contain excusively large
specimens. However, these are considered conspecific, because they comply
with all other character states of the other populations.

72 Peter Subai & Eike Neub ert
Distribution (Fig. 39): Lindholmiola pirinensis is well known from the moun-
tains in western and southern Bulgaria and in the northern part of Greece
north to the area between Kavála–Dráma and in Thracia. The westernmost
record is from Fyrom in the Vardar valley close to Katlanovo
Lindholmiola regisborisi (A. J. Wagner, 1927) Figs 40–42, 43, 46
1927 Caracollina regis-borisi A. J. Wagner, Annales zoologici Musei Polonici
Historiae Naturalis, 6 (4): 364, Taf., 19 Fig. 166–168.
1957 (Lindholmiola ?) regisborisi, – Jaeckel, Klemm & Meise, Abhandlungen und
Berichte aus dem Staatlichen Museum für Tierkunde in Dresden, 23 (2): 165.
1988 Lindholmiola regisborisi, – Reischütz, Annalen des Naturhistorischen
Museums in Wien, 90 (B): 347.
1990 Lindholmiola regisborisi, – Fechter & Falkner, Steinbachs Naturführer: 222.
Diagnosis: shell planispiral, strongly haired, umbilicus dish-like, perspec-
tive, long bursa copulatrix.
Fig. 39. Distribution of Lindholmiola pirinensis.

73
Revision of the genus Lindholmiol a Hesse, 1931
Description of shell: shell almost planispiral, basic colour light brown to
yellowish-brown; initial half of the protoconch whorl smooth, then with a very
fine and dense radial sculpture; teleoconch surface with strong radial riblets
of different size producing an undulating sculptural pattern (magnification
required), whole shell with a dense granulation, ventral side of the shell with
fine radial riblets and granulation; shells strongly haired, with a hair-free sub-
sutural zone followed by a zone of permanent hairs around the blunt periph-
ery of the shell (hairs in this area 0.8–2.3 mm long), ventral side of the shell
usually without hairs, the umbilicus with very short bristles; shell with 5¾–7
regularly increasing and well rounded whorls, with a blunt shoulder; suture
very deep; umbilicus dish-like, perspective, diameter 2.4–4.2 mm; aperture
very oblique, slightly curved in lateral view; in frontal view subquadrate to
obliquely oval, strongly cut out on the left side, peristomial insertions with
a gap of 2.6–4.1 mm; parietal callus very thin, undulating; peristome sharp,
strongly broadened apically and laterally, slightly reflected basally; oblique
columellar reflection, but only covering ca 1/10 of its diameter.
Measurements: H: 5–6.25; D: 12.5–16.5; aH: 4.85–6.4 (2–3.1); aW: 5.9–8.
Details of body (according to 8 animals from 400 m S of Kehrókambos):
body dark brown to blackish-greyish, head and eye-stalks grey with a dark-
brown tip; tale grey; sole cream with greyish lateral rims; mantle along the
lung vein with two stripes of dark-brown pigmentation of rounded spots; sec-
ondary ureter opens 0.5 –1.5 mm from the pneumostome, where it splits into
two canals.
Morphology of the genital organs (Fig. 46): penis relatively short, slightly
thickened, penial glandular tissue covers ca 1/3–1/2 of the total penis length,
penial lumen with two equally sized, serrated pilasters, starting at the distal
penial wall and stretching to the penial pore, sometimes with short perpen-
dicular folds between the pilasters; vagina very short, somewhat broadened;
glandula narrow with a short stalk and a long and scarcely folded part; bursa
copulatrix reaching twice the length of the glandula with a pedunculus occu-
pying half of the total length, vesicle strongly inflated; atrial lumen with a large,
broad and flat, serrated pilaster, connecting to the vaginal lumen and reach-
ing the proximal end of the vagina; opposite side of the atrial lumen with a
semi-circular, keeled fold.
Differential diagnosis: Due to the large planispiral shell with the shoul-
dered whorls, the long hairs and the large dish-like umbilicus, this species
cannot be confused with any other species of Lindholmiola. Other differentiat-
ing features are the unusually long bursa copulatrix with its clearly separated
vesicle, and the specific formation of penial and atrial pilasters.

74 Peter Subai & Eike Neub ert
Type specimens: Greece, Thráki, Xánthi, UTM LF 25, 41.14°N 24.89°E, syntypes:
NMNH 3167/4.
Additional specimens examined:
Greece, Makedonía, limestone mountain approx. 2.5 km NNE of Zarkadía (along
the road), 450 m alt., UTM LF 04, 41.0288°N 24.6417°E, leg. Fauer, 18.08.1979, S
12475/2; leg. Kiss & Pintér 7.7.1986, HNHM 29888/6 + 4 (juv.); leg. Fauer & Subai
22.09.1989, S 13977/58 ; leg. Subai 7.5.1995, S 14082/16; mountain E of Makri-
hóri, beside a marble quarry, 650 m alt., UTM LF 04, 41.0692°N 24.6298 °E, leg.
Subai 07.05.1995, S 14083/4; stone hedge in valley of two mountains 1 km from
Ágios Kosmás direction Makrihóri, 380 m alt., UTM LF 04, 41.0783°N 24.6536°E,
leg. Subai 26.05.19 97, S 15472/17; valley 2.5 km E of Makrihóri (= direction
Kehrókambos), beside village ruins, on limestone rocks, 180 m alt., UTM LF 04 ,
41.046 6°N 24.6462°E, leg. Subai 26.05.1997, S 15473/16; small river gorge 4.1
km from Kehrókambos direction Makrihóri, stone hedge, under stones, 485 m alt.,
UTM LF 05, 41.1292°N 24.6448°E, leg. Subai 26.05.1997, S 15474/1; 7.2 km from
Kehrókambos direction Makrihóri, at crossing to Skopós, under stones, 550 m
alt., UTM LF 05, 41.1043°N 24.6392°E, leg. Subai 26.05.1997, S 15475/3; 400 m S
of Kehrókambos, under (lime) stones, 390 m alt., UTM LF 05, 41.1514°N 24.6454°E,
leg. Subai 07.05.1995, S 1408 6/13 + 14 (alk.); 4 km from Kehrókambos direction
Stavroúpoli, on low limestone rocks, approx., 180 m alt., UTM LF 06, 41.1806°N
24.6687°E, leg. Subai 07.05.1995, S 14085/1; limestone mountain W of Stavroúpo-
li (= at the opposite bank of Néstos river), in forest, on limestone rocks, 120 m alt.,
UTM LF 06, 41.2031°N 24.7057°E, leg. Subai 27.07.1990, Maa/3, S 14059/35; leg.
Subai 24.05.1997, S 15471/25 + 5 (juv.); Parádissos Village, UTM LF 15, 41.08°N
24.76°E, leg. Kiss & Pintér 7.7.1986, HNHM 29889/12 + 2 (juv.); leg. Maassen 7.1980,
Maa/11 + 9 ( juv); NE border of Parádissos, on limestone rocks, 6 0 m alt., UTM LF 15,
41.0843°N 24.7612°E, leg. Subai 25.05.1997, S 15476/11 + 1 (juv.). — Thráki, 2 km
E of Stavroúpoli, UTM LF 06, 41.21°N 24.73°E, leg. Kiss & Pintér 5.7.1986, HNHM
29887/1 + 4 ( juv.); gorge near Xánthi, UTM LF 25, 41.14°N 24.88°E, leg. Schütt
2.5.1962, HNHM 39235/3, SMF 279328/4; leg. Maassen 7.1980, Maa/18 + 8 (juv.); 2
km N of Xánthi (= direction to Stavroúpoli), on limestone rocksW of the highway, 140
m alt., UTM LF 25, 41.1591°N 24.8806°E, leg. Subai 27.07.1990, S 14058/43.
Turkey, Marmara, Marmara Adasi, 40.59°N 27.56°E, 04.07.1976, leg. Oberling &
Gerber, NMBE 515643/2; ditto, 0 8.08.1977, leg. Oberling & Gerber, NMBE 515642/1,
NMBE 515633/8; Kaleh, Marmara Adasi, 40.62°N 27.62°E, 08.0 8.1977, leg. Oberling
& Gerber, NMBE 515630/20.
Remarks: The shells of L. regisborisi are not variable, there are no local
forms of this species known so far.
Figs 40–42. Lindholmiola regisborisi. Fig. 40 : syntype Caracollina regis-borisi, NMNH (Sofia) 3167/4,
Greece, Thrakia, Xanthi, D = 17.4 mm; Fig. 41: NMBE 515619, Greece, Makedonía, 400 m S of Kehrókam-
bos, D = 15.01 mm; Fig. 42: NMBE 515630, Turkey, Marmara Adasi, Kaleh, D = 14.78 mm. — All photos
Neubert/Bochud, × 3.

75
Revision of the genus Lindholmiol a Hesse, 1931

76 Peter Subai & Eike Neub ert
Distribution (Fig. 43): This species is known to live in a small area in the
Lekánis Mountains in Greece, west of the Néstos River, in the lower Néstos
valley between Stavroúpoli and Parádissos and east of the Néstos River to
Xánthi in Thráki. The record from Marmara Adasi is isolated from the Greek
distribution area, and is probably the result of an introduction due to anthro-
pogenic activities. This is the first record of this species for Turkey.
Lindholmiola reischuetzi Falkner, 1995 Figs 28, 47–50, 51
1907 Helix (Gonostoma ) lens var. insularis O. Boettger, Nachrichtsblatt der
Deutschen Malakozoologischen Gesellschaft, 39 (1): 34 [several times
preoccupied].
1918 Caracollina lens insularis, – Hesse, Nachrichtsblatt der Deutschen Malako-
zoologischen Gesellschaft, 50 : 109.
1960 Lindholmiola lens insularis, – Urbanski, Bulletin de l'Institut Zoologique
de l'Académie Bulgare des Sciences, 9: 91. Taf. 4 Fig. 4 a–c, 5 a–c.
1983 Lindholmiola lens insularis, – Reischütz, Annalen des Naturhistorischen
Museums in Wien, 85 (B): 135.
1988 Lindholmiola lens insularis, – Reischütz, Annalen des Naturhistorischen
Museums in Wien, 90 (B): 347.
1995 Lindholmiola reischuetzi Falkner, Heldia, 2 (3/4): 97 (nom. nov. pro Helix
(Gonostoma) lens var. insularis O. Boettger, 1907).
Fig. 43. Distribution of Lindholmiola regisborisi.

77
Revision of the genus Lindholmiol a Hesse, 1931
Figs 44 –4 6. Genital organs of Lindholmiola species. Fig. 44 : Lindholmiola gyria, a: situs, b: lumina of
genital organs, Turkey, Vilayet Antalya, Phaselis; Fig. 45: Lindholmiola barbata, a: situs, b: lumina
of genital organs, Greece, Island Kríti, Nomos Haniá, Soúda Bay ; Fig. 46: Lindholmiola regisborisi, a :
situs, b: lumina of genital organs, Greece, Makedonía, 400 m S of Kehrókambos.

78 Peter Subai & Eike Neub ert
Diagnosis: shell flat, lentiform, usually not haired, with a blunt keel, parie-
tum with a very thick callus, vagina completely reduced,
Description of shell: shell flat, lentiform, basic colour light brown to yel-
lowish-brown; initial 1–1½ of the teleoconch whorls smooth, then with a fine
and dense radial sculpture, becoming stronger on the subsequent whorls,
developing to small riblets; ventral side of the shell usually with fine radial
stripes only, sometimes with small riblets; whole shell with a dense granu-
lation; shell usually not haired, but if so, then restricted to a narrow stripe
just above the blunt peripheral keel (hairs 0.5–0.8 mm, yellowish); shell
with 5½–7 regularly increasing and flat whorls, with a blunt keel; shells tend
to produce scalarid forms (see remarks); suture shallow; last whorl slightly
ascending, but descending before the aperture with a white coloured area;
umbilicus dish-like, perspective, diameter 1.7–3.3 mm; aperture very oblique,
slightly curved in lateral view; in frontal view of semi-circle form, peristomial
insertions with a gap of 2–3.1 mm, parietal callus a very thick, bar-like struc-
ture; peristome sharp, broadened apically and laterally, often slightly reflect-
ed basally.
Measurements (in mm): H: 3.1–4.7; D: 9.1–13.3; aH: 2.5–4.7(1.4–2.8);
aW: 3.4–5.3.
Details of body (according to 6 animals from 4 km of Kehrókambos in direc-
tion of Stavroúpoli): head and dorsum dark brown to grey; sole cream with
brownish lateral rims; mantle above the secondary ureter with one stripe of
dark-brown pigmentation; secondary ureter opens 0.5–1 mm from the pneu-
mostome.
Morphology of the genital organs (Fig. 28): penis relatively short, slightly
thickened, penial glandular tissue covers only 1/4 of total penis length, penial
lumen with two large pilasters with one of them starting at the proximal
penial wall and stretching to the penial pore, the second branching into two
arms before reaching the proximal penial wall, several small, perpendicularly
arranged accessory folds present; vagina completely reduced, glandula and
bursa copulatrix branch off the atrium, glandula with a short stalk and a long
and scarcely folded part; bursa copulatrix reaching the length of the glandula
or somewhat shorter, vesicle almost not discernible; atrial lumen filled with a
large, broad and flat, pillow-like pilaster, connecting to a few short folds.
Differential diagnosis: On average, L. spectabilis is larger than L. reischuetzi.
Equally sized shells can always been identified as L. spectabilis by its larger
aperture, the weak palatal callus, and the usually narrower umbilicus. The
reduced vagina is a clear autapomorphic character that separates L. reischuetzi
from all other species of Lindholmiola.

79
Revision of the genus Lindholmiol a Hesse, 1931
Around the Bay of Korinthos, there is a strongly keeled form of L. lens (callo-
juncta), which somewhat resembles L. reischuetzi. This form of L. lens, how-
ever, has a larger shell, and its palatal callus is only slightly reinforced. Other
forms of L. lens have a more raised spire and always a weak palatal callus.
Figs 47–50. Lindholmiola reischuetzi. Fig. 47: syntype Helix (Gonostoma) lens var. insularis, SMF
7048/1, Greece, Island Thássos, D = 9.4 mm ; Fig. 48: NMBE 515640, Greece, Makedonía, Halkidikí
Peninsula, close to the cave of Petrálona, D = 10.13 mm; Fig. 49: NMBE 515639, Greece, Thráki, Can-
yon 2 Km N of Xánthi (form with large umbilicus and broad whorls), D = 12.43 mm; Fig. 50: NMBE
515641, Greece, Makedonía, N boundary of Taxiárhis (scalarid form), D = 12.01 mm. — All photos
Neubert/Bochud, × 3.

80 Peter Subai & Eike Neub ert
Type specimens: Greece, Thássos Island, UTM n. det., ex Speidel, ex coll. O. Boett-
ger, syntypes insularis: SMF 7048/1, 7049/1, 101953/4.
Additional specimens examined:
Greece, Makedonía, Halkidikí Peninsula, 500 m N of Eleohória, on low lime-
stone rocks, 170 m alt., UTM FK 86, 40.3434°N 23.1613°E, leg. Subai 05.05.1995, S
15485/16; Halkidikí Peninsula, in valley beside the cave of Petrálona, 360 m alt., UTM
FK 87, 40.3759°N 23.1681°E, leg. Subai 05.05.1995, S 15486/5; leg. Subai 29.5.1997,
S 15487/179; N N border of Galipsós, on limestone rocks, approx. 200 m alt., UTM
GL 42, 40.8062°N 23.9502°E, leg. Subai 06.05.1995, S 15484/13 + 10 (alk.); moun-
tain slope N of Podohóri, on limestone rocks, 450 m alt., UTM KF 42, 40.8473°N
24.0249°E, leg. Subai 27.05.1997, S 15505/25; Simvolo Mountains, 1.2 km from Sidi-
rohóri direction Foliá (= W of Karavangélis), on limestone rocks, 250 m alt., UTM KF
52, 40.8380°N 24.1375°E, leg. Subai 27.05.1997, S 15510/62; Kokkinohóri, UTM KF 52,
40.81°N 23.99°E, Neuteboom 11.7.1976, NNM/2 + 3 (juv.) + 16 (alk.); mountain slope
4.5 km W of the bifurcation to Loutrá Eleftheró (= 48 km W of Kavála on the street
along the bank), on limestone rocks, 30 m alt., UTM KF 51, 40.7211°N 24.0601°E,
leg. Fauer & Subai 21.09.1989, S 15492/26; N border of Taxiárhis, under crystal-
line stones, 500 m alt., UTM KF 66, 41.2328°N 24.1888°E, leg. Subai 22.05.1997, S
15508/36 + 4 (alk.); 11 km above Kalífitos, UTM KF 66, 41.25°N 24.29°E, leg. Fauer
31.7.1971, S 15497/1; 2.6 km SW of Makriplágio, on limestone rocks, 520 m alt., UTM
KF 66, 41.2082°N 24.2519°E, leg. Subai 08.05.1995, Maa/7, S 15491/37 + 13 (alk.);
above the ruins of Philippi near Krinídes, on limestone rocks, 85 m alt., UTM KF 74,
41.0128°N 24.2851°E, leg. Neuteboom 12.7.1976, NNM/23 + 4 (juv.) + 14 (alk.); leg.
Maassen 7.1980, Maa/91 + 5 (juv); leg. Kiss & Pintér 7.7.1986, HNHM 29894/18 + 3
(juv.); leg. Subai 7.5.1995, S 15496/2; 5 km from Limnia direction Zigós, on low lime-
stone rocks and under stones, 730 m alt., KF 74, 41.0589°N 24.3816°E, leg. Subai
26.05.1997, Maa/12, S 15499/118; on marble rocks at N border of Ipsili Ráhi, approx.
500 m alt., UTM KF 76, 41.2123°N 24.3469°E, leg. Subai 23.05.1997, S 15507/31; 1
km E of Nikifóros, UTM KF 76, 41.17°N 24.33°E, leg. Kiss & Pintér 5.7.1986, HNHM
29911/104 + 55 (juv. + fragment); bridge 1.8 km E from Nikifóros, on limestone
rocks, approx. 235 m alt., UTM KF 76, 41.1726°N 24.3345°E, leg. Subai 23.05.1997, S
15506/10; in Makriplágio Village, beech forest and border of forest with stone walls,
under stones and in leaf litter, crystalline rocks, 880 m alt., UTM KF 76, 41.2267°N
24.2713°E, leg. Subai 22.05.1997, S 15509/4; 2 km E of Kavála, UTM KF 83, 40.95°N
24.45°E, leg. Maassen 7.1980, Maa/17 + 4 (juv); 2.5 km E of Kavála, hill slope with
small limestone rocks, 20 m alt., UTM KF 83, 40.9486°N 24.4438°E, leg. Fauer & Subai
22.09.1989, S 15488/9; 3 km W of Paleá Kavála, on limestone rocks above the river,
200 m alt., UTM KF 84, 41.0009°N 24.3934°E, leg. Subai 07.05.1995, S 15494/74; 2.5
km from Paleá Kavála direction Korifés, on low limestone rocks and under stones,
500 m alt., UTM KF 84, 41.0170°N 24.4388°E, leg. Subai 07.05.1995, S 15493/57 + 5
(alk.); S exposed slope 4.2 km from Paleá Kavála direction Halkeró, S exposed moun-
tain slope, on limestone rocks, 440 m alt., UTM KF 84, 40.9994°N 24.4273°E, leg.
Subai 26.05.1997, S 15502/38; on limestone rocks at SE border of Likóstomo, 740 m
alt., UTM KF 85, 41.1078°N 24.4424°E, leg. Subai 26.05.1997, S 15498/14; 800 m W
of Dipótamos, on limestone rocks beside highway, 690 m alt., UTM KF 95, 41.1196°N

81
Revision of the genus Lindholmiol a Hesse, 1931
24.5762°E, leg. Subai 26.05.1997, S 15503/14; limestone mountain approx. 2.5 km
NNE of Zarkadía (along the road), 450 m alt., UTM LF 04, 41.0288°N 24.6417°E, leg.
Kiss & Pintér 7.7.1986, HNHM 29890/153 + 39 (juv. + fragment); leg. Fauer & Subai
22.09.1989, Maa/9, S 15489/179; leg. Subai 7.5.1995, S 15490/24; valley 2.5 km E of
Makrihóri (= direction Kehrókambos), beside village ruins, on limestone rocks, 180
m alt., UTM LF 04, 41.0466°N 24.6462°E, leg. Subai 26.05.1997, S 15500/24; stone
hedge in valley of two mountains 1 km from Ágios Kosmás direction Makrihóri, 380 m
alt., UTM LF 04, 41.0783°N 24.6536°E, leg. Subai 26.05.1997, S 15501/9; 4 km from
Kehrókambos direction Stavroúpoli, on low limestone rocks, approx., 180 m alt.,
UTM LF 06, 41.1806°N 24.6687°E, leg. Subai 07.05.1995, S 15495/47 + 7 (alk.); lime-
stone mountain W of Stavroúpoli (= opposite bank of Néstos river), in forest, on lime-
stone rocks, 120 m alt., UTM LF 06, 41.2031°N 24.7057°E, leg. Subai 27.07.1990, S
15482/3; leg. Subai 24.5.1997, S 15483/24 + 1 (juv.); Parádissos Village, UTM LF 15,
41.08°N 24.76°E, leg. Kiss & Pintér 7.7.1986, HNHM 29896/19 + 2 (juv.); leg. Maas-
sen 7.1980, Maa/45 + 3 (juv); NE border of Parádissos, on limestone rocks, 60 m
alt., UTM LF 15, 41.0843°N 24.7612°E, leg. Subai 25.05.1997, S 15504/9 + 2 (juv.) + 1
(alk.). — Thráki, 2 km N of Xánthi (= direction to Stavroúpoli), on limestone rocksW
of the highway, 140 m alt., UTM LF 25, 41.1591°N 24.8806°E, leg. Subai 27.07.1990,
S 15481/2. — Thássopoula Island, UTM LF 02, 40.82°N 24.70°E, leg. Valkoura, 1842,
SMF 158896/5. — Thássos Island, olive grove at S border of Alikí, under flat stones,
UTM LE 09, 40.6050°N 24.7416°E, leg. Fauer & Subai 21.09.1989, S 15477/10 ; 1 km W
of Alikí, UTM LE 09, 40.60°N 24.72°E, leg. Kiss & Pintér 6.7.1986, HNHM 29915/1; 15
km from Kalívia direction Alikí, on limestone rocks, 110 m alt., UTM LE 09, 40.5914°N
24.6683°E, leg. Fauer & Subai 21.09.1989, S 15478/4; Theológos, UTM LF 00, 40.66°N
24.69°E, leg. Kiss & Pintér 6.7.1986, HNHM 29903/12 + 14 ( juv.); on limestone rocks
above cemetery of Theológos, 300 m alt., UTM LF 00, 40.6634°N 24.7022°E, leg. Fauer
& Subai 21.09.1989, S 15479/37; narrow (partim) of a valley 2.8 km NE of Theológos,
on marble rocks, 530 m alt., UTM LF 00, 40.6696°N 24.7329°E, leg. Subai 25.07.1990,
S 15480/29 ; W of Thássos Town, UTM LF 01, 40.77°N 24.68°E, leg. Kiss & Pintér
6.7.1986, HNHM 29919/1.
Remarks: Shell size variations in L. reischuetzi are few. However, next to
the widespread typical form of this species, there are two local forms that
need to be mentioned: 1) the populations east and northeast of Dráma to
the Lekánis Mountains (for example in Makriplágio, Taxiárhis, Ipsilí Ráhi,
Nikifóros etc.) tend to build scalarid shells. The whorls of this form are
stepped, and often the ventral side of the shells is covered by riblets; 2) the
easternmost populations of L. reischuetzi (for example close to Stavroúpoli
and N of Xánthi) have flat shells with rapidly increasing whorls. It has to be
stressed that these character states are restricted to those populations men-
tioned. However, on the Island Thássos these forms can be found mixed with
the widespread form with all intergrades. For this reason, these local forms
cannot be separated as geographical subspecies.

82 Peter Subai & Eike Neub ert
Distribution (Fig 51): This species is recorded from north-eastern Greece
from Makedonía to western Thráki, in the coastal stripe between the Dráma to
Xánthi and the sea. It also lives on the islands of Thássos and Thássopoula, and
there is a small isolated population in the Southwest of the Halkidikí Penin-
sula.
Lindholmiola spectabilis Urbanski, 1960 Figs 13, 15, 52–54, 55–56
1936 Lindholmiola lens, – Fuchs & Käufel, Archiv für Naturgeschichte, N. F. 5 (4):
651 [partim, only Angista canyon close to Drama) (non lens Férussac].
1960 Lindholmiola lens spectabilis Urbanski, Bulletin de l'Institut Zoologique
de l'Académie Bulgare des Sciences, 9: 91, Taf. 4 Fig. 3 a–c.
1988 Lindholmiola lens lens, – Reischütz, Annalen des Naturhistorischen Muse-
ums in Wien, 90 (B): 347 (non lens Férussac).
1990 Lindholmiola spectabilis, – Fechter & Falkner, Steinbachs Naturführer:
222, Abb. 4. and 7.
1995 Lindholmiola spectabilis, – Falkner, Heldia, 2 (3/4): 97.
Diagnosis: shell depressed, last whorl with a sharp, often whitish keel,
umbilicus perspective, shell with a dense granulation,
Fig. 51. Distribution of Lindholmiola reischuetzi.

83
Revision of the genus Lindholmiol a Hesse, 1931
Description of shell: shell depressed, basic colour light brown to yellowish-
brown; initial ½–1 of the teleoconch whorls smooth, the following whorls with
fine radial stripes, becoming stronger on the subsequent whorls and devel-
oping to small riblets, sometimes increasing on the last whorls to form ribs;
ventral side of the shell with fine undulating radial stripes only; whole shell
with a dense granulation, fine on the upper but coarser on the last whorls;
shell usually not haired, but if so, then restricted to the area just above the
blunt peripheral keel of the last whorl (hairs 0.6–0.8 mm); shell with 6–7¾
regularly increasing whorls, the upper 2–3 whorls slightly rounded, the subse-
quent whorls flat and slightly stepped; last whorl with a sharp, often whitish
keel; suture shallow; last whorl slightly ascending, but descending before the
aperture with a white coloured area; umbilicus perspective, diameter 1.56 –
3.6 mm (in some populations almost cylindrical; aperture oblique, strongly
curved in lateral view, slightly undulating basally; in frontal view of semi-cir-
cle form, peristomial insertions with a gap of 2.3–4.7 mm, parietal callus very
thin, almost indiscernible; peristome sharp, broadened laterally, often slightly
reflected basally, but covering the umbilicus only slightly.
Measurements: H: 3.75–6.7; D: 9.1–16.1; aH: 3.6–6.4 (1.7–3.4) aW: 4.1–7.65.
Details of body (according to 9 animals from 2.6 km SW of Makriplágio/6,
and Falakró Mts., Ágios Pnevma plateau/3): dorsum light to dark brown to
grey, flanks and tail usually lighter; sole very bright whitish to cream, with
greyish lateral rims; mantle along the lung vein with two stripes of dark-brown
pigmentation, sometimes dissolving in groups of spots; secondary ureter opens
1 mm from the pneumostome.
Morphology of the genital organs (Fig. 13): penis relatively short, slightly
thickened, penial glandular tissue covers only 1/3 of total penis length, penial
lumen with three large pilasters starting at the proximal penial wall with some
knot-like warts, the two marginal folds longer than the central one, distal penial
lumen to the penial pore without folds; vagina short, slightly thickened, filled
by a single pilaster stretching from the proximal end of the vagina through the
atrial lumen to the atrial pore; glandula with a short narrow stalk and a long
and scarcely folded part; bursa copulatrix reaching the length of the glandula
or somewhat shorter, vesicle well discernible.
Differential diagnosis: This species is very similar to some local forms of
L. lens, particularly to those with the more flattened whorls and the wide umbili-
cus, which live around the Bay of Korinthos and the north-eastern Pelopón-
nisos, both far away form the distribution area of L. spectabilis. Comparing
equally sized specimens of both species, the aperture of L. lens is always
broader and more oval, its lower border is flat and not undulating, the gap
between the peristomial insertions is larger, its umbilicus is more perspective,

84 Peter Subai & Eike Neub ert
Figs 52–54. Lindholmiola spectabilis. Fig. 52: paratype Lindholmiola lens spectabilis, SMF 158640,
Greece, Makedonía, (peninsula) Athos, monastery Kireta, D = 17.34 mm; Fig. 53: NMBE 515645, Greece,
Makedonía, mountain slope N of Podohóri, (tightly coiled shell, narrow umbilicus), D = 12.02 mm;
Fig. 54 : NMBE 515644, Greece, Makedonía, N boundary of Taxiárhis (small, strongly ribbed, slightly
scalarid form), D = 11.09 mm. — All photos Neuber t/Bochud, × 3.

85
Revision of the genus Lindholmiol a Hesse, 1931
its surface granulation is finer, and if there are hairs, the haired area is larg-
er. Additionally, L. lens has only two and not three penial pilasters. Another
quite interesting difference is the mating behaviour: While in L. lens, the mat-
ing partners stimulate each other with "rhythmic swaying lasting for hours,
tender palming with the eye-stalks and kiss-like touches with the mouth",
L. spectabilis is obviously more prosaic without "kisses and eye-stalk palming",
which turned out to constitute a mating barrier between the two species (Fech-
ter & Falkner, 1990: 222).
Often, L. spectabilis lives syntopic with L. reischuetzi. These two species can
easily be separated, because on average, shells of L. reischuetzi are smaller,
their aperture is narrower with the strong bar-like parietal callus, its umbilicus
is more perspective, its vagina is reduced, and it has only two penial pilasters.
Figs 55– 56. Lindholmiola spectabilis. Fig. 55: NMBE 515646, Greece, Makedonía, Kalithia in direction
to Anthohóri (coarsely ribbed, slightly scalarid form), D = 11.48 mm ; Fig. 56 : NMBE 515647, Greece,
Angítis Canyon, well at the Mara cave (large, wide umbilicus), D = 14.2 mm. — All photos Neubert/
Bochud, × 3.

86 Peter Subai & Eike Neub ert
Type specimens: Greece, Makedonía, Athos [Peninsula] , Kireta monastery, leg.
Papasov 29.6.1936, paratypes spectabilis: SMF 158640/2.
Additional specimens examined:
Greece, Makedonía, on conglomerate rocks above Sidirókastro, 110 m alt., UTM
GL 06, 41.2398°N 23.3954°E, leg. Subai 20.05.1997, S 15623/13 + 3 (alk.); castel of
Séres, under stones, UTM GL 15, 41.09°N 23.55°E, leg. Fauer 29.7.1971, S 12457/6;
N border of Inoússa, SSE exposed mountain slope, on limestone rocks, 200 m alt.,
UTM GL 25, 41.1059°N 23.6312°E, leg. Subai 21.05.1997, S 15617/3; on conglomer-
ate rocks above Panórama, 650 m alt., UTM GL 36, 41.2511°N 23.8208°E, leg. Subai
21.05.1997, S 15619/126 + 8 (juv.) + 3 (alk.); marble quarry 3.8 km from Káto Vron-
toú direction Panórama, 750 m alt., UTM GL 37, 41.2620°N 23.7880°E, leg. Subai
21.05.1997, S 15614/35 + 2 (alk.); 4.3 km from Káto Vrontoú direction Panórama, on
limestone rocks, 750 m alt., UTM GL 37, 41.2620°N 23.7986°E, leg. Subai 21.05.1997,
S 15615/6; river valley at N border of Káto Nevrokópi, on low limestone rocks, 650
m alt., UTM GL 38, 41.3498°N 23.8810°E, leg. Subai 21.05.1997, S 15616/5; Kalithia
(= W of Dráma), UTM GL 45, 41.11°N 23.91°E, leg. Hemmen 24.06.1987, Maa/3;
river gorge 500 m from Kalithia direction Anthohóri, on rocks, 215 m alt., UTM GL
45, 41.1249°N 23.9293°E, leg. Subai 08.05.1995, S 15632/7 + 1 (alk.); Angítis, on
limestone rocks beside spring at cave of Maras, 150 m alt., UTM GL 46, 41.2210°N
23.8928°E, leg. Fauer 10.08.1979, S 12877/4; leg. Subai 21.05.1997, S 15613/37; leg.
Gittenberger & Uit de Weerd 21.05.1999, NNM/10; approx. 6 km N of Prossotsáni, on
lime conglomerate rocks, 340 m alt., UTM GL 46, 41.2225°N 23.9694°E, leg. Subai
08.05.1995, S 15626/23; approx. 8 km from Prossotsáni direction Granitis, on lime-
stone rocks beside a creek, 500 m alt., UTM GL 47, 41.2597°N 23.9737°E, leg. Subai
22.05.1997, S 15611/58 + 5 (alk.); 400 m from the highway Prossotsáni–Granítis
direction Vólakas, on limestone rocks, 680 m alt., UTM GL 47, 41.2821°N 23.9815°E,
leg. Subai 08.05.1995, S 15625/21; N border of Galipsós, on limestone rocks, approx.
200 m alt., UTM GL 42, 40.8062°N 23.9502°E, leg. Subai 06.05.1995, S 15630/14 +
7 (alk.); mountain slope N of Podohóri, on limestone rocks, 450 m alt., UTM KF 42,
40.8473°N 24.0249°E, leg. Subai 27.05.1997, S 15624/17 + 1 (juv.); limestone gorge
6.5 km SE of Alistráti, on rocks and under stones, 80 m alt., UTM GL 54, 41.0224°N
24.0067°E, leg. Subai 23.05.1997, S 15621/30 + 3 (juv.) + 3 (alk.); 3–3.5 km from
Petroússa direction Pirgi, small gorge, on low limestone rocks, 510 m alt., UTM KF
46, 41.2332°N 24.0135°E, leg. Subai 21.05.1997, S 15618/17 + 2 (alk.); 25.2 km from
Akrovouni direction Pangéo peak, meadow with limestone rocks below the peak, 1780
m alt., UTM KF 53, 40.9113°N 24.0973°E, leg. Fauer & Subai 23.09.1989, S 15933/12;
N border of Xiropótamos, on limestone rocks and under stones, 285 m alt., UTM FK
56, 41.1951°N 24.1002°E, leg. Subai 08.05.1995, S 15628/63 + 6 (alk.); Falakró Moun-
tains, 3 km from the bifurcation Vólakas direction skiing center, deciduous forest with
limestone rocks, 1200 m alt., UTM KF 57, 41.2952°N 24.0091°E, leg. Subai 08.05.1995,
S 15633/11 + 2 (alk.); Falakró Mountains, border of Agio Pnevma plateau, meadow S
of the skiing center, under stones, 1720 m alt., UTM KF 57, 41.2993°N 24.0637°E, leg.
Subai 08.05.1995, S 15934/4 + 9 (alk.); Kalifitos (= NE of Dráma), UTM KF 66, 41.17°N
24.21°E, leg. Schütt 30.4.1962, SMF 279325/20; 11 km above Kalifitos, UTM KF 66,

87
Revision of the genus Lindholmiol a Hesse, 1931
41.25°N 24.29°E, leg. Fauer 31.7.1971, S 12472/5; 2.6 km SW of Makriplágio, on lime-
stone rocks, 520 m alt., UTM KF 66, 41.2082°N 24.2519°E, leg. Subai 08.05.1995, S
15627/17 + 11 (alk.); N border of Taxiárhis, under crystalline stones, 500 m alt., UTM
KF 66, 41.2328°N 24.1888°E, leg. Subai 22.05.1997, S 15612/50; mountain slope 1
km S from Vatholakkos, on low marble rocks and under stones, 450 m alt., UTM KF
66, 41.2224°N 24.1670°E, leg. Subai 22.05.1997, S 15935/2 + 29 (alk.); 1 km W of
Peristéria (= direction Adriani), on limestone rocks, approx. 400 m alt., UTM KF 75,
41.1177°N 24.3837°E, leg. Subai 23.05.1997, S 15620/4 + 1 (juv.); 1.3 km from Agorá
direction Peristéria, on limestone rocks, approx. 380 m alt., UTM KF 75, 41.1165°N
24.3244°E, leg. Subai 23.05.1997, S 15622/50.
Remarks: According to Urbanski (1960: 92), the holotype and 4 paratypes
of L. spectabilis were deposited in the NMNH. Currently, these specimens could
not be traced back in the museum in Sofia, and thus a paratype from the col-
lection in SMF is figured.
Although the name spectabilis suggests that this is a large species within
Lindholmiola, there are also populations with smaller or even dwarf-like shells
known. Also, the shell’s form can be quite variable. The population from the
type locality has broad whorls and a wide umbilicus, and inhabits mainly val-
leys and the lower altitudinal ranges in the mountains. A quite differing form
has a more densely coiled shell and a narrower umbilicus, and lives along
the southern slopes of the Pangéo Mountain and the higher altitudes of the
Falakró Mountains. Other populations comprise shells of normal size but with
a mixture of shells with a dense or wide rib pattern, and populations, where
only the one or the other phenotype is represented. Usually, shells with the
wide rib pattern occur more often in the western part of the distribution area,
while shells with the dense rib pattern are more abundant in the eastern part;
the boundary is approximately along the Angítis River.
A very special form lives at the north-western boundary of the Lekánis
Mountains between Agorá and Peristéria. The shells in these populations have
extremely flat whorls with a very sharp keel and an extraordinarily wide umbili-
cus. Nonetheless, they are so similar in all other shell morphological characters
that it is currently not possible to separate them on a subspecific level.
Distribution (Fig. 15): Lindholmiola spectabilis was described from Athos.
Its distribution area covers the western boundary of the Vrontoū Mountains
(Sidirókastro, Séres etc.) to the north-western slopes of the Lekánis Moun-
tains east of Dráma. In northern direction, this species has been recorded up
to Káto Nevrokópi and the Ágios Pnevma plateau in the Falakró Mountains.

88 Peter Subai & Eike Neub ert
Discussion
Three species of Lindholmiola, L. corcyrensis, L. gir va and L. lens, inhabit
quite large areas, while the distribution areas of another three species, L. bar-
bata, L. pirinensis and L. reischuetzi, are much more restricted (the area of L.
reischuetzi is small, but the species is quite abundant there) (Fig. 57). Only
two species, L. regisborisi and L. gyria, have very small and isolated habitats.
Particularly L. gyria, which is only known from a small area in south-western
Turkey, has only been recorded from a very few places, and seems to be the
most endangered species within the genus.
Fig 57. Distribution of the genus Lindholmiola.

89
Revision of the genus Lindholmiol a Hesse, 1931
Judging from the voucher collections of the museums listed above, we
know that in historical times, some species had much larger distribution areas.
A good example is L. corcyrensis, which once could be found along the Dalma-
tian coast northwards to the delta of the Neretva River, and there were also
records from the Gargano Peninsula in Italy. All these populations have to
be considered extinct; there are no modern records for this species beyond
the Albanian border. Also in its southern distribution area in the northern
Pelopónnisos, this species has disappeared.
Sympatric or even syntopic occurrences of Lindholmiola species are numer-
ous. The most widespread species, L. lens and L. corcyrensis can be found
sympatrically on the Island of Kephallinia and the northwester boundary of
the Bay of Korinthos. Other sympatric occurrences are also known for L. girva,
L. regisborisi, L. reischuetzi, L. pirinensis and L. spectabilis, always in differing
combinations. Particularly eastern Macedonia in Greece is very rich in Lindhol-
miola species, and in some places, up to four species can be encountered (for
example 2.5 km north-northeast Zarkadia, at the western boundary of Stav-
roupoli or north of Podohori). Until now, a sympatric occurrence of L. girva
and L. corcyrensis has not been recorded.
Acknowledgements
We are very grateful for the support during the work on the manuscript. This
particularly hold true for all the responsible curators of the museums, which
freely gave access to their voucher collections of Lindholmiola. We are par-
ticularly grateful to Prof. Dr. E. Gittenberger for giving his first, but incomplete
version of the manuscript to the senior author. Dr. M. Szekeres helped a lot in
localising places and the UTM-Code particularly in Rumania and Moldavia. We
particularly thank Dipl. Biol. Eva Feltkamp for preparation of the distribution
maps, and MSc. Estée Bochud for the many photos she took during the prepa-
ration of this manuscript.
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94 Peter Subai & Eike Neub ert
Addresses of the authors:
Peter Subai
Kronenberg 143
D–52074, Aachen
E-mail: subai@bio1.rwth-aachen.de
Eike Neubert
Naturhistorisches Museum der Burgergemeinde Bern
Bernastr. 15
CH–3005 Bern
E-mail: eike.neubert@nmbe.ch