Lindholmiola spectabilis URBANSKI, 1960

Subai, Peter & Neubert, Eike, 2014, Revision of the genus Lindholmiola HESSE, 1931 (Gastropoda: Pulmonata: Helicodontidae), Contributions to Natural History 23, pp. 1-94 : 82-87

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 Museums 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,

Description of shell: shell depressed, basic colour light brown to yellowishbrown; initial ½–1 of the teleoconch whorls smooth, the following whorls with fine radial stripes, becoming stronger on the subsequent whorls and developing 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 subsequent 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-circle 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 umbilicus, which live around the Bay of Korinthos and the north-eastern Pelopónnisos, 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, its surface granulation is finer, and if there are hairs, the haired area is larger. Additionally, L. lens has only two and not three penial pilasters. Another quite interesting difference is the mating behaviour: While in L. lens , the mating 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 ( Fechter & 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.

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 conglomerate 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 Vrontoú 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 limestone 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 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 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 , 41 .25°N 24.29°E, leg. Fauer 31.7.1971, S 12472 /5 ; 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, 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 collection 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 valleys 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 umbilicus. 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 Mountains east of Dráma. In northern direction, this species has been recorded up to Káto Nevrokópi and the Ágios Pnevma plateau in the Falakró Mountains.

Figures

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 girva, a: situs, b: lumina of genital organs, Greece, Makedonía, Taxiárhis.

Fig. 15. Distribution of Lindholmiola lens and L. spectabilis.

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 Neubert/Bochud, × 3.

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.

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