taxonID	type	description	language	source
03DF87ECFFDF367FFF3887BD7F4F8CCA.taxon	materials_examined	Type species. Limax maximus Linnaeus, 1758; type by subsequent designation	en	Dedov, Ivaylo K., Schneppat, Ulrich E., Glogger, Fabia Knechtle, Cornu, Regula, Hristovski, Slavčo, Borissov, Simeon (2025): Morphological, anatomical, and molecular notes on two neglected high-altitude taxa of the genus Limax (Stylommatophora: Limacidae) from North Macedonia and their unusual adaptation to extreme alpine conditions. Zootaxa 5719 (1): 91-107, DOI: 10.11646/zootaxa.5719.1.4, URL: https://doi.org/10.11646/zootaxa.5719.1.4
03DF87ECFFDF3670FF38860179B08EFC.taxon	description	(Figs. 4 A, B; 5 A; 7 C, F)	en	Dedov, Ivaylo K., Schneppat, Ulrich E., Glogger, Fabia Knechtle, Cornu, Regula, Hristovski, Slavčo, Borissov, Simeon (2025): Morphological, anatomical, and molecular notes on two neglected high-altitude taxa of the genus Limax (Stylommatophora: Limacidae) from North Macedonia and their unusual adaptation to extreme alpine conditions. Zootaxa 5719 (1): 91-107, DOI: 10.11646/zootaxa.5719.1.4, URL: https://doi.org/10.11646/zootaxa.5719.1.4
03DF87ECFFDF3670FF38860179B08EFC.taxon	materials_examined	Material examined. North Macedonia • Jakupica Mountains (= Mokra Planina), Karadzhica ridge, eastern slopes of Solunska Glava, in open habitat, rocky alpine meadow, on limestone, 41.7136 ° N 021.3956 E °, 2279 m, leg. I. Dedov, 11.07.2011. BNM 063258 – 063260, BNM 064074 – 064081 / 11 specimens. (Fig. 2 A); • Jakupica Mountains (= Mokra Planina Mountains), Karadzhica Ridge, Marina Rupa area, a sinkhole / dolina, situated little north of Čeples peak and little southwest of Kantarica peak, in open habitat, rocky alpine meadow on limestone, 41.6781 ºN 021.3999 ºE, 2128 m, leg. I. Dedov, 10.07.2017, IBER-BAS, Coll. No. 10765 / 1 specimen; 1 more specimen observed (Fig. 2 B); • Jakupica Mountains (= Mokra Planina Mountains), Karadzhica Ridge, Marina Rupa area, a sinkhole / dolina, situated little north of Čeples peak and little southwest of Kantarica peak, in open habitat, rocky alpine meadow on limestone, 41.6776 ºN 021.3949 ºE, 2217 m, leg. I. Dedov, 10.07.2017, IBER-BAS, Coll. No. 10767 / 1 specimen; 2 more specimens observed (Fig. 2 C). Diagnosis Limax macedonicus macedonicus is a small sized species (55 – 80 mm) in comparison to other species of the genus living in Macedonia and elsewhere on the Balkans. Limax macedonicus macedonicu s is almost totally deep black invariably all over the body including the keel, which is not the case with any of the other accepted species of Limax with penial appendage, and a striking and distinctive character. Only the sides of the neck are cream-whitishyellowish, underpart of the free mantle flap, and the center field of the sole. The lateral sole fields are always dark-grayish-black, even in relatively small juveniles. The penis is c. as long as the body. The penis ends with a bent blind penial appendage, which is strongly coiled, massively thickening towards the insertion of vas deferens (VD) and penis retractor muscle (PR); VD and PR are inserting at the (more or less) same spot with no or almost no space between them. The duct of the bursa copulatrix (BCD) is attached close to the atrium. The subspecies L. macedonicus macedonicus is only known from the type locality. Morphology Body measurements are presented in Table 1. All measurements are taken from specimens preserved in ethanol 75 % + 3 % glycerol (N 2), in millimeters. No measurements of live specimens. Limax macedonicus macedonicus, unlike most species of the genus, are of an astonishingly uniform deep black color (Fig. 4 A, B). This black is covering any visible parts of resting or crawling specimens in any of the developmental stages known up to now. Even ommatophores and tentacles are entirely deep black as well as the skin between wrinkle rows and in deep folds. No other pigmentation or markings are visible. Cream-white are only the lateral sides of the neck, main underpart of the free mantle flap and the center field of the sole. The sole pigmentation fades partly after fixation in ethanol. In preserved specimens the black pigmentation is unusually dense and not to be compared with dark specimens of any of the known species of Limax species with penial appendage, e. g. Limax cephalonicus Simroth, 1886 and L. graecus (Fig. 4 C). In these species, even in very dark (“ black ”) pigmented specimens, there is always a hue of brown, ocher, etc., often a lightening to a very dark gray (personal observations). The only species known to us of this general dark black pigmentation is Limax wohlberedti Simroth, 1900, and L. cinereoniger but those species have no penial appendage. Mantle of L. macedonicus macedonicus is showing the typical “ fingerprint ” structures of Limax spp. when alive (Fig. 4 A, B). These are lost in preserved specimens. As already described by P. Hesse (1928), the wrinkle rows in L. macedonicus are fine and there are more wrinkles than in other species of the genus (Table 2). In fact, all wrinkles are slightly carinate and shallow in live specimens. Live animals are almost smooth at first sight, and especially when disturbed and secreting more slime. The thin and very fine sculptured skin is a distinctive character for both subspecies. The keel is totally black, very short and not especially high. Among the posterior dorsal wrinkle rows, it remains almost invisible without magnification (Fig. 4 A, B). P. Hesse (1928) mentioned the end of the dorsum to be characteristic and distinctive: the species exteriorly is distinguished by the high, suddenly steep falling down end of tail. In our opinion it is an erroneous observation, because P. Hesse had only preserved specimens at hand which were killed in ethanol and therefore contracted. We observed that the shape of the dorsum end varies when the specimens are fully stretched and / or crawling or the specimens are contracted (for example in rest position). Shell Shell is typical for the genus — strongly flattened, thin, asymmetrical (see Wiktor 1983). Embryonic part shifted posteriorly and to the left of longitudinal axis, more clearly defined in L. macedonicus macedonicus, than in L. macedonicus leucopus (see Wiktor 1983) (Fig. 5 A). Size: c. 10 mm. Anatomy The anatomy of L. macedonicus macedonicus follows the anatomy of L. graecus (Fig. 6): relatively long penis comparable to body length with bent blind penial appendage. The main differences that distinguish the alpine taxa from the other Limax species with penial appendage are: gray-black pigmented upper penis part (towards the penial appendage), including the penial appendage itself; the duct of the bursa copulatrix is attached close to the atrium, contrary to L. graecus in which the duct of the bursa copulatrix is attached to the free oviduct; VD and PR are inserting at the same spot with almost no space between, contrary to L. graecus in which there are distance between VD and PR. The main parts of L. graecus type anatomy are indicated on Fig. 6. The reproductive anatomy of L. macedonicus macedonicus is visualized on Fig. 7 C, F, and the differences with L. graecus (Fig. 7 G) are highlighted (Fig. 7 F). According to our observation, the size and the shape of bursa copulatrix, hermaphrodite gland, and albumen gland vary and are not species-specific (compare P. Hesse, 1928). Biology Limax macedonicus macedonicus, unlike the members of the genus, seem to inhabit only open limestone habitats, rocks and rocky alpine pastures, up to 2000 m. Early in the morning, in cool foggy weather (at about 8.00 h, 11. July 2011), the subspecies of L. macedonicus macedonicus was registered to be actively crawling on the grassy vegetation and on stones. A large number of active young and adult specimens were found in depressions, which remain longer under shadow of rocks during the day. At midday and afternoon single specimens only were found inactive, concealed under stones even in sunny places.	en	Dedov, Ivaylo K., Schneppat, Ulrich E., Glogger, Fabia Knechtle, Cornu, Regula, Hristovski, Slavčo, Borissov, Simeon (2025): Morphological, anatomical, and molecular notes on two neglected high-altitude taxa of the genus Limax (Stylommatophora: Limacidae) from North Macedonia and their unusual adaptation to extreme alpine conditions. Zootaxa 5719 (1): 91-107, DOI: 10.11646/zootaxa.5719.1.4, URL: https://doi.org/10.11646/zootaxa.5719.1.4
03DF87ECFFD03676FF38841A786C8D59.taxon	description	(Figs. 4 D; 5 B; 7 A, B, E; 8 A, B)	en	Dedov, Ivaylo K., Schneppat, Ulrich E., Glogger, Fabia Knechtle, Cornu, Regula, Hristovski, Slavčo, Borissov, Simeon (2025): Morphological, anatomical, and molecular notes on two neglected high-altitude taxa of the genus Limax (Stylommatophora: Limacidae) from North Macedonia and their unusual adaptation to extreme alpine conditions. Zootaxa 5719 (1): 91-107, DOI: 10.11646/zootaxa.5719.1.4, URL: https://doi.org/10.11646/zootaxa.5719.1.4
03DF87ECFFD03676FF38841A786C8D59.taxon	materials_examined	Limax (Limax) graecus, Simroth, 1889 — Wiktor, 1996: 77 – 78. Material examined. North Macedonia • Pelister Mountains (= Baba Planina Mountains), high-mountain pasture, 40.9939 ° N 021.1798 ° E, 2370 m, leg. S. Hristovski, 21.06.2023, IBER-BAS, Coll. No. 11010 / 6 specimens (Fig. 2 D). Diagnosis Limax macedonicus leucopus has similar reproductive system but differs from the nominate species by its smaller size (Table 2) and the creamy-yellowish-whitish colouration of the whole three sole parts (middle and both lateral sides). Limax macedonicus leucopus is only known from the type locality. As already described by P. Hesse (1928), the wrinkle rows from pneumostome to the back edge of the mantle in L. macedonicus (both subspecies) are finer and more numerous than in other Limax species with penial appendage (Table 2). Morphology Body measurements are presented in Table 1. All measurements are taken from specimens preserved in ethanol 75 % + 3 % glycerol (N 2), in millimeters. No measurements of live specimens. Limax macedonicus leucopus, like the nominate subspecies, has a strikingly uniform deep black color. (Fig. 4 D). The only difference from the coloration of L. macedonicus macedonicus is that in L. macedonicus leucopus the entire sole is creamy white. In L. macedonicus leucopus some blackish spots could be observed on the edge of the lateral zones of the sole, but never total pigmentation as in L. macedonicus macedonicus. The mantle of L. macedonicus leucopus has similar colouration and pattern as in L. macedonicus macedonicus. Shell Shell is typical for the genus (see comments on the previous subspecies) (Wiktor 1983) (Fig. 5 B). Size: c. 8.5 mm. Anatomy The anatomy of L. macedonicus leucopus is similar to L. macedonicus macedonicus. The reproductive anatomy of L. macedonicus leucopus is visualized on Fig. 7 A, B, E. Biology and mating behaviour We observed unusual mating behaviour of L. macedonicus leucopus from the alpine area of Pelister Mountains. Both pictures (Fig. 8 A, B) were taken on 07 July 2013, between 13.00 and 13.15 h. The specimens copulated in circle position, attached to vertical rock surface, without mucus thread, and their penises are hanging down. Molecular phylogeny The p-distances were calculated on a 1206 bp dataset and are given in Table 2. The dataset used for phylogeny included 31 DNA sequences and consisted of 573 bp from which 209 variable and 176 parsimony-informative sites. The new tree is similar to the one shown by Nitz et al. (2009). Although some of the branches received weak support, the populations from the Pelister and Jakupitsa mountains grouped together to form a strongly supported clade sister to a L. graecus + L. conemenosi clade (Fig. 3).	en	Dedov, Ivaylo K., Schneppat, Ulrich E., Glogger, Fabia Knechtle, Cornu, Regula, Hristovski, Slavčo, Borissov, Simeon (2025): Morphological, anatomical, and molecular notes on two neglected high-altitude taxa of the genus Limax (Stylommatophora: Limacidae) from North Macedonia and their unusual adaptation to extreme alpine conditions. Zootaxa 5719 (1): 91-107, DOI: 10.11646/zootaxa.5719.1.4, URL: https://doi.org/10.11646/zootaxa.5719.1.4
