YU ISSN 0351 -0077
PRIRODOSLOVNI MUZEJ SLOVENIJE MUSEUM HISTORIAE NATURALIS SLOVENIAE
SCOPOLIA
12
Botanica
Geologica & Palaeontologica
Museologica
7 Zooiogica
Georg DŽUKJČ:
Taxonomic and Biogeographic Characteristics of the Slow-Worm {Anguis fragilis LINNAEUS 1758) in Yugoslavia and on the Balcan Peninsula
Taksonomske in biogeografske značilnosti slepca (An^is fragilis LINNAEUS 1758) V Jugoslaviji in na Balkanskem polotoku
SCOPOLIA
No 12
pp. 1-47 Ljubljana
Dec. 1987
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Journal ofthe Museum ofNatural History ofSlovenia. Ljubljana. Edited by the Museum of Natural History of Slovenia, subsidized by Research Community of Slovenia, Cultural Community of Slovenia. Centre of Scientific Research of the SASA and Univ. Institute of Biology E. Kardelj. Editional Staff: Jože BOLE. Ernest FANINGER. Janez GREGORI (Editor). Boris KRYSTUFEK, Ignac SIVEC, Kazimir TARMAN, and Tone WRABER. Readers: Cvetana TAVZES (for Slovene) and Helena SMOLEJ (for English). Address of the Editorial Officc and Admninistration: Prirodoslovn; muzej Slovenije, YU 61000 Ljubljana. Prešernova 20. The Journal appears twice a year, 500 copies per issue. Issue price for institutions and establishments 1000 din. for individuals 500 din. Current account at LB No 501(X)-603-40l 15. Printed by tiskarna Kurir. Ljubljana
TAXONOMIC AND BIOGEOGRAPHIC CHARACTERISTICS OF THE SLOW-WORM (ANGUIS FRAGILIS LINNAEUS 1758) IN YUGOSLAVIA AND ON THE BALCAN PENINSULA
DŽUKJČ Gcoib
Institut za biološka istraživanja
»Siniša Stankovič«
YU 11060 Beograd, 29. novembra 142 Received: 10. 12. 1986
UDC (UDK) 598.112.6 (497) (045) = 20 Anguis Jragilis
iz VLECEK" TAKSONOMSKE in BIOGEOGRAFSKE značilnosti slepca (ANGUIS FRAGILIS LINNAEUS 1758) V JUGOSLAVIJI IN NA BALKANSKEM POLOTOKU - Proučevana je variabilnost zunanjih morfoloških značilnosti vrste Anguis fragil is LINNAEUS 1758 v Jugoslaviji in na os bvbv Balkanske m polotoku. Dokazana je prisotnost nominantne (A.ffragilis) in kolhidske (A.f.colchicus) podvrste slcpca ter kontaktne cone med njima. Določeni so areali v Jugoslaviji za vrsto Anguis fragilis ter obe njeni podvrsti.
ABSTRACT - The object of study is the variability of external morphological characteristics of the species Anguis fragilis LINNAEUS 1758 in Yugoslavia and the rest of the Balcan Peninsula. Proved is the presence of the typical (/4./ fragilis) and the Colchidic {A.f. colchicus) subspecies of slow-worm and a contact zone between them. Determined are the ranges of the species Anguis fragilis and both of its subspecies in Yugoslavia.
Introduction
After WERMUTH'S introduction of variational statistics (1950) which set the basis for the solution of the majority of the problems concerning intraspecific and secondary sex variability, the lizard Anguis fragilis became an object of intensive taxonomic investigations in the countries covered by its range. A number of works were published dealing with the material from the authors* countries or elsewhere (VOIPIO 1956, 1962, FUHN 1961, STUG-REN, FUHN, POPOVICI 1962, LUKINA 1965, BEŠKOV 1966, LÄC 1967, PETZOLD 1971, DELY 1972, 1974, 1981. MUSTERS, BOSCH in den 1982). Most of the authors adopted the standpoint of WERMUTH (1950) and MERTENS & WERMUTH (1960), namely, that the slow-worm is a polytypic species represented in its vast range by three subspecies: a typical - Anguis fragilis fragilis L., a Colchidic - Af. colchicus (NORDMANN) 1840, and a Peloponnesian - Af. peloponnesiacus STŠPANEK 1937. Thus, they determined the intraspecific appurtenance of the slow-worm populations of their own national territories, and where several subspecies were present, also their interrelations and delimitations. Though not denying the polytypic character of the species Soviet authors consider the identification of the Colchidic subspecies unfounded (NIKOUSKI 1913, 1915, TEREN-TJEV&CERNOV 1936,1949, LUKINA 1965) or suspicious (BANNIKOV, DAREVSKIJ & RUSTAMOV 1971. BANNIKOV, DAREVSKIJ, iSČENKO, RUSTAMOV & ŠČER-BAK 1977).
Due lo its central position in the general range of the species, as well as the central position it occupies on the Balcan Peninsula - the only territory in which all three species meet - Yugoslavia could be neglected in no paper on the origin of the species and the subspecies, the history of the formation of the present range, the intraspecific variability, the distribution and the comparison of single populations in geographic relationships etc. However, with reference to our contry the conclusions presented in the works referring to the above mentioned problems are, to say the least of it. quite generalized. A lack of the evidential material led the authors to speculate and proceed on the basis of analogy with the surrounding regions so that in the majority ofcases the opinions are definitely divided. Thus, some authors (WER-MUTH 1950. MERTENS & WERMUTH i960) believe that exclusively An^^iiis f. colchivus lives in our country, others (VOIPIO 1962. FUHN & VANCEA 1966. POZZI i966) state, beside the typical one. also the Colchidic subspecies, with arbitrary delimitations between them, while some of them (MUSTERS & BOSCH in den 1982. HENLE 1985) have taken no definite standpoint yet.
Considerable disparities to be noticed among the characters of the populations of.l./ra-giUs from the territory of Yugoslavia make the solution of the microtaxonomy and the zoogeography of the species and the research thereof quite diflicult. Trying to moderate this conflict I decided upon researching the intraspecific variability of the slow-worm in our country. By means of a complex analysis of the external morphological characters on a larger number of specimmens I wished to obtain such results as to eliminate any arbitrariness from the interpretation of the taxonomic and biogeographic characteristics of this species in Yugoslavia. These results would represent also the missing link, making it possible to speak more reliantly on the geographic delimitation of the taxa within the species Angtiis fragHis.
Acknowledgements
The author would wish lo lhank his colleagues from the Natural His(or>' Museum in Belgrade, the National Zoological Museum of Croatia in Zagreb, the State Museum of Bosnia and Herzegovina, the Institute ofZoology at the Faculty of Natural and Mathematical Sciences in Belgrade, the Natural Hislor> Museum of Slovenia in Ljubljana, as well as the »V. Mandič-Manda« Society of Reserchers in Valjevo. all of whom not only made it rx'^sible for him to have access to the relevant collections but most generously helped him in his work.
I would particularly wish to express my gratitude to my friends and colleagues Prof. S. BRELIH. N. TVRTKOVIČ. MSc., and Dr. Ü. PASU UEVIČ for their unselfish help in collecting and studyng the material.
1. Suney of the past investigations dedicated to (he intraspcciflc variability of the species A. fragilis from Yugoslavia
The principal hcrpetological works dating back to the end of the 19»^ century' i.e. »Her-pctologia Europaea« (SCHREIBER 1875) and »Die Reptilien und Amphibien (^terreich-Ungams und der Occupationsländer« (WERNER 1897) abound with imprecisions with respect to the denominations and the distribution of the Colchidic subspecies, so it is not surprising to find a similar state of the matter rcfiected also in our herF>etological literature of that time and the beginning of the 20»'» centur>'. Even though fine experts in our herpctofauna. SCHREIBER and WERNER could state nothing definite nor generalized on this subspecies in our country. The reasons therefore could be found in the absence ofthe Austro-Hungarian domination over the eastern parts of Yugoslavia (cf. DŽUKIČ 1979) and a lack of the material from this region, respectively.
The first confirmation of the presence of the subspecics A.f.cokhicm in Yugoslavia can be found in the monography »Amphibians and Reptiles of Serbia« (DJORDJEVIČ 1900).
DJORDJEVIČ dearly distinguishes between the Colchidic and the typical subspecies, staling: »Some specimens of this variety make part of our collection.«' He does not enter into considerations on the intraspecific variability and the geographic distribution of single »varieties« but merely slates the presence of the typical and the Colchidic subspecies in Serbia.
Published in 1916 was another Yugoslav work on the Colchidic subspecies i.e. SELIŠ-KAR's paper »Blue-Spotted Slow-Worm {Anguis fragilis L. var inceria KRYN)«. In this work SELISKAR specifically pointed out an interesting fact, namely, that the above-mentioned »variety« had been found at Kamniška Bistrica, far from all localities known by then, however, he made no attempt to explain it but dedicated all his attention to the description of external morphology and the problems of dependence of such a morphological type upon various factors.
In the period between the Wars, the time which saw the work of three of our greatest herpetologists (BOLKAY. KARAMAN, RADOVANOVIČ) no special importance was attributed to the species Angitis fragilis and the problems connected to its intraspecific variability. Whenever discussing the slow-worm they consider it a monotypic species. It is interesting to note that not even CYR^N (1941) who dedicates a detailed study to the relationships within the species .'l.yröj^/7/.v on the Balcan Peninsula considers the Colchidic subspecies valid. As evident from his work »Beiträge zur Herpetologie der Balkanhalbinsel« (»Contributions to the herpetology of the Balcan Peninsula«) no material from Yugoslavia was at his disposal.
After the World War 11 the slow-worm continues to be neglected by ourauthors whereas elsewhere in Europn; this species becomes an object of unusually intensive studies. Thus, as late as the seventies we can only speak of such investigation results concerning the problems of the intraspecific diflerentiation of the Yugoslav population of slow-worms as presented by foreign authors.
As pointed out before, the scientific zeal in the research ofthe species A.fragilis was initiated by WERMUTH (1950). In his principal work forthe understanding of intraspecific variability he included also a material from Yugoslavia. Yugoslavia could then be found together with parts of Italy and Austria, Hungary, the major part of Rumania, Bulgaria, Albania. Greece and the European pan of Turkey in the region - southeastern Europe. The population oiA.fragiiis from this region was treated as unique and having been subject to a statistical analysis it revealed a scries of characteristics in comparison with other regions that WERMUTH singled out from the slow-worm range.
Published in 1951 was a monography »The British Amphibians and Reptiles« (M. SMITH) the subject matter of which reaches far beyond the limits indicated by the title. In the chapter dealing with the slow-worm i.e. that part ofthe monography in which SMITH discii'v'sc'i inini<pecific categories, he analyses also a material from Yugoslavia. At the time Smith was undoubtedly not familiar with WERMUTH's work so he was confronted with the same kind of problems as numerous authors before him i.e. the problems of ascertaining discontinuity in a seemingly continuous series of morphological variants so that he supported the idea ofthe non-existence of the Colchidic subspecies. He nonetheless devoted much attention to the blue-spotted »variety«. However, like TERENTJEV and CERNOV (1936, 1949, 1965) he connected this phenomenon exclusively with sexually mature males. In the part in which he discusses the distribution ofthe blue-spotted »variety« he states that among the slow-worms from different countries he researched also a material from Yugoslavia.
MERTENS and WERMUTH (I960) generalized WERMUTH's conclusion (1950) on the microtaxonomy ofslow-worms and the distribution of single subspecific taxa, consequently, there prevailed a belief that in Yugoslavia the species Angiiis Jragilis was represented by
Collection ofthe Zoological Institute of High schod in Belgrade
the Colchidic subspecies. As also according to WERMUTH the delimitation with respcci to the typical subspecies is fairly approximative it is drawn rather arbitrarily in a scries of works by different authors (FUHN & VANCEA I960, VOIPIO 1962, STUGREN, FUHN & PO-POVICI 1962, PAVLETIČ 1964, POZZI 1966, BESKOV 1966. LÄC 1967, DELY 1972. 1974, 1981, MUSTERS & BOSCH in den 1982).
VOIPIO (1962) makes an interesting remark, namely, that even though SMITH (1951) stated specimens with blue spots in Yugoslavia, they did not make part of the six specimens from our counlr>' as available to WERMUTH. On the basic of these facts VOIPIO drew an important conclusion i.e. that in the region of »southeastern Europe« the geographical distribution was not homogeneous. He believed that the »colchicus« characters were concentrated in the extreme southeast of this region (Rumania, Bulgaria, and Greece).
In her notes at the end of the index of amphibians and reptiles figuring in the collection of the Croatian Zoological Museum in 2^grcb Jela PAVLETIČ (1964) says that FUHN and VANCEA (1961) state the Colchidic subspecies for the eastern parts of Yugoslavia, so she considers it necessary to verify the subspecific appurtenance of the material from the localities in the region of Slavonia.
An extensive study on the zoogeography and taxonomy of Yugoslav amphibians and reptiles by the Italian author POZZI (1966) represents another example of a subjective delimitation of the intraspecific slow-worm taxa in our country. According to POZZI the typical subspecies is associated with the territory ofSlovenia. Croatia and some islands whereas the Colchidic one with the central and the southern part of Yugoslavia: Bosna, Herzegovina, Montenegro, Serbia and Macedonia.
Published on the basic of preliminary investigations were the first native results on the intraspecific differentiation and distribution of A. fragilis in Yugoslavia (BR EL IH & DŽU KIČ 1974). We then established the presence of both subspecies, whereas A. fragilis fragilis is characteristic of the west of our country whereas in the east it appears only on highlands. while A. f. colchicus is associated with the eastern and southeastern parts of Yugoslavia, appearing merely locally in the west. We were not quite sure about the demarcation line between the subspecies, which was duly pointed out also in the paper.
A detailed analysis of slow-worms in Yugoslavia was presented by DZUKIČ in his Master's Thesis the results of which are incorporated in the present work.
Subjective views on the microtaxonomy, the status and delimitations of single intraspecific taxa are disclosed in the works of DELY (1981), MUSTERS and BOSCH in den 1982. and HENLE (1985), while the presence of slow-worms revealing the caracteristics of the Colchidic subspecies on the coast under the Velebit Mis. is mentioned by BANK, KRUYN-TJENS & PAULISSEN (1982).
2. Material and methods
In determining the intraspecific differentiation and horizontal and vertical distribution of subspecific categories of A. fragilis in Yugoslavia 1 made use of a material from the entire ierritor>' of our country. In total I analysed 310 specimens from 135 localities. For the purposes of a comparison with populations from other territories 1 dcali with, in addition to the references from the literature, also with 12 specimens from Greecc. Mbania. and Spain.
Of totally 310 specimens 91 specimens make part of my own collection stored at the »Si-nisa Stankovič« Institute for Biological Research (IBIS) in Belgrade. 66 specimens belong to the Natural History Museum ofSlovenia (PMS) in Ljubljana, 41 to the Croatian Zoological Museum (HNZM) in Zagreb, 34 to the State Museum of Bosnia and Herzegovina (ZMBH) in Sarajevo, 23 to the herpetological collection of the late Prof Dr. Milutin Radovanovic (MR), 21 to the Natural History Museum (PMB) in Belgrade, 20 to the collection of Prof Dr. Gojko Pasuljevic (GP), 12 to the herpetological collection of the Young Researchers
»Vladimir Mandioč - Manda« (VMM) in Valjevo, 3 lo the collcclion of Boban Zečevič (BZ) from Zaječar, and 2 specimens are studied on the basis of the data presented by Dr. G. Dely.
A survey of localitiers is given in Table I and their geographical position in Fig. I. The reference number of a locality corresponds lo its number in the map.
Most of the material is preserved in 75 % alcohol and a smaller part in 4 % formalin. The specimens from the collection of B. Zečevic are dermoplastic preparations.
Considerable difficulties presented themselves in the case of museum exhibits kept in scaled cylinders, as well as the material fixed in spiral form. I was not in position to study such specimens in detail. In the former case I took into consideration only the characters visible through the glass walls of the vessels whereas in the latter exact linear measures could not be taken.
In selecting morphological characters to be analysed I decided on those that proved most reliable in the course of the past studies dedicated to the problem of intraspecific differentiation of the slow-worm. In the first place the characters concerned are qualitative morphological characters, namely: 1. presence or absence of clear ear openings: 2. relation between the homy plates of the pileus: 3. colour and appearance of blue spots.
With respect to quantitative morphological characters I took into account their meaning as already ascertained and primarily made use of the number of scales in an average series around the body. Other quantitative characters arc less significant than those mentioned above, that is why they are less frequently applied in analyses. I nonetheless took the following measures: I. length of the head and the trunk (measured from snout to vent); 2. length of the tail (measured from vent to the tip of the tail); 3. length ofthe pileus (measured from the snout to the back margin ofthe occipital plate).
Statistical methods were also applied in describing and analysing the results, as well as drawing conclusions. Taken as basic parameters are the values ofthe above-mentioned measures (X), the number of specimens in a sample (N), and the frequency of single qualitative characters in difTerent populations of slow-worms. Calculated by means of these parameters are arithmetic mean (X) and standard error of the difference between two arithmetic means (Sx, - Xj).
The T-test was applied to define the statistic meaning of differences between two populations.
Taking into account the fact that the analysis of qualitative morphological characters yielded incomparably better results than the analysis of quantitative morphological characters. when comparing populations o^A.fraplis from difTerent parts of its range I considered mainly the comparison of the first characters, applying the method of hi-square test (x^).
When testing two independent samples I entered the results into a 2 x 2 and calculated x^ - lest by applying i he nu called Yalcs's cuiiection.
3. Results
3.1. Variability of basic difTcrential charactcrs
In the chapter on the material and the methods applied I explained in detail which morphological characters would be analysed and in what way. In presenting the results of researching the morphological characteristics of every single specimen, I made use of modified DELV's tables (1972, 1974). By adopting this manner of presenting the results one gets an insight into group as well as individual variability of slow-worms in Yugoslavia. Besides, other authors are offered a possibility of comparing the results anc getting into closer contact with the material 1 treated. In the appendix of tables roman numbers arc used to designate the following characteristics: I - running number of the specimen: II - number ofthe specimen in the corresponding collection; III - sex; IV - length of the head and the trunk; V - length of the tail; VI - length of the pileus; VII - presence of ear openings: VIII - pileus
i \ ä 1 u
V)

Fig. t. Survey of (he localities from which the material analysed in (he present paper originales. Slika i. Pregled lokalitet. na katerih je bil zbran analizirani material.
Table Survey of localities.	Tabela I: Pregled lokaliiel
Locality
No. Locality
1	Slovenia: the Julian Alps: Rombon (1200-1300 m a.s.l.)
2	Slovenia: the Julian Alps: Poiovnik
3	Slovenia: the Julian Mp^- Mangart: Planina (highland) below Mangart
4	Slovenia: the Julian Alps: Kmsko jezero (Lake of Krn)
5	Slovenia: Kranjska gora: Martuljek: Srednji vrh
6	Slovenia: Karavanken Mts. above Gozd Martuljek: behind Lepi vrh
7	Slovenia: the Julian Alps: Triglav group: Krma (900 m a.s.l.)
8	Slovenia: the Julian Alps: Komarča
9	Slovenia: the Julian Alps: Bohinj: (Jkanc
10	Slovenia: Koper
11	Croatia: Istria: Buje
12	Croatia: Istria: Rovinj
13	Slovenia: Ajdovščina
14	Slovenia: Radovljica
15	Slovenia: Kranj
16	Slovenia: Domžale
17	Slovenia: Ljubljana: Bežigrad (300 m a.s.l.)
18	Slovenia: Javor
19	Slovenia: Preserjc
20	Slovenia: vicinity of Ig: Kremenica
21	Slovenia: Vrhnika: Borovnica: Kopitov grič
22	Slovenia: Mokrec
23	Slovenia: Rašica
23 a Slovenia: Travna gora
24	Slovenia: Snežnik (highland)
25	Croatia: Gorski Kotan Risnjak National Park: Cmi Lug: Markov brlog and Velika voda
26	Croatia: Gorski Kotar: Delnice: Rogozno
27	Croatia: Gorski Kotar Vrbovsko: Zapeč
28	Croatia: Gorski Kotar Plešce 28 a	Slovenia: Kočevje
29	Croatia: Bakar
30	Croatia: Krk (island): Soline
31	Croatia: Novi Vinodol
32	Croatia: Crikvenica-Vinodol: Kotor
33	Croatia: Krk (island): Baška Draga
34	Croatia: Krk (island): Krk
35	Croatia: Senj
36	Croatia: Croatian seaside: Draga
37	Croatia: Jablanac
38	Slovenia: Pohorje
39	Slovenia: Pohorje: Pungert (1500 m a.s.l.)
40	Slovenia: Boč (highland): Poljčane (850 m a.s.l.)
41	Slovenia: Podčetrtek
Locality
No. Locality
42	Slovenia: Slovenske Gorice: Miklavž
43	Croatia: Mali Kalnik
44	Croatia: Medvednica (highland): Medvedgrad
45	Croatia; Medvednica (highland): Kraljičin zdenac
46	Croatia: Medvednica (highland):
47	Croatia: Zagreb
48	Croatia: Samobor
49	Croatia: Turopolje: Peščenica
50	Croatia: Ogulin
51	Croatia: Velika kapela (highland): Razvala
52	Croatia: Vrhovine: Doljani: Topoluša (800-1000 m a.s.l.)
53	Croatia: Vrhovine: Donji Babin Potok: Boriča Borik (850 m a.s.l.)
54	Croatia: Velebit (highland): Zavizan
55	Croatia: Velebit (highland): Golic (1450 m a.s.l.)
56	Croatia: Velebit (highland): Ostarije
57	Croatia: Lika: Gospič
58	Croatia: Velebit (highland): Paklenica
59	Croatia: Velebit (highland): Predzid (660-750 m a.s.l.)
60	Croatia: Slavonia: Vocin
61	Bosnia and Herzegovina: vicinity of Jajce
62	Croatia: Split: Maijan (locality under justified suspicion)
63	Croatia: Vis (island)
64	Bosnia and Herzegovina: Glamoc
65	Bosnia and Herzegovina: Livno
66	Bosnia and Herzegovina: Vranica (highland)
67	Bosnia and Herzegovina: Dcrventa
68	Croatia: Mljet (island): Govcdjari
69	Bosnia and Herzegovina: Igman (highland): Donja Grka-Veliko polje
70	Bosnia and Herzegovina: Treskavica (highland)
71	Bosnia and Herzegovina: Zelengora (highland): Gornje bare
71	a	Bosnia and Herzegovina: Čemerno
72	Bosnia and Herzegovina: Crvanj (highland)
73	Bosnia and Herzegovina: Bjelašica
74	Bosnia and Herzegovina: Korito
75	Bosnia and Herzegovina: Trcbinje
75	a	Montenegro: Vilusi
76	Croatia: Slavonia: Valpovo
77	Croatia: Baranja: Haljevo (forest)
78	Croatia: Slavonia: Djurdjanci
79	Bosnia and Herzegovina: vicinity of Višegrad
80	Montenegro: Durmitor (highland): Zminje jezero (lake)
81	Montenegro: Nikšič
82	Montenegro: Donja Lastva (locality under justified suspicion)
83	Montenegro: Podgorica-Titograd
84	Montenegro: Crmnica: Brčeli
85	Montenegro: Novi Bar
86	Montenegro: Ivangrad: Rovca
87	Serbia: Voivodina: Fruska gora (highland)
Locality
No. Locality
88	Serbia: Voivodina: Fruška gora (highland): Andrevljc
89	Serbia: Debrc: Vlasanica
90	Serbia: Sokolske planine: Šljivova
9)	Serbia: foot of Jablanik (highland): Pocuta: Bebica Luka
92	Serbia: Valjevo: Ribnicka klisura
93	Serbia: Maljen (highland): Divčibare 93 a	Serbia: Osečenica: Bunčevica
94	Serbia: Ljubovija: Gornja Trešnjica
95	Serbia: vicinity of Užička Požega: Glumač. Bakionica, Zdravčiči
96	Serbia: Tara (highland): Vidikovac
97	Serbia: Tara (highland): Tarabica brdo
98	Serbia: Tara (highland): Djurdjevo brdo
99	Serbia: Tara (highland): Vežanja
100	Serbia: Belgrade: Topcider )01	Serbia: Belgrade: Košulnjak
101	a	Serbia: Avala (highland)
102	Serbia: Kosmaj (highland)
103	Serbia: Bukuija (highland)
103	a	Serbia: Rudnik (highland): eastern slopes
104	Serbia: Batočina: Rogot (forest) (129 m a.s.l.)
105	Serbia: Glednicke planine
106	Serbia: Vitanovac
107	Serbia: Goč (highland): Brezjanska kosa
108	Serbia: Studenica: Savovo
109	Serbia: Jasirebac: Prokopačka kosa
110	Serbia: Kosovo: Leposavic: Gnje, dani
111	Serbia: Kosovo: the East 111a	Serbia: Kosovo: Babaj Boks
112	Serbia: Kosovo: Priština: Grmija
113	Serbia: Kosovo: Šara: Brezovica
114	Serbia: Kosovo: Šara: Piribeg
115	Macedonia: the Treska Canyon
116	Macedonia: Pelister (highland): Begova česma
117	Serbia: Voivodina: Vršački breg
118	Serbia: Djcrdap: Dobra
119	Serbia: Donji Milanovac
120	Serbia: Tekija: the Kasljinska reka (river)
121	Serbia: Majdanpek: Blagojev Kamen
122	Serbia: Sisevac
122	a	Serbia: vicinity of Zajecar
123	Serbia: Ozren (highland)
124	Serbia: Vučje: canyon of the Vucjanka river
125	Macedonia: Kožuf (highland)
126	Serbia: Bistar
SIov.	Slovenia without a precisely stated locality
Hrs'.	Croatia without a precisely staled locality
Srb.	Serbia without a precisely staled locality
Kos.	Kosovo without a precisely stated locality
E	Spain: Mont Serrat
Locality
No.	Locality
GR I	Greece: Olympia: Kladeos (valley)
GR 2 Greece: Akamanien
AL I	Albania: Kanina
AL 2	Albania: Valona: Parha Liman
AL3	Albania: Velipoja
AL 4	Albania (northern part)
type; IX - number of scales in an average series around the body; X - presence of blue spots; XI - date of the finding; and XIl - number of the locality. Presence and absence of a qualitative morphological character is designated as + and respectively. Only in the case of the existence of a depression in the place of an ear opening the sign = (according to DELY 1974) is used. The abbreviations reg. and def accompanying the linear measures in the tail length column indicate whether the tail is regenerated or defective.
As ensuing from the table, the number of localities and their geographic position are directly interdependent. Already in identifying the localities by means of numbers I decided upon a succession from low r to higher numbers, starting from the northwest towards the southeast. My decision was \ ased on WERMUTH's conclusions (1950). namely, that from a global point of view morphological characters of the slow-worm, especially those taxono-mically important, vary in the function of longitude. In my belief such an approach was methodologically the most appropriate as it provides answers to the fundamental questions of morphological variability and geographical distribution of single variants in the territory of Yugoslavia.
With respect to the frequency of positively linked inlraspecific characters the material investigated was divided in three groups.
3.1.1. Groups of individuals with dominant characters of the typical subspecics
Gathered in the first group (Table 2) are all individuals characterized by the domination of the characters of the typical subspecies, as well as some specimens with a more marked presence of the »colchicus« characters originating from localities in the region otA.fragilis fragilis. The majority of slow-worms, no less than 215 specimens (72 %) proved to belong to this group. Not only does this population dominate in number but it also covers more or less the entire territory of our country. The region includes the whole of Slovenia, almost all of Croatia, Bosnia and Herzegovina, a large part of Montenegro and western Serbia with Kosovo (Fig. 2).

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Slika 2. Razi»iijcnost nominantnc in kolhidskc podvrste slepca v Jugoslaviji.
Due to the reasons explained in the chapter on the material and the methods applied as well as the inadequacy ofjuvenile specimens, it was impossible to subject every single specimen to the analysis of all basic differential characters. An analysis of presence or absence, respectively, of clear ear openings was performed on 186 specimens of this population, furthermore. an analysis of the relation among homy pileus plates on 192 specimens, and the presence of blue spots on all of 215 specimens. The presence of blue spots could only be ascertained in males so that for further treatment and comparisons calculations were prepared only on 76 males of this population. The number of scales in an average series around the body was established in 182 specimens.
3.1.1.1. Ear openings
With respect to the existence of clear car openings in slow-worms making part of the first group the situation is as follows:
+ + 1.1
+=
l.I % 27.4% 4.3% 66.1
n
2+2+51+8+123=186
Only four specimens had ear openings two of which had an ear depression on one side of the head only. The specimens with symmetrical ear openings originate from Kranj and
GlamoČ, whereas those with one ear opening from Domžale and the Tara highlands. Even though in a geographical sense the specimens with ear openings are markedly dispersed, a certain grouping tendency of the »colchicus« character can be noticed in the specimens from the area of Kranj-Kamnik-Domžale. Taken into consideration in the present case is also SE-LlŠKAR'sdatcon Kamniška Bistrica. Procentually speaking 2.2 % of the specimens had ear
100
90 80 70 60 50 40 30 20 10
with an ear openings z ušesnimi odprtinami
without ear openings brez ušesnih odprtin
1	A.f.fragllls
2	A.f.fraglll$ ^ A.f.colchicus
3	A.f.colchicus
Fig. 3. Proccntual frcqucncy of the occurrcncc of ear openings within three slow-worm populations in Yugoslavia.
Slika 3. Odstotna zastopanost pojavljanja ušesnih odprtin znotraj treh populacij slepca v Jugoslaviji.
openings whereas in 97.8 % (Fig. 3) the ear openings were perfectly covered. A marked feature of this population of slow-worms in Yugoslavia is the absence of ear openings.
3.1.1.2. Pileus
Various possibilities of the position of prefrontal plates arc shown in Fig. 4 and the following table:
Pileus Type A 62.0%
Pileus Type B
29.2%
Pileus Type C 8.8%
119 + 56+17=192
As evident from the above, in the majority of specimens the prefrontal plates come into contact at considerable lenght, in one third thereof they touch one another in one point, whereas the number of specimens in which the prefrontal plates are separated from one another by the inter nasal shield and the frontal shield is very small.
!! I
pileus type tip ptieusa
WM pileus type g ŽS^ tip piieusa
pileus type tip piieusa
1	A.f.fraailis
2	A.f.fragilis ^ A.f.colchicus
3	A.f.colchicus
Fig. 4. Proceniual frequency of the occurrence of single pileus types wiihin three slow-worm populations in Yugoslavia.
Slika 4. Odstotna zastopanost pojavljanja posameznih tipov pilcu&a znotraj treh populacij slepca v Jugoslaviji.
3.1.1.3. Blue spots
Blue spots represent the third variable character important to the microtaxonomy of slow-worms. Relative frequency of this character within the first group of individuals is as follows:
30.3% 69.7%
n
23 + 53 = 76
A graphical illustration of the result is shown in Fig. 5. The appearance of spots in the specimens making part of the slow-worm group with dominant characters of the typical subspecies is quite frequent, taking into account the fact that it is characteristic of one third of the males.
3.1.1.4. Number of scales
The number of scales in an average series around the body represents one of the most important as well as stable difTerential characters of slow-worms. In the first population the frequency of specimens with a definite number of scales around the body is evident from the table and Fig. 6.
100* 90 80 70 60 50 40 30 20 10
with blue spots s plavimi pegami
without blue spots brez piavih peg
1	A.f.fragllis
2	A.f.fragills % A.f.colchicus
3	A.t.colchicu8
Fig. 5. Proccnlual frequency of the occurrence of blue spots within three slow-worm populations in Yugoslavia.
Slika 5. Odstotna zastopanost pojavljanja piavih peg znotraj treh populacij slepca v Jugoslaviji.
23 24 25 26 27 28 1.1% 32.4% 25.8% U.6% 3.9% 2.2%
2 + 59+47 + 63 + 7 + 4= 182
The number of scales varies from 23, in fact the lowest number of scales in a slow-worm recorded in Yugoslavia, to 28. The mean value of this character for the entire population amounts to: X = 25.1.
3.1.2. Group of animals from Che contact /one of the typical and the Colchidic subspecies
Another group (Table 3) is represented by the animals from the contact zone of the typical and the Colchidic subspecies, which endows this population with a »mixed« character. Of 303 specimens of slow-worms from Yugoslavia 43 (14 %) belong to this group. Contrary to the previous case, one cannot speak ofa range ofboundaries of the range ofthis population but much more appropriately of a zone in which the two subspecies contact each other without clearly drawn border lines. Its position is presented in Fig. 2 and a detailed description in the following chapter.
Not all of the specimens of this group of animals could be subject to the analysis.
100 90 80 70 60 50 40 30 20 10
22-23 24-25 26-27 28 -29
30.31
1	A.f.fragilis
2	A.f.fragilis % A.f.colchicus
3	A.f.coichicus
1
Fig. 6. Procenlual frequency of the number of scalcs (in an average scries) within three slow-worm populations in Yugoslavia.
Slika 6. Odstotna zastopanost Števila lusk (v povprečnem nizu) znotraj treh populacij slepca v Jugoslaviji.
3.1.2.1. Kar openings
42 specimens were analysed to determine procenlual frequency of ear openings in the contact population.
+ + + = 2.4% 7.1%
47.6% 42.8%
l+3 + 20+l8:=42
In one specimen symmetrical ear openings were stated whereas three specimens had ear openings on one side of the head only. There exist indications of an increasing number ofspn:-cimens with an asymmetrical ear opening in the contact population. Totally« 4 specimens (9.5 %) had ear openings and 38 specimens (90.4 %) had none. The results oflhe analysis are summarized in Histogram I. With respect to the previous populations one notices a negli-geable increase in the frequency of ear openings.
3.1.2.2. Pileus
Relative frequency of this character was calculated for 43 specimens. The relation among (he prefrontal shields does not depend on sex and age.
Pileus Type A
41.9%
Pileus Typ)e B
34.9%
Pileus Type C
23.2%
18+15 + 10 = 43
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3.1.2.3. Blue spots
Relative frequency of the occurrence of blue spots was calculated on 23 males, by taking into account the same reasons as in the previous group of individuals.
+	-	n
74.0% 26.0%	17 + 6 = 23
In addition to the table the results are pjesented also in Fig. 5. Already in conncection with the previous group attention was called to a relatively high frequency of the occurrence of blues spots in comparison with west-European populations. An increase in frequency is even more markedly expressed in the contact population where no less than 74 % of males have blue spots.
3.1.2.4. Number of scales
In the contact population the number ofscales ranged between 24 and 28. The frequency of the specimens with a definite number of scales ensues from Fig. 6 and the folowing table:
24 25 26 27 28	n
2.9% 22.9% 57.1 % 11.4% 5.7%	I +8 + 20 + 4 + 2 = 35
With regard to the previous group the variation range is somewhat narrower. One can notice the prevailing of the specimens with 26 scales around the body. The arithmetic mean of the number of scales in this population amounts to X = 26.
3.1.3. Group of individuals with dominant characters of the Colchidic subspecies
The individuals characterized by the domination of the »colchicus« characters are classified into the third group (Table 4). Of the total number of the specimens as studied 45 (14 %) make part of this group. The range of this population comprises the eastern and the southeastern F>arl of our country. The contact zone separates it from the range of the typical subspecies (Fig. 2).
Due to the reasons as already stated it was impossible to attend to a detailed treatment of all slow-worm specimens from this group.
3.1.3.1. Kar openings
The variability of this character within the population with the domination of »colchi> CUS« characters was analysed on 42 specimens. The frequency of single variants is shown in Fig. 3 and ensues from the table below:
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Dominating in this group are individuals with clearly visible symmetrical car openings. The number of animals with an ear opening on one side of the head only or those without car openings is quite small.
3.1.3.2. Pileus
Proccntual frequency of single types of pilei in this population is established on 45 specimens.
Pileus Type	Pileus Type	Pileus Type
ABC	n
17.8 %	8.9 %	73.3 %	8 + 4 +33 = 45
Prevailing in this group of slow-worms are individuals with separated prefrontal plates. The pwrcentage of the animals with the B pileus type is surprisingly low, whereas the number of the specimens with the pileus characteristic of the western subspecies is somewhat higher. A graphic presentation of the results is revealed in Fig. 4.
3.1.3.3.	Blue spots
Relative frequency of the occurrence of blue spots was calculated for 20 spKicimens.
+ -	n
80.0% 20.0%	16 + 4 = 20
A tendency to an increasing frequency of blue spots in connection with the Yugoslav populations of slow-worms is most obviously expressed in the group with dominant characters of the Colchidic subspecies. By the presence of blue spots (80%) our population, together with the Slovakian one (80.3 %, LÄC 1967), distinguishes itself from the populations of this subspecies in other countries e. g. Bulgaria (68.7 %. BESKOV 1966) and Finland (73 %, VOI-PIO 1962). Relative frequency is presented in Fig. 5.
3.1.3.4.	Number of scales
In the population with dominant »colchicus« characters the proccntual frequency of the occurrence of a definite number of scales around the body was calculated on 43 specimens. The results of calculation are shown in the table below and Fig. 6.
24 25	26 27 28 29 30 31
7.0 % 4.6 % 34.9 % 16.3 % 23.3 % 7.0 % 4.6 % 2.3 % 3 + 2 + 15+7 + 10+3 + 2 + I =43
This population proved to have the widest variation range of the number of scales. The diapason of variation corresponds to the situation in Slovakia (LAC 1967) and Rumania
(STUGREN, FUHN, POPOVICI 1962). It is interesting to note that in Hungarv the variation range is narrower (DELY 1972, 1974). VOIPIO (1962) and BEŠKOV (1966) did not analyse this character. A specimen from the eastern slopes of the Rudnik highlands possessed a maximal n^ber of scales (31) stated for Yugoslavia. The arithmetic mean of this character amounts to X=: 26.909.
3.2. Comparative analysis
As pointed out already in the chapter on the material and methods, I applied the method of hi-square lest with Yates's correction to establish the differences among the populations identified on the basis ofthe calculated frequency of single characteristics as observed. Subject to comparison were all three qualitative diiTerential characters. Analysed were three existing possibilities of combining the populations.
The t-test was used to test the differences between the arithmetic means ofthe number of scales within the populations identified. 1 preliminarily calculated the standard error ofthe difierence between two arithmetic means.
3.2.1. Comparison of a ^Mip of individuals with the domination of »fragilLs« charactcrs with the popularion from the contact zone
The first comparison was made between the population ofslow-worms with the domination of the characters of the typical subspecies with the population from the contad zone. The results ought to give a response to the question on the type of delimitation between two »great« subspecies ofslow-worms, and to confirm or else refute whether the identification ofthe contact population was justified or not.
3.2.1.1. Ear openings
With respect to previous results (DELY 1972, 1974, BESKOV 1966 etc.) the existence of clear ear openings represents the stablest diflTerential character in the microtaxonomy ofslow-worms. therefore, a comparative analysis was introduced on the basis of this property.
A. f. li^Iis	A. f. fra^is $ colchicus
Ni= 186	Nj=42
Without ear. op. = 182	Without car op. = 38
With ear op. = 4	With car op. = 4
/2 = 2.382
As ensuing from the hi-square table, the limit value of;f^ at one degree of freedom attains a level of significance from 0.05 (5 %), x^ = 3.841. As our hi-squre is smaller, no significant difference exists between these two populations with respect to the presence of clear car openings.
3.2.1.2. Pilous
A. f. fragilis	A. f. fra^lis ^ A. f. cokhkrus
Ni = I92	N2 = 43
A=119	A=I8
B + C= 73	B + C = 25
X^ = 5.076
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3.2.1 J. Blue spots
A. f. fragUls	A. f. fragilis % A. f. cokrhicus
N. = 76	N2=23
Without bl. sp. = 53	Without bl. sp. = 6
With bl. sp. = 23	with bl. sp. = 17
X'=\2.555
Taking into account the feet that the hi-square value is much above the limit value, the populations as compared essentially differ from each other in this property.
3.2.1.4. Comparison of artithmetic means of the number of scales and test of the difference between them
A. f. fragilis	A. f. fragilis 5 A. f. colchicus
XI = 25.41= 25	X2 = 25.940 = 26
SI = 7.2= 7	S2= 1.604= 2
^'=•82 sx,-x2 = 0.62I	N. = 34
t= 1.610
On the threshold of significance of0.05 (1.96) the difference between the populations is not statistically significant. Not even on the threshold of significance ofO.OI (2.58) does the difference between the populations become important.
3.2.2. Comparison of the contact and the Cokhidic population 3.2.2.1. Ear openings
A. f. colchkrus	A. f. fragilis % A. f. cokhiciis
Ni=42	N2=:42
Without ear. op. = 2	Without car op. = 38
With ear. op. = 40	With car. op. = 4
X' = 58.46
The hi-squre value points to significant differences between the two populations when the most important character for idcntif>ing the typical subspecies is in question.
3.2.2.2. Pileus
A. f. cokrhicus	A. f. fragilis % A. f. cokhkrus
Ni=45	N2 = 43
A= 8	A= 18
B + C = 37	B + C = 25 = 5.024
The comaci populaiion importantly diflčrs from ihc Coichidic one also with respccl lo dcllniie pilcus l>pcs as rcprcseniLxl in ihis case. Though ihe diflcrcnce does not aliain Ihc difference level of ihe previous characler. ii is nevertheless significant.
3.2.2^1. Blue spots
A. f. cokhicus	A. f. fragilis ^ A. f. cokhicus
Ni = 20	N2 = 23
Without bl. sp. = 4	Without bl. sp. = 6
Withbl. sp.= 16	withbl. sp.= 17
/' = 0.010
The hi-square as obtained is coasiderably lower llian the significance threshold, which means that with regard lo the frequency of the ocx'urrence of blue spots no statistically significant difference exists between the populations compared.
3.2.2.4. Comparison of arithmetk means of the number of scaks and testing of the difference between them
A. f. cokhkus	A. f. fragilis % A. f. cokhicus
XI = 26.909 = 27	xj = 25.940 = 26
si = 3.444 = 3	S3 = 1.604 = 2
Ni=4l	N2 = 24
Sxi-X2 = 0.579 1= 1.727
At the threshold of significance ofO.05 (1.96) the difference between the populations is not significant. which is true also of the significance threshold of 0.01 (2.58).
3.2.3. Comparison of the t>picaJ and the Cokhidk population
We are finally going to compare two populations of slow-worms, one of which Is spread more lo the northwest and the olher to the southeast of Yugoslavia.
3.2.3.1. tlar openings
A. f. fragilis	A. f. cokhkus
Ni= 186	Nj = 42
Without car. op. = 182	Without car op. = 2
With car. op. = 4	With ear op. = 40
184.711
The limit value of hi-square at one degree of freedom on the significance level of 0.05 (5 %), X' = 3.841. The hi-square value is exceptionally high, which indicates that with respect lo the existence of car opening ihe differences between the typical and the Coichidic subspecies in Yugoslavia are most important.
3,2.3.2. Pileus
A. f. fragilis	A. f. colchicus
Nt= 192	N2 = 45
A= 119	A= 8
B + C = 73	B + C = 37
X^ = 26.89
Also in this character a marked difTercnce is to be observed between these two populations, since the hi-square value considerably exceeds the significance threshold.
3.2.3.3. Blue spots
A. f. fragilis	A. f. colchicus
Ni = 76	Ni = 20
Without bl. sp. = 53	Without bl. sp. = 4
With bl. sp. = 23	With bl. sp.= 16
= 14.240
The typical and the Colchidic subspecies differ least in the existence of blue spots in males, even though also in this case the difference between them is statistically most significant.
3.2.3.4. Comparison of the arithmetic means of the number of scales and testing of the difference between them
A. f. fragilis	A. f. colchicus
Ki =25.41 =25	xj = 26.909 = 27
Si = 7.2 = 7	S2 = 3.444 = 3
Ni= 182	Nj = 41
Sxi -X2 = 0.698 t = 2.865
The t-iest value indicates thai ihc difference between the populations compared is statistically significant, both on the trhreshold of significance of 0.05 (1.96) and that of 0.01 (2.58).
3.2.4. Comparison of the typical subspecies of slow-worms of Central Europe and Yugoslavia
In addition to testing the differences between the identified slow-worm groups from Yugoslavia I checked possible digressions of the f>opulations thus identified from those that undoubtedly make part of the typical and the Colchidic subspecies, respectively. The material to be compared with the typical subspecies was taken from DELY's works (1972,1974). The specimens originate from Centra! Europe (Austria and Hungary).
A comparison of the typical subspecies A. fragilis with our slow-worms classified into this group was made for all three differential characters and the number of scales around the body in one series.
3.2.4.1. Kar openings
A. f. fragllis
Yugoslavia Ni= 186 Without ear op. = 182 With ear op. = 4

A. f. fragilis Central Europe N2 = 52 Without ear op. = 51 With ear op. = 1
These two populations do not differ from each other with respect to the existence of ear openings.
3.2.4.2. Pileus
A. f. fragilis
Yugoslavia Ni = 192 A= 119 B + C= 73
A. f. Fragilis Central Europe Nj = 46 A = 32 B + C= 14
= 0.613
From a statistic point of view« the Yugoslav population of slow-worms, classified into the typical subspecies, does not differ from the typical subspecies from Central Europe.
3.2.4.3. Blue spots
A. f. fragilis
Yugoslavia Ni = 76 Without bl. sp. = 53 With bl. sp. = 23
= 0.02!
A. f. fragilis Central Europe
N^ = 23 Without bl. sp. = 15 With. bl. sp. = 8
Not even in the third qualitative differential character do the populations compared differ from each other.
Due to negligible differences in the arithmetic means of the number ofscales and an identical standard error between the arithmetic means in the case of the Yugoslav slow-worms classified into the first group and the indubitable A. f. fragilis from Central Europe, the t-test value amounted to t = 0.
3.2.5. Comparison of the Colchidic subspecies of slow-worms from Eastern Europe and Yugoslavia
Analysed in the above-mentioned works by Dely is also a material on A.f.colchiats from Eastern Europe (Rumania, Bulgaria, Ukraine, Poland, Russia. Gruziya, and Azerbaijan). With respect to the existence of blue spots 1 was not in position to attend to a comparison since Dely's material did not include a sufficient number of male slow-worms from these countries.
3.2.5.1.	Kar openings
A. f. colchicus	A. f. coichicus
Yugoslavia	I:aslcrn Europe
Ni = 42	N2 = ?6
Without car op. = 2	Without car op. = 3
With car op. = 20	With car op. = 33
/2 = 0.0316
The hi-square value does not attain the significance level (3.841 for one degree of freedom) so that no dilTercnces in the presence of ear openings exist between the slow-worm population from eastern Europe, making part of the Colchidic subspecies, and the slow-worm population from our country that I classified into this subspecies.
3.2.5.2.	Pileus
A. f. colchicus	A. f. colchicus
Yugoslavia	Eastern Europe
N. = 45	N2 = 60
A= 8	A = 23
B + C = 37	IJ + C = 37
= 4.278
In this character the slow-worm population from Yugoslavia that I classified into the Colchidic subspccics differs from the population of/1. ./. colchiam from eastern Europe.
For these two populations the results of calculating the arithmetic means of the number ofscales and the standard error between them were perfectly identical. No further calculation was necessary as the samples obviously do not differ in this feature. The t-value was nevert-heles calculated and proved to be equal to 0.
4. Discussion
The basic prerequisite in giving any sort ofjudgement on the taxonomy and biogeogra-phic characteristics of the species Afifiuis J'ra^ilis in Yugoslavia was to possess a sufficient number of specimens for the purpose of analysis. In my opinion the treatment of the Yugoslav population of slow-worms was carried out on a material satisfying the above prerequisite. For the sake of comparison I am stating the material which served as a basis for solving the problem of systematics and distribution of this species in the countries where these problems have been an objects of intensive studies. Of the neighbouring countries, the sample for an analysis of .1. fraailis from Bulgaria comprised 120 specimens (BESKOV 1966). from Rumania 55 specimens (STUGREN. FUHN & POPOVICI 1962). and from Hungar> 126 spe-cimcns(DELY 1972, 1974). In Slovakia the study was done on 105 specimens (LAC 1967). in Finland on6l (VOIPIO 1962), in Norway on l2(VOIPIO 1962), in Sweden on 101 (VOl-PIO 1962). in Holland on 138 (MUSTERS & BOSCH in den 1984) and in the Soviet Union on 46 specimens (LUKINA 1965).
The material as treated and the data from (he literature render it possible to get an insight into the present range of slow-worms in Yugoslavia, to respond to the question on the in-traspecific differentiation of the species in our country-, and to deliver a judgement on the geographic delimitation of the taxa within the species A. frafiHis.
When discussing the distribution of the species in Yugoslavia, the authors most usualK present generalized conclusions wherefrom it follows that it is spread everywhere with the ex-
ception of the islands. This probably results from a poor knowledge on the recent range of one of the most common species of reptiles in Yugoslavia, as well as a loo tepid an interest in the problems connected with its distribution.
The results 1 obtained, as well as the latest results of other authors (BRUNO 1980) reveal a number of new facts on the distribution of slow-worms in Yugoslavia. First, A. fragilis not live in certain parts of our country. The question is of larger areas ofVoivodina: the major part of Bačka and the whole of Banat where the cultured steppe offers no conditions for the life of slow-worms. However, two data refer to this area, namely: Kovilj and Titel (MOJSl-SOVICS 1897). In my opinion the localities in question might be connected with Fruska Gora (Mt.), a great dispersi9n center of slow-worms, and a possibility of survival of this lizard in gallery forests (DŽUKIČ 1980) - a possibility in which Mojsisovics, author of the afore mentioned data, did not believe. DŽUKIČ's presumption was confirmed by finding slow-worms in forests on the alluvium at Doroslov in Bačka in May 1985. It is highly probable that slow-worms are likewise absent from those areas that from a biogeographic point of view make part of the province of Aegean-Anatolian semideserts (MATVEJEV 1961) which in our country arc restricted mainly to some parts of Macedonia.
It used to be believed that A.fra^ilh does not inhabit the islands of the Adriatic Sea, with the exception of Isle of Kosljun (WERNER 1891). After P02:zrs (1966) generalized statement that it can be found on certain islands. BRUNO's work (1980) was published in which a series of localities is stated referring to the islands Cres and Krk. It is interesting to note that the localities on Krk from which our material originates do not coincide with BRUNO'S data. This points to the fact that on Krk the slow-worm is rather wide-spread. Included in this group of data isalso one most kindly communicated to me by Konrad KLEMMER from the Natur-Museum Senckenberg. It refers to a male from the Brioni Islands which is kept in the museum collection. In mybeliefa much greater meaning should be ascribed to an unexpected occurrence of slow-worms on the south-Dalmatian islands. The first finding is connected with Govcdjari on Island of Mljet and the second on Island of Vis. But the following datum is in fact unexpected and, I think, not perfectly proved. Namely: working on the collection of Prof. RADOVANOVIČ I found, in a pot. a juvenile specimen o^ A.frafiilis with a designation »Vis«. Signed on the label as collector is Anaf, and beside the slow-worm itself there was also a tail ofHemiJaciyhts lurdcus. In any case this means that the Dalmatian islands shoudl not be excluded from the Yugoslav range of slow-worms before it is proved that, as a matter of fact, on one of them this lizard does not live. The above corrections of the slow-worm range in Yugoslavia represent a contribution to a better knowledge of the actual distribution of the species, as well as a signpost to future investigations dedicated to the range and eventual changes in it.
In order to realize the foremosi objective of this work and to solve the problem of the mtraspecific differential ion oft he species An^uis fragilis and the horology oft he intraspecific taxa. an extensive analysis of external morphological characters was performed for the first lime on a material which made it possible to obtain valid results.
In accordance with my previous results as well as those of other authors the material available from the territory of Yugoslavia was divided into three groups. The first group consists of individuals characterized by the domination of the characters of the typical subspecies {A.f. fragilis). The second group is characterized by mixed characters with an indication that it originates from a contact zone between two subspecies, whereas the specimens marked by the domination of »Colchidic« characters were classified into the third group.
For every group in question I ascertained the variability of those qualitative and quantitative characters in the case of which differential nature has been proved in the history of investigating the microtaxonomic problems of the species. The procentual frequency of the occurrence of definite difTereniial characters is graphically presented in Figs. 3,4. 5, and 6. The results obtained by calculating the frequencies of definite qualitative and quantitative characteristics of the groups as identified reveal differences among them. The most marked
difference among the above-mentioned group is expressed in the existence ofclear ear-ope-nings. Thus, in the group of individuals with the domination of the characters of the typical subspecies only 2.2 % of animals had ear openings whereas in that with the domination of the features of the other »great« subspecies (A. f. cokhicus) ear openings appear in as many as 92.5 % of the specimens. Somewhat less pronounced differences reveal themselves in the presence of a certain type of pileus and the number of scales in a series around the body, and the least important differences in the occurrence of blue spots. In order to find out whether the differences observed among the groups identified are statistically significant, the method of hi-square test was applied in the case of qualitative features and the t-test method in that of quantitative characters. In the first case I had to applly YATES*s correction due to an unequal sample. I first compared the groups characterized by the domination of the characters of the »great« subspecies with the contact population, and then the two of them with each other
As proved by the analysis, no statistically important difTerences present themselves between the groups of individuals with dominant characteristics o^ A. f.fraRilis2kX\*l the contact population in the existence of ear openings and the number of scales around the body. The populations differ essentially from each other in the pileus type and the presence ofblue spots in males.
Comparing the individuals characterized by the domination of the Colchidic characters with the group of t he contact population one can take note of a great difTerence between these two groups in the existence of ear openings or the presence of a certain pileus lyF>c. as well as the fact that the differences in the presence ofblue spots and the number of scales arc not statistically significant.
The results of comparing all differential characters revealed important statistical differences between the groups of individuals which with respect to their characteristics make pari of one or the other »great« subspecies.
Beside testing the differences between the groups of identified slow-worms from Yugoslavia I also checked eventual deviations of the populations thus identified, with respect to the populations which undoubtedly belong to the typical and the Colchidic subspecies, res-F>ectively. The results show that there exist no statistically significant differences between the typical subspecies ofslow-worms from Central Europe and the Yugoslav population with the domination of the characters of A. f. fragUis^ that on the contrary, the differences are in fact negligible. A similar result was obtained also in the case of comparing A. f. colchicus of Eastern Europe with a group of individuals from Yugoslavia with prevailing characters of the eastern subspccies. The only difference between them was disclosed in different types of pilei. Namely, in the populations from the above-mentioned east-European countries the A type of pileus is more common than in the corresponding group of individuals from Yugoslavia. These differences are believed to result from the heterogeneity of the material from Eastern Europe, because no difTerences made themselves evident in comparing our data with the literature data from Bulgaria. Slovakia and Hungary.
The above results lead to a conclusion that in Yugoslavia the species fiagilis is differentiated as two subspecies: the typical -A. fragilisfragilis^ and the Colchidic - A. fiagilis colchicus. Existing in the zone of a secondary contact between the said populations is a population marked by hybrid characteristics, however, it comes somewhat closer to the typical subspecies.
The sübspccics A J'ragilisfiagHis is much more widely distributed in our country; its range includes Slovenia, almost all of Croatia and Bosnia, a large part of Herzegovina and Montenegro. as well as western Serbia and most of Kosovo (Fig. 2). Outside this compact pan of the range the subspecies appears also on the highlands of Serbia and Macedonia. A contact zone of uneven width, without clearly drawn limits (Fig. 2), lies between the typical and the Colchidic.subspecies.
The contact zone is narrower in the highland part of our country and broader in the lowlands, as well as the hilly region, ft is widest in Šumadija, Srem and Slavonia. Due to a lack of the material from the western parts of Slavonia and Podravina I was not in position to precisely define the said zone in this region. Narrow contact zones take the form of strips on highlands whose roots and environs are populated by the Colchidic subspecies. Our knowledge on the vertical differeniion of slow-worms in the eastern part of Yugoslavia has not as yet attained a level of definitive conclusions. However, the fact is that in the mountains the typical subspecies appears in the range oiA.fra}iUis colchicus. A fine example thereof is to be observed on Šar planina (highlands) where the Colchidic subspecies lives in the lowlands on the Macedonian and the Metohija side, while the typical on the mountain ridge: Brczovica and Pirib^ (our data), and Karanikola and Lukovo polje (KRIVOKAPIČ 1969). A similar situation is to be found in the region of northern Albania. However, the material available to me from Albania (see Table I) leads to a diflerent conclusion. This apparent contradiction results from the fact that DELY's material origites from highlands (Lura) and mine from lower sites of Albania. BEŠKOVs data (1966), as well as mine, referring to Stara planina may serve as another proof to this height differentiation. Here, on the peaks along the Yugoslav frontier appears the typical subspecies while the Colchidic subspecies populates the valleys of the Lom river and the environs of Zaječar.
The subspecies Aniiuis fragilis colchicus has a narrower but a most interesting distribution in Yugoslavia (Fig. 2). Its range includes the major part of Serbia, almost all of Macedonia, and a part of Montenegro and Herzegovina. The populations of slow-worms from parts of the ranges of the Colchidic subspecics in Herzegovina, Montenegro, and Metohija establish contact with the compact part of the range through the Albanian and the Greek populations (Table 5). The western delimitation of the range of the Colchidic subspecics A. f. colchicus in the mediterranean zone runs along the known turn of the Neretva river setting also a limit to the distribution of some east-mcditcrrancan elements of our herpetofauna {Stauremys caspica) and terriofauna {Talpa caeca. Piiymys thomasi and others, PETROV 1979), while transitional forms of this zone were slated also by M.^TVEJEV (1969)- birds, MIKSIČ.S.(l97l)-Orthoptera. DROVENIK (l973)-Coleoptera. and SIJAR1Č (1974)-Rhopalocera.
This subspecics has no disjunct parts of the range in the west of Yugoslavia, even though single occurrences of specimens (SELIŠKAR 1916; BRUNO 1976: BANK et al. 1982) have been recorded bearing the characteristics of/f./ colchicus. Such appearance of the characteristics of the eastern subspecies can be obtained by a »relay« passing-over of the genie material. by the appearance of ancestral characters in a determined genie combination or as a result of periodic pulsations of the range of the subspcncs in the post-glacial epoch.
Single occurrences of specimens phenotypically belonging to a subspecies in the range of another cause confusion among the authors (BANK ct al. 1982: MUSTERS & BOSCH in den 1982; HENLE 1985) who derived their herpeiological experiences from biogeograp-hically simpler areas so they tend to a complete simplification of the biogcographic situation, there at abstracting an exceptional biogcographic, geological, and orographic composition of the Balcan Peninsula, as well as the historical past of the herpetofauna. As a result ofthis tendency their conclusions on the problem of delimitation of the intraspecific taxa of A. fraf^ilis on the Balcan l^eninsula do not repose on forcible arguments. In addition to the above, the lack of a representative sample induces MUSTERS & BOSCH in den (1982) to seek the arguments for their conclusions by disputing undoubtedly unreliable consideration of these problems (POZZI 1966) or by questioning the validity of the data which geographically do not make part of the Balcan Peninsula (LAC 1967) or are rather distant from it, as in the case of Finland (VOIPIO 1962). It is not clear why in their analysis they do not discuss the results of BEŠKOV (1966), FUHN (1961), FUHN & VANCEA (1961), STUGREN et al. (1962). Not even the results of DELY (1972, 1974) to which they refer are interpreted quite correctly
as ihey overlook the existence of a »mixed« population between the typical and the Colchidic subspecies on the Dunazug highlands.
Unacceptable is also their rigid attitude to the »purity« of the subspecies where in spite of a generally accepted belief no broader diapason of variability is acknowledged within the populations making part of one subspecies.
The results thus interpreted were summarized by MUSTERS & BOSCH in den (1982) in a map which, they believe, represents »the real state of affairs« but which» in an extremely schematized representation where precise geographic positions and even less so their altitudes. as well as animals with »mixed« characteristics arc not duly taken into account, instead of classifying things leads to complete confusion. Generally speaking, their conclusions abound with presumptions which should not serve to identify the subspecies A. f. colchictis. which collides also with the essence of their remarks on the previous results.
The crucial remark of MUSTERS & BOSCH in den (1982) is that the authors opted for different characters, which represents an insurmountable obstacle to the drawing of concrete conclusions. As this remark is not supported by suitable arguments, it is our belief that in the previous analyses there are not concerned options for different characters but merely single instances of the omission of one of the characters which, if duly studied, would undoubtedly contribute to a greater importance of the analyses. I am convinced that in spite of the absence of absolutely identical analyses we are witnessing a positive trend in solving the problems of microtaxonomy and biogeography of slow-worms through a series of successive, complementary phases bringing us closer to ultimate solutions.
In principle, the explanation of the range of the two subspecies of slow-worms in Yugoslavia such as disclosed in this work, reposes on the theory put forward by VOIPIO(1962), irrespectively of certain inconsistencies and isolated beliefs (MUSTERS & BOSCH in den 1982) that the theoo' is precocious. VOIPIO (1962) presumed that in the period od glacia-lionsa unique population of slow-worms from Central Europe divided itselfinto a »western« population (A) finding a refuge on the Pyrencan Peninsula, and an »eastern« population (B) that withdrew to Persia and Asia Minor (Fig. 7). In the time of glaciations the species was divided into subspecies two »great« of which established a secondary contact with Central Europe at the end of the glacial period. Taking into account the results of this work it is important to note that Voipio. as well as the majority of other authors, believes that the contact of the subspecies was brought about on the eastern slopes of the Alps and that it is there that A. f. frafiilis is delimited from A. f. cokhicus. A certain progress of the contact zone in the direction of the east can be observed in LAC (1967). whereas a considerable, though a speculative one in DELY (1981). An obvious contribution to the perfection of VOIPIO's theory (1962) was presented by BEŠKOV (1966). He was the first to show that with regard to the results of slow-worm investigations in Bulgaria and Rumania, during one of the glaciations, the typical subspecies must have populated almost all of the territory of Central Europe and the Balcan Peninsula, but that as a result of a later penetration of the Colchidic subspecies, assisted by a global moderation of the climate, the typical subspecies retreated on one side towards the west, and on the other to the mountains. At the same time BEŠKOV (1966) was the first to raise the question of autochthonous populations originating from the Tcniar>' ones (BOLKAY 1913. MLYNARSKI 1962). and their infiuencc upon the recent phenotype of slow-worms on the Balcan Peninsula.
My results of 1980. as well as those disclosed in the present work, largely support BEŠ-KOV's amendment of VOI PIO's hypothesis in addition to geographically defining the border between the typical and the Colchidic subspecies which, with respect to the material available to him. BEŠKOV was not in position to do. Likewise substantiated are indications that the delimitation of the typical and the Colchidic subspecies in Albania and Greece might be similar. Though as yet not possessing concrete prooft. we believe that on the highlands of northern Greece, more precisely, in the habitats populated by Laceria aailis and Vipera hcrus
(CHONDROPOULOS 1986, ONDIRAS 1968) slow-worms make part of the typical subspecies.
There remains an insufficiently eiucidated origin of the Peloponnesian subspecics (A.f. petoponnesiacus) AT\d its link, originally, with the autochthonous populations which undoubtedly had absolute chances to survive on the Greek land, and secondarily, with the arriver i.e. the Colchidic subspecies, taking into aaccount a broad contact zone between Peloponnesus and the Greek mainland which existed several times for relatively long periods, thus also in the period of the last glaciation in the Valdai epoch (GERASIMOV & VELICHKO 1982).
5. Conclusions
The study of the Yugoslav population od At\guis fra^ilis was carried out on 307 specimens, a sample that made it possible to obtain valid results. Researched is the distribution ofthis species and its intraspecific differentiation in ourcountry andon the Balcan Peninsula, as well as the biogeographic characteristics of the intraspecific taxa.
Fig. 7. Penetration of the typical and the Colchidic subspccics of the slow-worm (Angnis fragilis L.) to the north after the glacial period. According to VOIPIO (1962).
Slika 7. Smeri prodiranja nominantnc in kolhidske podvrste slepca {AnguisfrasUis L.) na sever po ledeni dobi (po Voipiu 1962).
Defined is the present slow-worm range in Yugoslavia, a range that does not cover the entire territory but includes larger areas in the cultured steppe ofVoivodina from which this species is absent, in addition to a strong probability that A. Jragilis does not populates such provinces that biogeographicaily make part of the subprovince of Aegean-Anatolian semi-deserts {MATVEJEV 1961) which in Yugoslavia are limited to single parts of Macedonia. Slow-worms, in some cases in large numbers, were proved to be present on single islands of the northern Adriatic, whereas the first mention of Anguis J'ragilis is connected with the south-Dalmatian islands Mljet and Vis.
On the basis of the frequency of positively linked intraspecific characters the Yugoslav population of slow-worms is divided into three groups. The first group consists of individuals with dominant characters of the typical subspecies {A. f fragilis), the second of individuals of the contact population, and the third of specimens of the Colchidic subspecies.
By analysing qualitative and selected quantitative characteristics we established their variability within each group as indentified while the hi-square and t-test methods were applied to determine the statistical significances of the difTerences observed among single groups. It can now be stated that both the typical and the Colchidic species exist in Yugoslavia. A dominant difference between them expresses itself in Yugoslavia. A dominant difference between them expresses itself in the presence of ear openings, a lesser one in the number of scales around the body and the pileus type, and the least, though statistically an important difference in the presence of blue spots.
By testing the differences between the slow-worm groups from Yugoslavia and the population of A. fragilis from the neighbouring countries, undoubtedly making part of the typical and the Colchidic subspecies, respectively, we proved that our popuations wholly belong to one of the two »great« subspecies. The sole difference in the frequency of the type A of the pileus was observed between the Yugoslav specimens of the Colchidic subspecies and the specimens of this subspecies from an area of Estern Europe as kept in DELY's collection. Such difTerences were not recorded when comparing our specimens with the material from Bulgaria, Slovakia, and Hungary.
Defined was the range of A. f./ra^ilis; we also proved that in Yugoslavia this subspecies is more widely spread than the Colchidic one, and that its range comprises Slovenia, almost all ofCroatia and Bosnia, a considerable part of Herzegovina and Montenegro, as well as western Serbia with a larger part of Kosovo. Outside the compact part of the range the subspecies appears on the highlands of Serbia and Macedonia, which is stated also in the »Catalogus Faunae Jugoslaviae« (BRELIH & DŽUKIČ 1974) and which HENLE (1985) erroneously associates with the highlands of the western part of Yugoslavia.
Even though of a narrower span, the range of the Colchidic subspecies (A. f. colchLsuc) has more indented contours. It covers the major part of Serbia, almost all of Macedonia and parts of Montenegro and Herzegovina. In Herzegovina, Montenegro and Metohija the slow-worm populations of the Colchidic subspecies come into contact with the compact part of the range through Albanian and Greek populations. In the mediterranean zone the western border of the range the Colchidic subspecies runs along the turn of the Neretva river. This subspecies has no disjunct part of the range in the west of Yugoslavia in spite of the fact that single occurrences of specimens with phenotypic characteristics of A.f. cokhicus have been recorded.
Where the typical and the Colchidic subspecies encounter each other the existence of a »contact« population can be established. The geographic position of the contact zone forms also the delimitation line between the typical and the Colchidic subspecies. The contact zone is narrower in the mountainous part of Yugoslavia and broader in the lowlands and the highlands. Narrow contact zones take the form of strips on the highlands whose roots and larger environment are populated by the Colchidic subspecies.
The delimitation among the subspccies such as evidenced for Yugoslavia and Bulgaria (BEŠKOV 1966) is valid for the entire southeastern part of the Balcan Peninsula.
Al present the most acceptable to ihc explanation of the intraspecific difiercnliation of the species Anguis J'ragilis and the horology of the intraspecific taxa in Yugoslavia and on the Balcan Peninsula are VOIPIO's hypothesis and an amended division of the Balcan Peninsula by BESKOV (1966), respectively.
Summary
The object of our study is the variability of morphological characters of the species An-ffuisfragilis LINNAEUS 1758 (Reptilia: Sauria: Anguidae) in Yugoslavia nad on the Balcan Peninsula. Included in the research were 307 specimens. It was found that the geographic distribution of external morpholigical characters of slow-worms was not homogeneous. A detailed study of the variability of outer ear openings, the number of scales in a series around the body, the pileus type, and the existence of blue spots served to prove the presence of the typical {A. f. fragilis) and the Colchidic {A. f. colchicus) subspecies of slow-worms in Yugoslavia, with a contact zone between them. Described are characteristics of these populations, in addition to a comparative analysis, both with respect to the identified groups of populations in Yugoslavia and the populations from different parts of Europe which undoubtedly belong to the typical or the Colchidic subspecies of slow-worms.
Moreclo.sely determined are ranges of the subspecies in our country, and defined is also the contact zone. The typical subspecies was found to cover a larger range in our territory including Slovenia, almost all of Croatia and Bosnia, the major part of Herzegovina and Montenegro, as well as western Serbia and Kosovo. In addition to the compact part of the range the said subspecies appears also on the highlands of Serbia and Macedonia. The range of the Colchidic subspecies encompasses the eastern and the southeastern part of Yugoslavia, while from Greece and Albania a part extends to Montenegro, Herzegovina and southern Dalmatia. Extending between these populations i.e. subspecies is a contact zone of an uneven breadth and without sharp outlines. It is broader in the highland parts and the lowlands of Šumadija. Srem and Slavonia. The contact zone takes also the form of narrow strips on the highlands with isolated parts of the range of the typical subspecies.
Beside the ranges of the subspecies this works ofTers also an insight into the present range of the species An git i s fragil is in Yugoslavia. The species was found to be absent from larger spans of the cultured steppe in Voivodina and it is presumed that it likewise does not live in the provinces of Aegean-Anatolian semi-deserts in Macedonia. The species was stated for the first time on the south-Dalmatian islands.
The most acceptable to the explanation of such a type of the slow-worm range in Yugoslavia and the Balcan Peninsula, as well as its historical formation is V0lP10*ss hypothesis (1966) amended by BESKOV (1966).
Povzetek
Proučevana je variabilnost zunanjih morfološk-h značilnosti vrste Angtiis fragilis LINNAEUS 1758 (Reptilia: Sauria: Anguidae) v Jugoslaviji in na Balkanskem polotoku. Proučevanje je zajelo 307 primerkov. Ugotovljeno je, da ni homogene geografske distribucije zunanjih morfoloških značilnosti. Natančna obdelava spremenljivosti zunanjih ušesnih odprtin, števila lusk v enem nizu okoli telesa, tipa pileusa in plavih peg je dokazala nominantno {A.f. fragilis) in kolhidsko {A.f. colchicus) podvrsto slepca v Jugoslaviji in njuno medsebojno stično območje. Opisane so značilnosti teh dveh populacij, narejena je komparativna analiza med tema skupinama populacij kot tudi med populacijami iz različnih delov Evrope, ki nedvomno pripadajo nominantni ali kolhidski podvrsti.
Bolj natančno so določeni areali podvrst v naši deželi in njihovo stično območje. Ugotovljeno je, da ima nominantna podvrsta širši areal na našem ozemlju, ki zajema Slovenijo, skoraj celo Hrvatsko in Bosno, večji del Hercegovine in Črne gore, zahodno Srbijo ter Kosovo. Razen na tem strnjenem območju se ta podvrsta pojavlja na visokih planinah Srbije in Makedonije. Areal kolhidske podvrste zajema vzhodni in jugovzhodni del Jugoslavije. En del areala kolhidske podvrste zahaja preko Grčije in Albanije v Črno goro, Hercegovino in južno Dalmacijo. Med temi populacijami, pravzaprav podvrstami, je stično območje brez ostrih meja in neenake širine. Sirse je v hribovitih in nižinskih predelih Šumadije, Srema in Slavonije. Stično območje se pojavi kot ozek višinski pas na planinah, kjer so izolirani deli areala nominantne podvrste.
Razen areala podvrst je v delu podana tudi analiza areala vrste Angitis fragilis v Jugoslaviji. Ugotovljeno je, da vrste na širšem prostoru kulturne steF>e v Vojvodini ni in domneva se, da je ni na območju egejsko-anatolskih polpuščav v Makedoniji. Vrsta je prvič ugotovljena na južnodalmatinskih otokih.
Za obrazložitev takšnega tipa areala slepca v Jugoslaviji in na Balkanu kot tudi njegovega zgodovinskega nastanka je sprejemljiva Beškovljeva (1966) dopolnitev hipoteze Voipia (1962).
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Conienis
Introduction....................................................................................................................................................................................................................................................I
Acknowledgements..............................................................................................................................................................................................................................2
Survey of the past investigations dedicated to the intraspecific variability of the species
A./ragiiis from Yugoslavia......................................................................................................................................................................................................2
Material and methods ....................................................................................................................................................................................................................4
Results......................................................................................................................................................................................................................................................................5
1.	Variability of basic difTerentia! characters ......................................................................................................................................................5
1.1.	Groups of individuals with dominant charactcrs of the typical subspecies ..............................................10
1.1.1.	Ear openings..................................................................................................................................................................................................................................................19
1.1.2.	Pileus..........................................................................................................................................................................................................................................................................20
1.1.3.	Blue spots............................................................................................................................................................................................................................................................21
1.1.4.	Number of scales....................................................................................................................................................................................................................................21
1.2.	Group of animals from the contact zone of the typical and the Colchidic subspecies ..........22
1.2.1.	Ear openings..................................................................................................................................................................................................................................................23
1.2.2.	Pileus..........................................................................................................................................................................................................................................................................23
1.2.3.	Blue spots............................................................................................................................................................................................................................................................26
1.2.4.	Number of scales ....................................................................................................................................................................................................................................26
1.3.	Group of individuals with dominant charactcrs of the Colchidic subspecies........................................26
1.3.1.	Ear openings..................................................................................................................................................................................................................................................29
1.3.2.	Pileus..........................................................................................................................................................................................................................................................................29
1.3.3.	Blue spots............................................................................................................................................................................................................................................................29
1.3.4.	Number of scales....................................................................................................................................................................................................................................30
2.	Comparative analysis ...........................................................................................................
2. i. Comparison of a group of individuals with the domination of>»fragilis« charactcrs with the
population from the contact zone ..............................................................................................................................................................................30
2.1.1.	Ear openings ..................................................................................................................................................................................................................................................30
2.1.2.	Pileus..........................................................................................................................................................................................................................................................................30
2.1.3.	Blue spots............................................................................................................................................................................................................................................................32
2.1.4.	Comparison of arithmetic means of the number of scales and test of the diflemece between them ............................................................................................................................................................................................................................................................................32
2.2.	Comparison of the contact and the Colchidic population....................................................................................................32
2.2.1.	Ear openings..................................................................................................................................................................................................................................................32
2.2.2.	Pileus..........................................................................................................................................................................................................................................................................32
2.2.3.	Blue spots............................................................................................................................................................................................................................................................33
2.2.4.	Comparison of arithmetic means of the number of scales and testing of the difference between them ......................................................................................................................................................................................................................................................33
2.3.	Comparison of the typical and the Colchidic population ....................................................................................................33
2.3.1.	Ear openings ..................................................................................................................................................................................................................................................33
2.3.2.	Pileus..................................................................................................................................................................................................................................................34
2.3.3.	Blue spots............................................................................................................................................................................................................................................................34
2.3.4.	Comparison of the arithmetic means of the number of scales and testing of the difference between ihcm................................................................................................................................................................................................................................................34
2.4.	Comparison of the typical subspecies of slow-worms of Central Europe and Yugoslavia	34
2.4.1.	Ear openings ..................................................................................................................................................................................................................................................35
2.4.2.	Pileus..........................................................................................................................................................................................................................................................................35
2.4.3.	Blue spots............................................................................................................................................................................................................................................................35
2.5.	Comparison of the Colchidic subspecies of slow-worms from Eastern Europe and Yugoslavia ..........................................................................................................................................................................................................................................................................35
2.5.1.	Ear openings..................................................................................................................................................................................................................................................36
2.5.2.	Pileus..........................................................................................................................................................................................................................................................................36
Discussion ..........................................................................................................................................................................................................................................................36
Conclusions......................................................................................................................................................................................................................................................41
Summary..............................................................................................................................................................................................................................................................43
Povzetek................................................................................................................................................................................................................................................................43
References..........................................................................................................................................................................................................................................................44