Ytf ISSN 0351 — 0077
PRIRODOSLOVNI MUZEJ SLOVENIJE MUSEUM HISTORIAE NATURALIS SLOVENIAE
SCOPOLIA
Botanies
Geologica &
Palaeontologica Mtiseologica 5 Zoologies
Boris KRYSTUFEK
Forest Dormouse Dryomys niiedula
{PALLAS, 1778)
— Rodentia, Mammalia — in Yugoslavia
Drevesni polh Dryomys niteduia (PALLASj 1778) — Rodentia, Mammalia V Jugoslaviji
SCOPOLIA I No 9 [ pp. 1—36 Ljttbljana
Oct. 19S5
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FOREST DORMOUSE DRYOMYS NITEDULA (PALLAS, 1778) - RODENTIA, MAMMALIA -IN YUGOSLAVIA
KHYSTUFEK Boris Yu ßiooo Lijubljana, Prešernova 20 Natural History Museum of Slovenia sprejeto: 14. 3.1985
XJDC (UDK) 5B9.32S.2{4D7.1)(M5)=20Dn/omi;smtedtila
IZVLEČEK — DHE VESNI POLH dryomys W/TECH/L^ (PALLAS, 1778) — RODENTIA, MAMMALIA — V JUGOSLAVIJI — Areal -drevesnega polha Hryomys utfedwJo se v Jugoslaviji sklada z glavnimi ©arskimd masivi (Alpe, Dinar Idi, Sarska-Piindako in Karpatsko-Ealkamako gorovje), medtein ko živi v Maikedonijl tudi v nižavju. Vižinski raapofn mahaJallSi je med 60 in 2000 m nadmorske vdiine.
Na ozemlju Jii^oalavije (preponnaimo dve podvrsti; D.»;, Intermetiim (Alpe. severni Dlnanidl) in D. n. favi^ojla (Maikedtoffiiija). Raiilikiijeta se po riäanl loibanje 1 bulami. Večja je iprl i>odvratf ravijojla. Drevesni ,pcdhi 6z Hercegovine, Orne gore in Kosova kažejo anake iprehioda med obema (podvrstama.
ABSTRACTS — The area of the foreat dormouse	niteditla in Yugoslavia
mostly coinciideE with tlie prindipal nwuntain massifs (.the Alps, the Dinaiic Alpe, the Pin die and the Carpatho-Balcaii Tnauntain regions), 'Whereas in Macedonia it lives also in ithe lowland, The vertical nange of the localittea is between 60 and 2000 m.
In the tecrMOT^ of Yugoalaraia two subspecies were identified: l>,n. intermedius (the Alps, the NOTthem Dinaric Alps) aiid D. n. rouijajla (Macedonia). They diEEer fnwn each oither toy the liraJinoaise heiight iper JjuHae Virhich is larger liin the sub^äes ravijojla. Foreist darmice from Herzegoivina, Montenegro -and Kosovo display signs of transiti<m between the two subspecies.
CONTENTS — VSEBINA
Introduction ....
1.	Material and Methods
1.1.	Material.....
2.	Distribution . , .
3.	Variability .... Age Variability , .
3.2.	Sexual Variability .
3.3.	Individual Variability
3.4.	Geograplilc Variability
4.	Subspecies of d, n. in Yugoslavia , ...........
2 2
4
5
7 !) 10 10
5.	Vertical Distribution.....18
6.	Habitat Requireinents . . , ,	22
7.	Dominance .	36
8.	Period Of Activity......30
0.1. Heproduction........30
9.	Conclusions and Discussion . .	31 AclinoWiedgements ......	31
Povzetek..........34
References.........35
Intioduction
The forest dormouse Dryomys niteduJc {hereinafter D. n.) is one of the less known mammals ol tlie Yugoslav fauna. Dae to its looks and unfrequent apprerance it tias always attracted the attention of mammalogists. Thus, one often comes across short notes concerning new localities of ta^conomic problems but all of them, with no exception, are based on a small n-omber of specimens, frequently on one single animal. In his study on the TyroUan subspecies of the forest dormouse fD. n. iTitermedius) SCHEDL (1968) included also the northwestern region of the D.n, area in Yugoslavia (Slovenia and Croatia), however, for our territory no new, unpublished data were at his disposal.
From the territory of Yugoslavia two new subspecies were described (D, ti. ravijoila P ASP ALE V, MARTINO and PEC HE V, 1952 and D. n. diamesus LEHMANN, i9&9), but so far their taxonomic status has not been elucidated in detail. On the whole, our knowledge on the infraspecific differentiation is rather poor, for all conclusions have so far been ^sed on few specimens only.
Some scattered data exist also with regard to the selection of the habitat and the vertical distribution of D. ti. However, the species area of the territory of Yugoslavia as a whole remains undetermined.
1. Material and Methods
51 specimens of D.n. were examined from the entire territory of Yugoslavia, One part of the material used in. this study is kept at the Natural History Museum of Slovenia (PBIS). The other part was most kindly given at our disposal by Dr, B, PETBOV (Belgrade), for which we sincerely thank him.
Most of the material (91,7 •^/o) was caught into snap traps. Of the animals caught the skull and usually also the skin have been preserved. On the animals just caught the following dimensions were measured;
1.	Weight (in grams)—W
2.	Head and body (from the snout to the anus) — HE
3.	Tail length (from the anus to the tail tip) — Tl
4.	Hind foot length (without daws) — Hf
5.	Ear length — E
Seventeen linear measurements were taken on each of the skulls (Fig. 1) using a vernier caliper with an accurancy to the nearest 0,1 mm. Their definitions and symbols are as follows:
1.	Profile length — PL
2.	Condylobasal length — CbL
3.	Rostrum length — RoL
4.	Maxillary tooth-row length (alveolar) — MxT
5.	Diastema length ^ DiL
--a — J
I
f4
I

----,5^-J
FUg. 1. Method of iaMng skuU measurements of Dryomyi nifeduia (see the test for
further explanation) SI. 1. Natin merjenja lobanje pri drevesnem polhu. Za raala^o glej tekst
6.	Braincase breadth — EcB
7.	Zygomatic breadth — ZyB
8.	Breadth over mollars — MoB
9.	Rostrum breadth — RoB
10.	Interorbital constriction — IC
11.	Braincase height — BH
13. Braincase height per bullae — BbH
13.	Bullae length —BuL
14,	Rostrum height — RoH 13. Mandible length — MdL
16. Height of ramus mandibulae — RMH IT, Mandibular tooth-row length — MdT
The following quotient indices were calculated:
L Zygomatic breadth X 100 : Condylobasal length = QH 2. Braincase height per bullae K 100 : Condylobasal length = QI2 3> Braincase height per bullae X 100 : Braincase breadth = QZ3
4.	Rostrum length X 100: Condylobasal length = QI4
5.	Bullae length X 100 : Condylobasal length = QI5
6.	Height of ramus mandibulae X 100 : Mandible length = QI6
7.	Braincase breadth X 100 : Condylobasal length = QI7
8.	Mandible length X 100 : Condylobasal length = QI8
Calculation was made of average values (x), standard deviations (SD) and coefficients of variation (CV).
1.1, MaterUl
Slovenia: Julijske Alpe, Planica, 1000 m, 12,—19. 8. 1932, 3 (J <5 1 $ (PMS 2611, 2636, 2639, 2643); 8. 1963, 1 5 1? (PMS 4423^) leg. A. and B. KRYSTUFEK Jelovica, Goška ravan, 970 tn, S5,—30. 1903, 1 5 {PMS 3224) leg, I. KHYSTUFEK Pohorje, Velika Kc®«, 3520 m, i. 7. 1953, (PMS 3430) leg. B. KHYSTUFEK ßakitna, Smrekovec, 750 m, 17. 9. J968, 2 sex? (PMS 4388—&) leg. F, l^BEM; Borovnica, 23. 9. 196B, 1 sex? (PMS4390) leg. F. LEBEN; Snežnik, SvinSčakl, 13431«, ja, 7, 1967, 1 299 (BP 65/67) leg. B. PETROV.
Croatia: Hisnjak, 1450—1500 m, 9. B. 1967, 3 ^ <5 (BP 296—297/67) leg. B, PETROV; VelebJt, Zavižan, 1500—1650 m, 1 S (BP 26/71) leg. G. DZUKIC; Velebit, Alan, 1300 m, 15. 6. 1976, 19 (PMS 807) leg. B. KRYSTUFEK.
Bosnia and Herzegovina: Sator, Satorsko jezero, 1550 m, 5. 9. 1954, 1 d (PMS 4174) lee. B. KavSTUFEK; KupreSka vrata, Eaprdvnica, PoriCe, 8OB m, 10. 9. 19B3, 1 (J (PMS 3616) leg. B. KRYSTUFEK and V. POPOV; Z^engara, OrOovačko jezero, IS00 m, 9. 1984, 1 9 {PMS 4313) leg. V. POPOV; Magill, Prljevor, 1700—ISOOm, 10. 3. 1970, 13 299 {BP 216—218/70) leg. B, PETROV.
Montenegro; Durmitor, Cmo jezero, 1450 m, B.—0. 6. 1083, 3 99 (PMSM04, 3415—6) leg. B. KRYSTUFEK; Bjelaisica, Jelovica, 1450—1500 m, 16,—17. 7. 1976, 2 <3 <3 (PMS 592—3) leg. A. and B, KRYSTUFEK; SO. 9. 1981, 1 5 (BP 130/81) leg. B. PETROV; Komavi, Kom Vasojevii, 1800m, 16. 7. 1978, l9 (PMS 1012) leg. B. KRYSTUFEK; Orjen, Crkvice, 1050m, 1 9 Isex? {PMS 1021—2) leg. B. KRYSTUFEK,
B. Kiyätuf^äi: Fora&t Dta-mouse Dryomi^i mfedtila (PALLAS, 1170)... 5
Serbia; 2:jeb, Kula, n&O m, 14. 6. 1970, 1 S 1? (Bp03—9/77) leg, B. PETÄOV; Sar planina. Črni Kamen, KaCanik, 1300 m, 13. 1940, 1(3 <BP 116/40) leg. B, PETROV; Niä, SidevaČka ki is ura, 300 m, 26. 9. 136D, 1 sex? (BP 303/09) leg, b. PETROV.
Macedonia: Prilep, 700—BOO m, 21. 1563. 6 5 (Bp 53—03/00) leg. B, PETROV; Prcspanslio jezero, Asamati and Kurbinovo, 800—820 m, 16,-23, 7, 1977, 3 9$ (PMS 7no—leg. A. and B. KRYSTUFEK; Kajmaktalan. Eeder (5—7 km below), approx, 1000 m, 33. 9. 1373, I? [BP 79/72) leg. B. PETROV; Gevgeli, near Vardar, OD m, 26. 4. 1971, 2 3 5 {BP 33—4/71) leg. B. PETROV; Valandovo, AnaKa reka, 65 m, 7.-8, 4. 1968, 2 <5 <3 (BP 16/68, 33/68) leg. B, PETROV; OvSe pole, Sv. Nikole, 400 m, 17. 4. 1963, 1 (3 (BP 13/69) leg. B. PETROV.
2. Dlstfibutioii
The data on the diatributian of D. n. in Yugoslavia can be found in the works of FREYER (1842), PANCIC (1869), MILLER (1912), PASZLAVSKY (1918), BOLKAY (1924), WETTSTEIN (192B), MARTINO (1934, 193S), ZALESKY (1938), PETROV (1939, 1969, 1979), LEHMANN (1959), HENEBERG et al. (1964), FELTEN and STORCH (1965), MIRIC (1966, 1931), DULiC and VIDINI(1967), STORCH (1978) and KRyStUFEK (1979, 1&82). They are supplemented by private data as given at our disposal by Dr. B. PETROV, the data from the files of the collections kept at the Natural History Museum of Slovenia and those from the Collection Martino of the Zoological Institute, USSR Academy of Sciences, Leningrad (Fig. 2).
Let us shortly comment on some older citations;
For the territory of Yugoslavia ("bei R^ifnitz" - near Eibnica) H. FREYER (1842: 2) was the first to mention D. ti. under the name of Myaxv-s nitelta. DESCHMANN (1866: 214) reexamined the specimen on which FREYER's determination was based and came to the conclusion that Myojcu^ dry as (= D.n.) was in question. So FREYER doubtlessly han a D. n, in his hands and not Eliomys quercinus as erroneously stated in the literature,
For the forests of eastern Serbia PANCIC (1869: 82) cites Myoxus nitella and M. dryas, whereas Eliomys quercinus does not live in Serbia (PETROV 1979: 117).
MOJSISOVICS (1897; 183) and PASZLAVSKY (1913: 39) cite D.n. for Voivodina. Their citations are not supported by concrete data and have not been, confirmed ever since. In the list of Serbian mammals PETROV {1979: 117) no longer mentions this species in connection with Voivodina. Thus we can say that no proof exists on the presence of D. n. in the lowland north of the Sava and the Danube,
MILLER (1912: 569) mentioned D.n. for Belgrade ("Serbia: Belgrade"). The material (two females) was collected by Dr. H. B, S HARPE. This citation, too, has never been confirmed in spite of an intensive research in this area (for a longer period of time also Prof, MARTINO participated in this work). It is possible that the two specimens originated from the surrounding countryside of Belgrade. Thus, for instance, D. ti. has recently been found in Der dap (MIEIC 1931; 36) i. e. about 10(J km to the east of Belgrade,

Pig 2. Disfcribirtdonail records of Dryomys niteiivla in Yugoslavia SI. 2, NahajailäSa drevesnega poJha v Jugoslaviji
1. St Wolfgang (= Boifedk), 700 m (ZALESKY 103G); 2. »use (HANEBERG &t al. 1364); S. Vdika Kopa, 1520 m; 4. Mežica (M. KUNC in titt); 5. Jarvornilt (the files of the Natural History Museum of Slov.); 6. Ratete [ZIN-L); Planica, 1000m; 7. Trenta (S. BRELIH in U«.); a. Jeloviica. GoSka ravan. 970 m; 9. Boroviidca; Haikitna, Smreikovec, 750 m; 10. HeoifiniHz (= Ribnica), 491 m (FREYER 1S43, DESCHMAWN 1S83>; 11. Forests Of Kočevje Ko^«vsiki gixsäi) (the filas Of the Natural History Muiiemn of Slav.); 12. Snežariik, Svln£6alkii, 1242m (B, PETROV in Jdtt.); 13. Cabar (-Cataar), 550 m (ZALESKY 1938); 14. Riisnjak, H50—1550 m (B. PETHOV in Ktt,); 15. VÄlehit, Zai/Iišan, 1500—1650m (B. PETROV m litt,); Krasno (ZIN-L); 16. Velebit, Alan, 1300 m; H. Velebit, StirovaCa, approx. 900 m (WETTSTEIN 1020); 18. Sator, Satorsko jezero, 1550 m (BULlC et VIDINIC 1907); 19. Kuprežka vrata, Koprivnica, Poride, SSO m; 20. Veliki Igman, Veliko polje, 1340 m (ZIN-L); Gr kali ca (ZIN-L); 21. Tara Mts, (STORCH 1978); 22, Gola Jahüfina, 1800 m (ZIK-L); 24. Tr^afcawica, Kutski grad, 1450 m (2IN-L); 24. Zelengora, OrloVa^ko Jezero, 1500 m; 25. Maglič Pri je vor, 1700—1800 m (B, PETROV in litt.); 26. Kalinovik, Vlaholje (ZIN-L); 27. Prenj, Cmo polje, 1300 ni (B. PETROV in litt.); 2ß, Durmi tor, Valoviti Do, 2000 m (PJrrHOV 1939); Crino jezearo. 1450m; Grepifljna ipoljana, 17Q0m; 29. Bjelasica,
B, KryStufak: Forest EJoraiouse Dryomys mted«la (PALLAS, 177B)... 7
Jelovica, 1450—1500m (KRYSTUFEK 1979); 30. Komovl, Kom Vasojevl^, 1800 m (KRYäTUFEK. 1970); 31. Or]en, Crkvice, 1050 m (KEYStUFEK 1979}; 32, LovCen, Ivanova Korita {LEHMANN 1959); 33. RiJgOTO, Lisic (ZIN-L); 34. Žljeb, KuU 1750m (B. PFTEOV in litt,); 35. Sar planina. Črni Kamen, Kačanik, 1300m (PETROV 1969>-, 36, Pri aren. Kopane vode, ie20m (MIRIC 1966); 37. Bistra, Senečki Suvati, 1300 m (MARTINO 1934}; 38. Korab, Cos-Alija, 1500 m (MARTINO 1939); 39, Priiep, Markova kula (ZIN-L); Prilep, 700—800 m (B. PETROV in litt.); 40. Perister, near Magarevo, 1000 m (MARTINO 1939}; 41. Pres p a lako {FELTEN and STORCH 1965); Asamaiti, 860 m; KrurttifuoW, 920 m; 43. KajmakCaJan, 5—7 km below Reder, approx. 1000 m <B. PETROV in litt,}; 43. Kožut, Dve Uii, 1600 m {PETROV 1909}; 44. Gevgeli, near Vardaf, 60 m (E, PETROV in litt.); 45. Val and ovo, Anska reka, 65 m (PETROV 1909); 46. Sv, Nikole, 400m fB. PETROV in litt.); 47. SičevaCka klisura, 300 m {B. PETROV in litt.); 48. Berdap, Grobljanski potok near Dobra
(MIHIC 1981)
The major part of the D. n. area in Yugoslavia coincides with the principal mountain massifs, nanjely; the Alps in the northwest, the Dinaric Alps towards the southeast, above all their karst higlands. In the southeastern Yugoslavia the area corresponds to the Sar-Pindic and the Carpatho-Balcan highlands. In Macedonia it is widespread, populating also the Macedonian lowland (Fig. 3).
D. n, does not live on the coast and the islands of the Adriatic. It comes closest to the coast on the Velebit and along Boka Kotorska (Or j en, Lovcen). All of these localities are on heights exceeding 1000 m, i.e. in the montane zone. It might be worth mentioning that D. n. has not been found on the Dalmatian highlands inhabited by l^üomys quercinus (Mosor, Dinara).
D. n, most probably does not live to the north and the east of the moun-tamous region and is likely to be absent also from the entire Pannonian lowland.
Climatically the D. n. area reveals an outstanding variability. Characteristic of certain highlands to the northwest of Yugoslavia and of Montenegro are great quantities of precipitation representing extreme values in Yugoslavia or even in Europe (the Julian Alps 3141 mm of precipitation yearly, 1921 mm thereof in the period between May and October; Risnjak 3709^2260; Northern Velebit 3013/1482; Or j en 5317/1970}. Macedonian mountains are drier {Pel ister 1000/526), whereas Macedonian fields are positively dry (Sv. Nikole 453/273). Likewise diverse are tha average July temperatures which in the mountains vary between 10 and	{or even lower) and exceed 20 ^C in the lowland
of Macedonia.
3, Variiibiiity 3,1. Age Vanability
Different methods have been applied to determine age groups of D. n. The method based on the degree of the wear of tooth enamel permits precision up to the age of 1 year (HOMOLKA 1979, LOZAN 1961). Our material was divided into two groups:

Fig. 3. Hyptrthetical distiribiutian area of Dj-i/omys nitedula. Ln YugoiJavia SI. 3. HLpotetii&na raaäirienjost drevesnega polha v Jugoslaviji
Group: Juvenile animals born within the same year. These specimens retain juvenile pelage as late as September. The tail is less bushy. The weight does not exceed 19,5 g.
In thG material from Austria SPITZENBERGER (1983: 30) classilied specimens from 15 to 26 g into a juvenile category.
2nd Group: Adult animals which have hibernated at least once. According to LOZAN (1961: 1741) D. n. does not survive a fourth hibernation, consequently, it does mot live more than four years. Weight between 19 and 38 g (Tab. 1).
Mointh	n	lat Group min—max	n	2n£l Group min—max
June	0		3	2a —22,5
July	2	ID—n	9	21,5—29
August	4	13—19,5	5	22 —30,5
September	2	1&—18,5	8	10 —38
Total	8	10—19,5	25	19 —^B
Tab 1. Variations in weight of Dryamys nitedula from Vnigosla-via with respect to
age fffouips
Tab. 1. Var tiran je tefe pni Dryomj/s nitedula (iz Jugoslavije) gl&de na etarostne
isliuipdne
B. Rry&tuiek; Forest Dormomae Draotnys mtedwla (PALLAS, 1778)
HAI.ES	FEHAIKS
	n I ss	CT	n 2 5D	C7	t
*	22/ 52,a? t5,B52	23,07	13/ +	13,70	1,50
IIB	23/ 57,52 14,		13/ + 3,826	3.»	0,51
n	IV/ 03^2 +.S,aB7	7,Iff	B/ 82,oa +.3,665	4,47	
Iff	23/ 30,71 + 0,052	^,26	13/ 20,51 + 0,653	3,18	0,72
E	23/ 13,711 0,043	6,15	13/ y,OS t		1,W
PL	22/ 26,^ J. 0,741	£.75	13/ 36,46 + 1,024	3,67	1,53
CbL	24,66 +0,751	3.05	13/ 24,25 + 0,732	3,23	1,55
üab	23/ 12,03 + 11,347	2,es	13/ 11,79 + 0,547	4,64	1,?3
War	23/ 3,S5 + 0,170	4.42	13/ 3,79+0,171	4,51	0,55
DIL	23/ i 0,2^0	4,36	U/ 5,78 + 0,255	4/1	1,76
BaB	22/ 13,35 + o,^^3	3,"	X3/ 13,19 + 0,299	2,27	1,24
ZTB	IV 15,20 + 0,504	3,30	IV 15,0B + 0,579	3,8+	0,9s
KoB	23/ e,23+0,19a	3,1a	13/ 6,12+0,196	3,20	
IlDl)	23/ 5,07 + 0.236	4.65	IV 4,99+0,229	4,59	1,01
3C	23/ 4,17 +	f,05	IV ■ 4,051 0,151	3.69	1,34
m	W ?,19+0,294	3,10	13/ + D,44!)	4,fl9	1|45
BhIT	20/ 11,25 + 0,352	3.13	13/ il,l£ + 0,512	4,60	0,S5
	Z2/ 7,3+ i. 0,408		13/ 7*21+0,2»		1,^3
noR	2!/ 4,61 + 0,235		12/ +,S3tV?7	+,DS	0,30
HdL	iy i 0,52s	3,77	ly 13,85 + 0,463	3,34	0,39
R^		4,41	13/ 6,73 + 0,373	5,54	1,43
MdT	23/ 3,99 t 0,m	3,11	13/ 3,97+0,14+	3,63	0/9
QU	1?/ 62,22 > 1,976	3,ia	IV 62,41 + 1,873	3,00	0,25
HE	20/ 45,fll tl,SB5	3,Ss	13/ 45,69+1,861	4,06	0,12
an3	20/ 84,63 t 2,^3	3,46	13/ 04,36 + 3,27*	3,88	0,25
114	23/ 4B + 0,301	1.S5	13/ 48,59 + 1,041	2,1+	0,+Ö
	22/ 23,7B tl,477	4,96	13/ 29^7 + 1,061	3,59	0,44
a^	24/ 4?,57 t	4,35	13/ 48,58 +.2,U7	+ ,36	1,34
ai7	54,16 i 1,127	2,00	13/ S4,U 11.70«	3,15	0,10
aia	22/ 5^.49 11,759	3.17	M/ 57 + 1,459	2,55	1.13
Tab. 2, Monphcjnieitrrical characterisiäcs of Dryomys nitdeula fnam Yugoslavia Crom
the sex aspect
Tab. 2, MorfoTTi&triMe lastUciSti Dryomys nifedula (iz Jugoslavije) gicde na apci 3.3, Sexual Variability
Even though males attain somewhat higher values of metric indexes, the differences between the sexes are negligible and cannot be statistically corroborated (Tab. 2). Thus, we can affirm that in the case of D. n. secondary sexul dimorphism is not expressed.
3.3. Individual Vaitability
Of the morphometric indexes weight varies most (CV 22,07 with adult males). Body dimensions (CV between 4,13 and 1,07 for males and between 3,18 and 5,93 for females) are more variable than those of the skull (CV between 2,75 and 5,56 for males antj between 2,27 and 5,54 for females). The males display a greater variability than the females.
Considerable variability is due also to the colour, which is most evident in the D, n. from Montenegro. By way of example: of the three specimens from the Durmitor two have a greyish and one a yellowish-grey back.
3A Geographic Variability
For the purposes of studying geographic variability the material was grouped into four populations (Fig. 4);
L the Alps
2, the Northern Dinaric Alps

FS^. 4, Geographic jJioSititxriB of four (pojjulatioris of DryCrnys nitedula (sec the text for
further explanation)
SI. 4. Geografska lega stir ill ipapulaclj drevesnega .polha (ja razlago glej tekst)
B, Kry&tufek; Fwest Dormouse Dryom^s nUcduJo (PALLAS, 1718).,. 11
3,	the Southern Dinaric Alps
4.	Macedonia
The population of the Alps has a relatively (QI5) and an absolutely smaller bullae and a relatively longer rostrum (QI 4) than the population of the Northern Dinarlg Alps (Tab, 3, 4). From the population of the Southern Dinaric Alps it differs by a relatively lower skull (Ql 2, QI 3) and from the population of Macedonia by a lesser zygomatic breadth and a relatively (Qlä, QI 3) as well as an absolutely lower skull, The population of the Northern Dinaric Alps distinguishes itself from that of the Southern TJinaric Alps by a greater profile length, greater loC, relatively (QI 5) and absolutely longer bullae and a relatively lower skull (QI2), It differs from the Macedonian population by a smaller weight, a relatively (QI 2, QI 3) and an absolutely lower skull, relatively longer bullae (QI 5) and a relatively narrower skull (QI 1). The population of the Southern Dinaric Alps differs from that of Macedonia by a relatively (QI 1) and an absolutely narrower skull, a relatively (QI 2, QI 3) and absolutely lower skull and a longer mandibula.
The D. n. from the Alps differs from the D, n. from other parts of Yugoslavia by a relatively lower skull. The braincase height seems to reveal a dinal growth from the north towards the Eou1;h of the state. The population of the D. n. from Macedonia distinguishes itself from the D. n. from the rest of Yugoslavia by a relatively and an absolutely higher skull. The D. n, from the Northern Dinaric Alps has considerably larger bullae, exceeding S mm {the range for the rest of Yugoslavia is between 6,6 and 7,3 mmj.
In the scatter diagram (Pig, 5) in which the height of the skull per bullae is plotted against condylobasal length the population of the Alps and the Northern Dinaric Alps are well distinguishable from the Macedonian population. The population of the D. n. from the Southern Dinaric Alps reveals intermediate characteristics.
The coloration of the pelage displays a considerable variability. The most homogeneous are the two populations from the extreme areas of Yugoslavia. Thus, the back of thü D. n, from the Alps is grey, with indistinct brown tones or without them. In the population of the Northern Dinaric Alps the brown tones are more obvious. Considerable homogeneity of coloration is characteristic of the population of Macedonia clearly deviating from the D. ti. of the northwestern part of Yugoslavia by a markedly reddish-brown back. Sped mens with greyish tones are extremely rare in Macedonia.
The population of the Southern Dinaric Alps is characterized by an out-« standing variability of coloration. Animals with a grey or else a reddish-brown back can be found in one and the same locality (e.g. the Durmitor). On the other hand, however, the disposition of two colour types in the range of the Southern Dinaric Alps shows a disjunctive charactor. The D, n. from Maglit and Žljeb is perfectly grey, The adult female from Žljeb (BPS9/77) has a black-grey back, darker than of any other D. n. from the Alps that we examined. The specimens from Zelengora and Bjelasica are characterized by well pronounced brownish tones,

ä
	ALP3		KflElHfilJ/ DlKAfilC
	n I SD	CT	n I SD
n	4/ 27,25 +	14,57	5/ 24,00 + 2,915
HB	4/ + 0,957		6/ 96,5a t 4,037
TI	4/ e+,00 i i.m		4/ 64,38 t
Bf	</ 20,eQ +0,666		20,82 + 1,216
£	4/ 13,95 ± 0.173		6/ 14,40 + l.yS
PI,	4/ ±	2,50	i/ 27,42 i 0,736
ObL	4/ 24,35 + 0*433	1,97	6/ 24,96 + 0,674
HOL	+/ i	1,60	6/ 12,K i 0,337
KiT	4/ 3,83 i 0,096		6/ 3,00 1.0,063
DIL	4/ t 0,133	3,10	6/ 5,93 +0,314
BaB	4/ 13,30 ± 0,35£	2,es	6/ 13,50 i 0^79
ByB	4/ 14,05 ± 0,507	3,«	5/ 15,32 t 0,259
KD3	4/ £,ia + 0,116	1,89	6/ 6,27 1.0,175
R«B	4/ 4,95+(J,ISX	3,as	5,12 t 0,306
IC	4/ 4,Z&tO,141	3,3^	4,25 + 0,164
sa	4/ 9t50 + 0,£?4	3,16	6/ 9,03+0,234
BUI	V 10,73 + D.512	4,77	^ 11,12 t 0,E04
Sal	4/ 7,13 ±0,287	4,03	^ 7.77±5,4B9'
Boti	4/ 4,5&+0,i5Ü	5,OS	^ 4,93 + 0,137
HilL	4/ 13,90 + 0,^9*	2,16	6/ 13,97 1-0.071
mm	*/ +		C/ i 0^356
KIT	4/ 4,03 ± 0,050	1,24	C/ 4,00 t o,uo
Uli	4/ Sfl.jB £ I,£a3	2,10	5/ 61,011 1,566
012	4/ 44,05 t 1,457	3,31	6/ 44,51 1 0,957
ei3	4/ 20,62 +2,64B		6/ 02,3b i 2.0£9
ai4	4/ 49,0B i 0,251	0,51	i/ 46,10 + o',59e
015	V 2$t76 t 0,900	3,0fi	£/ 31,07 t lv365
äxS	4/ 47,i 0,653		£/ 4?,69 + 2,57^
IV}	V 54,sa t 0,^37	1.17	^ t C,549
dJS	4/ 57,w ± 1,459		^ 55,69 t
SOUlilERR DlflARIS ALPE
WkCfiUCim.
cv	nI SD	OT	al ED	CT
12,15	11/ 27,27 15,533	20,47	14/ 29,71 ± 3,632	12,90
4,10	iv ±3,145	3,2a	14/ 90,71 t 4,064	4,14
2,62	10/ 81,50 + 5,386	7,34	7/ 52,711.7,477	9,04
5,04	12/ 20^1 ± 0,614	3,02	W/ 20,83 + 0,762	5,66
7,'P	iV 13,8? i	6^27	W 13,58 + 0,710	5,29
2,6a	11/ 26,39 i 0,556	Zj22	13/ 2^,89 1.0,996	5,70
	U/ Z4,32 10,590	2,42	13/ 24,60 + 0,921	3,74
	12/ U,85 +0^7	3,25	14/ la,00 10,53a	4,48
	IV 3,90 + 0,217	5,56	14/ 3,ea t D,10?	4,9+
	5,77 ± 0,227	3,94	14/ 6,0110,259	4,31
2,BI	11/ 13,21 + 0,4lS	3,15	13/ 13.31 t 0,335	2,52
	S/ 15,00 + 0,500	3,33	y 15,61 +0,426	2,73
2,79	la/ i,2310,230	3,69	14/ 6,19 ±.0,18^	3,Off
5,90	lÄ' 5,Ö7+0,2£4	5,21	14/ 5,03t°il90	3,78
	IV 4,13 +; 0,103		13/ 4,10 +.0,204	4,98
	IV B,?iiO,35e	4,00	u/ 9,32 10,352	3,78
	11/ 11,16 i; 0,336	3,01	IV 11,54 i 0.354	2,20
6,3(1	11/ 7,1S + 0,21B	3,04	13/ 7,24 + 0,293	4,05
2,7a	IV 4.73±.0,iee	3,92	u/ 4,76+0,SI5	4,52
	IS/ 13,^3 + 0,542	3.90	M/ 14 ,U + 0,254	1,80
	IV 6,73 + 0,363		W 6,95 + 0,268	4,U
2,74	U/ 4,03 + 0,122		14/ 3,94 + 0,150	3,61
2,57	W 61.57+1,705	2,75	s/ 64,04 t 1,439	2,25
2,15	U/ 45,«? ± 0,307	1,99	11/ 4?,27 + 1,71?	3,64
2,51	IV 84 .47 1 1,?S0	i,36	11/ 07,35 11|735	
1,S4	iv 4S,a3 + 0,955	i. 95	13/ 48,6a + 1,155	2,37
4,39	iv 29,35 i 1,050	3,50	13/ 29,47 i 1,366	4,64
5.IS	11/ 49,39 s. 2,493	5,06	IV +9*25 t 2,142	
1,02	11/ 54,00 + 1,7S7	3,27	13/ 54,14 + 1,481	2,74
3,Ee	11/ 56,26 + ij7oi	3.(ß	13/ 97,52 + 1,506	2,6Z
B. K^y^tufek: For-est Doranonise Dryomys nitedula (PALLAS, 1778) ... 13
Alpa	il|i4	A tili	ir-HlasTl«	R-Dlnarla S^JUnul«
B-Diaarta S-flimrdO	IfaBodimlS EJJijiarjlq	Ma^samin HKSfrdfflllt
T 1,«	0,01	1,J3	3,02^	1,30
HB 1,31	0,03	0,43	1,12	lfS9
n ö,2g	o,&i	0,33	0,3a	Q^z	0,37
Hf 0,03	n,07	0,15	0,03
t 0,77	0,17	1,0a	1,08	1,96	0,?r3
PL l,e7	0,73	0,44	1,^0	1,45
CÜ 1,60	0,10	0,51	2,10	0,S5	0,8B
Rai.	0,3^	0,2^'	0,13	0,£S
Ktt	0,49	0,04	1,03	0,39
Iii 0,19	1,06	0,7^	1,29	1,57	2j50
BeB	0,84	0,39	1(41
1,01	1,00	2,02^	1,32	1,39	£.77
HOB 1,67	1,04	0,9*	0,J9	0,82	0,40
RoB D,^	0,84	0,73	0,60	0,«
10 0,50	1,20	0,91	0,77	o,za
™ 1,60	1,96	0,09^	0,7a	1,76	2,74^
IM 1,72	0,36	4,20	0,25	3,43	5,00
EaL i,34	0,41	6,16	2,9^	0,S2
See 0,35	1,56	1,71	1(6+	1,S4	b,+3
m. 0,1?	0,93	1,35	1,15	0,72
BKH 1,55	0,55	1,11	0,11	1,70
MT 0,42	0,ÜÖ	1,14	0,4 2	0,94	1,65
an	0,03	1.49	1.05^
™ 0,76	2,97^	3,3+^	2,?0	3,^0	2,3^^
ai3 l,6fi	3,05	1,05	S,27^
atü 3,05	0,35	0,^7	it«	1,15^
ins 2,32	0,1s	0,30	2,38	0,33
aifi 1,52	1,34	1,44	o,5B	0,39	0,15
air 1,55	0,62	ü,04	o,ie	0,?o
aia 1,10	Ü,a7	0,49	2,05	hP-
Tab, 4. Interpopuilation t-test values of four papuiatkins erf Driiotnys ttitediilo fram
Yrugoslavia. P =0,05 (+) and P 0,01 (++)
Tab. 4, Interpapiilacljske vrednosti l-testa pri &ÜTih	populaaijah vpite Dryomus
rtitediiJa '!^z Jugaslaviije). P = 0,05 (+) üi	P = 0,01 ( + +)
Subspecies of D. n. in Yugoslavia
The subapecific status of D. ti. in Yugoslavia has been discussed by the followmg authors: MILLER {1912: 569J, WETTSTEIN (1928), MARTINO (1934: 8B, 1939), PETROV (1939: 78), PASPALEFF, MABTINO and PECHEFF (1933), FELTEN and STORCH (1965: 351), DULtČ and VIDINiC (1967: 151) and SCHEDL (196G: 392).
For the territory of Yugoslavia two subspecies were described, namely:
BbH
U
11 -
111 -
13

15
!B
Ct>L
Fig. 5. Discriniiinatkwi of subspecies of Dryomj/s nitedula Ln Yugoslavia by means of a scatter diagram wliere ithe bradincaise height iier tmUae (BbH) is .ploMad agaimat the Kmdylobasal. length (CbL). Solid circles reiJr^eiit D. n. intermedius; half solid
circleE D. n. iTiterTnedius > D,Tavtjojia ijitergraides; open circles D, n. Tavijojla;
crosses, avK-age values SI. 5. Raallfeovanje ipodvrst dreivesnega ^polha v Jugoeloiviiji s ipomočjo Jioreiacijsirega diagrama. It j er nanažaimo wüino lobanje 2 buJamd (BbH) proti kandilobazairi dolžini lobanje (ObL). &inii Isrogi predistavlja^o D, rt. irtteTJnedtUS; f>ol5rni kPOgl OBnačujeJO
prehodno obliiko D- jl intertnediiis ^ D. n. ravijojla; beli Jcrogi D. n. Tooijojla; križi
— srednje vrednosti
D, n. ravijojla PASPALEV, MAETINO and PECHEV, 1952 (loc, typ.: Macedonia, Bistra Mts., Sensčki Suvati).
D. 71. diamesiis LEHMANN, 1959 (loc, typ.: Montenegro, Lovčen Mts,, Ivanova Korita).
Most of the authors overlooked the denomination "ravijojla", and only COHBET (1978; 146) and STORCH (197B: 229) drew attention to this name once again.
In the last decades several subspecific classifications of D. n. have been proposed with respect to the Balkans. PASPALEV et al. (1952: 202) identified four subspecies which they keyed as follows:
s, KrystiUfek: Forest Doirmoiuse Dryomys niieduiji (PALLAS, 1778).., tb
1.	Skull massive, zygomatic breadth exceeding 15^5 mm
(on an average)	D. ti, robusttts
—	Skull delicate, zygomatic breadth not eitceeding 15,5 mm . . ....... , , ......2
2.	Back dari: grey..............D. n, inlerinedius
—	Back yellowish...............3
3.	nostrum shorter than 8 mm..........D. ti. wingei
—	Rostrum longer than Ö mm..........O. ravijojla
Ascribed to D, n. ravijojlti were the specimens from Macedonia, Kosovo and Bulgaria.
DULIC and MIRiC (1967; 28) cited 5 subspecies for Yugoslavia:
D.n. nitedula (PALLAS, 1779) for Croatia, Serbia, Bosnia and Herzegovina
D.n. wingei (NEHRING, 1903) for Macedonia
D. n. intermedius (NEHBING, 1902) for Slovenia, Bosnia and Herzegovina, northern Montenegro, and Croatia with a question mark
D.n. robuseu5 (MILLER, 1910) for Macedonia and Serbia with a question mark
D. n. diamesns (LEHMANN, 1959) for Montenegro For all of these taxa (MIRiC, 1970) contributed short diagnoses. ONDRIAS (1966: 22) identified 5 subspecies for the Balkans and mapped their distribution. He cites D.n. intermedius for western Yugoslavia, D. n, nitedula for eastern Voivodina and Rumania, D. n. Tobustus for Bulgaria, D, n. diamesus for Montenegro, and D, n. wingei for Macedonia and Greece. His classification reposes on pelage coloration.
STORCH (1978: 232) proposed the following classification for the Balkans: D, n. iritermedius (ober sei tes dunkel grau mit höhstens schwachem olivgrünem odor braunem Anflug, schmale Schädelkapsel, relativ kleiner Schädel): Alpen, Abgrenzung nach E (Ungarn) und SE (Slovenien-Bosnien) unklar.
D. 71, wing^i (bei Adulten oberseits lebhaft rostrote Tönung, relativ geringe Schädelmasse, sehmale und flache Hirnkapsel): Griechenland und S-Ju-goslawien (Makedonien), wahrscheinlich auch S-Albanien und SW-Bulgarien.
—	Synonim; Tavijojla,
D, 71. nitedula (Obersoitenf ärbung graubraun, gelbbraun oder rotbraun mit grauem bi^ rostrotem Uberflug, Schädel relativ gross, HLrnkapsel robuster als bei den vorhergehenden): Übriges europäisches Art-Areal, Terra typica der Nominatrasse ist die Gegend von Kasan, UdSSR. — Synonyme: robustus, car-pathicus, diamesws.
We can see that most of the authors lay great stress upon the colour of the pelage whereas we consider it an auxiliary character. In our opinion a material distinction between the population of the Alps and possibly that of the Northern Dinaric Alps (in fact the latter has larger bullae) on one hand and the population of Macedonia on the other as two different subspecies can be made by taking into account the relative height of the skull. The former would be identical with the ssp. i»itermediHS (Tab. 6). In our belief the D. n.
?
I H ■ a = s a =
t- LA lA r^	INJ IM ITV O
S 3 S a"» a	3 a 3 s a Ä*
«D (4	«
0\ C9 CQ	"C^ O QV Cr -Crv, ^
biS
JH u to Si» VD	(D ff^ ^ ^
^ Jt	Ä ^ ffS o^i
^ r-t >-f rH *H
■»1 #1 »I •	® I »1 • •• »I •• •
C^ O", O	cd CO t^cdfiCiGA
f? C> o	ODO0
4] fc^ «	4. > *!	4'
□I ^AC^HC^LAIA	fli O fr^ lA
p C ni 'HrH O	OM^ fs #H O r4
J
C? « g ü
• " ä 11 a ^
S-Jd r iS H 5 ^
a
ir.
" s jj
I
s.. _
■ ja Tt	H »t
^ S S M Jt
M ^ S ^ S A£ is^ssS^ss
»	v^ ^ irt
^ EiS ^ E^l fe g	StflS^J
f
u
E
01
« . ^ g
s
u
J
9

Tab. 5. Data tm Dryomvs ntteduia fiTtan Yugpdslavia as gathered froin the coJlectton
of V. MARTINO (ZtJoio^ical toatitiite, USSR Acadmey of Sci&ncas, LeÄigrad) Tab. 5. Podaitikii o vrati Dryom^^s nitectula i\z Jus osla vlje), fci so v zbdrid V. MARTINO {ZooJoäki mi&tiitut. Akademija znajwsti ZSSR, Leningrad)
B. Kry^'bufek; Forest DarmOmSe DryOmys niteduEo (PALLAS, 17Va) ...
17
Geographic Region	Subspedes	N	min— S —^jnox	Authors
Austuria	Lntermedius	IC	23,a—24.33—25,5	Spitzen berger 1903
Yugoslavia {Alps, North, Dinaric Alps)	intermediiE	10	23,0—24,73—25,8	the present study
Yugoslavia (Magedonia)	raviijojla		32,9—24,60—26,4	the present study
Greece	Wilngei	13	23,2—Ä3,5 —34,0	Ondrias I96fi
Bulgaria	wingei	110	22,0—23,95—25,5	Peshev, Mi lev 1979
Europeain Soviet Union	nitedula	114	23,0—25,12—27,0	Rossolimo 1971
Asia Minor	phrygius	14	23,0—23,37—24,8	Spitzenberger 1973
Tab, e. CoTidylobasal length nn UÜferent subapeDies of Di-yomys tiifsdula Tab. 6. K:ciiiidiloba2ailna dolžina lobanje ipni razlifinih ipodvrstah Dryomys nitedula
from Macedonia should not be identified as D. n. wingei from Greece since the former is greater and has a higher skull. It is denominated rnvijojla, Macedonia beitig its type locality.
The population of the Southern Dinaric Alps reveals rnixed characteristics both with regard to coloration as well ss in relation to BbH/CbL, therefore, it is designated as an intermediate (mixed) population fD. n. intermedins > D, fi, ravijojla). If recognized as an independent subspecies, this population should he denominated -»diamesiis«. For the present, however, we consider it more adequate to treat it as a population of transitional properties.
No data are available on the morphologic characteristics of the D.n. from the Carpatho-Balcanic mountain region. Consequently, nothing can be said of its taxonomic status.
Summing up, the following picture is obtained:
D. n. intermedius fNEHRlNG, 1902)
Type locality; Near lienz, Tyrol, Austria
Distribution: Alps and Northern Dinaric Alps (Snežnik, Ri&njak, Velebit, western Bosnia},
Diagnosis: Skull lower (BbH 10,0—11,4mm). Back mostly grey, brownish tones can also be expressed.
D. n. Tavijojla PASPALEV, MAKTINO and PECHEV, 1952
Type locality: Senečki Suvati, Bistra Mts., Macedonia, Yugoslavia.
Distribution: Macedonia
Diagnosis; Skull higher (BbH 11,2—11,9). Back mostly reddish-brown, rarely with visible greyish tones.
D. ri, intermedius ^ D. n, rnuijojla
Distribution: Bosnia, Herzegovina, Montenegro and Kosovo.
Diagnosis; Mixed population. BbH ID,4—11,6mm. From the coloration of the back the characteristics of both subspecies are evident.
.J \


t
/"v.
1


O
Fig. 6. DiÄtribUiättnal reCOTids Ol Druomyi nitedlila subsl>eQiefi in Yugoslavia. Symbols
as in fig. 5
SL e. Kae^irjenoet podvrst Dryamys n.itedu!a v Jugoslarviji. SimtwU sü taMni, kt>t
na si. 5
5. Vertical Distribution
All known localities of D, n. in Yugoslavia Ee within 60 and 2000 m of altitude. The lowest Locality is that in the lowland of Macedonia (Valandovo and Gevgeli; PETROV 1S69: 1), the highest in Montenegro {Durmitor; PETROV 1939: 78). The vertical range of D.n. in Slovenia is between 490 and 1780 ni, in Croatia between 550 and 1650 m, in Bothis and Herzegovina between 330 and IBQD m, in Montenegro between 1050 and 2000 m, in Serbia between approx. 60 and 1750m, and in Macedonia between 60 and 1900 m.
67,4®/o of all localities of D.n. lie above 1000m. One half (53,1 %) belongs to the highland zone from 1201 to IßOO m. D, rt, is rarest in the lowland zone (up to 600 m.). In this respect the lowland of Macedonia is an exception. Most localities (65,3 ®/o) are to be found in the montane zone {600 to 1600 m of
B. KryÄtufek: Foreat Dormause Dtyomys jiifediitc (PALLAS, 177S)... 10
Fig. 7 0, 3, gkulls öf Dryomys nitcdulcl fromi Yugoslavia, a — D. n. intermedius (Slovenia, Julian Alps); b — D.n. ravijojla (Macedonia, Pjespa lake}. Fig, 8. — Nofrma dansalis; ög. 9, — Kturna ventralLs; tig, lO. — Norma lateralis. (Foto M.
Stangelj)
SI. 7, (J, 9. — Ijobanji Dryomys nttcdula iz J.ugoslavije. a — D. Ji. intermedins (Slovenija, Juli jske Alpe); b — D. U. Tavijojla (MaJiedflniiija, E^espaiu^ jezero). Si. S. — Norma dtmsali^; si. — Noima venitralis; b1. 10- — Norma laterailiis. (Foto M.
Stangelj)
B. Kry&tufek; Foiresi Dormootee Uryomy^ nifeduia (PALLAS, 177B) ... 21
altitude). This means that D. n. is a species of the upper montane and the lower Alpine zone.
The narrowest vertical distribution of D, 71. Ls observed in the Dinarle Alps (Velebit, Bosnia and Herzegovina, Montenegro) where it is limited to the montane and the sub-Alpine zone (verticals range from 900 to 2000 rn), whereas the broadest verticale range is known in Macedonia (60 to 1000m of altitude).
4,08
16,33
6. IIabital Requirements
In Yugoslavia D. n. inhabits the forests of the order Fagetalia, Vcn:cinia-Piceetalia and Quercetalia., as well as their degradational stadia. In beech forests {AnemancFagetum, Abieti-Fagetum, Fagetum mantanum, Fagetum subclpin-um,
ALT (m)
20Ü0
1800 -
1600 -
1400 -
1200 -
1000 -
800 -
600 -
400 -
200 ' 0
20,41
16,33
10.20 10,20
8,17
4,08 4,08 6,12

—I— 10
15
I
20

Fig. 10. Vertcal ddstrLbution of Drvom^a nitednlfl dm Yugoslatvia. Itiriicasted ig the
frequency of presence wfijthlii a two-hundred meter highlamd zone 31. 10. VeJ^LJcalna raizSirJsniCBt Drt/ornjs nitedula v Jiigoslaviji. Oznaiena je frelpvenca prisotnosti znatt-aj dvestomatrake^a viSinsäiega ipaeu
B, Kry^tufdf: Forest Donraoiuse Dryomas nitedula (PALLAS, 17Ta)...
33
Seslcno-Fagetumj it has been found above all in the Dinaric region. It seems to pref&r tJie rocky ground which is quite frequent in the beech forests of said region. It has also been found in rocky places once overgrown by beech forests where as a result of degradation there remain just single beech trees. It is less keen on pine and fir woods {LuEuta siluaticacPtceetum, PtceettiTn excelsas montanum, Piceetum excelsae croaticum, Pinetujn leucodermis, Pinetum peucis), preferring the dwarf pine (Rhndotamno-Rhoiiodendrctum, Pinetum mug i dinaricum) to all other environments. In Macedonia and Serbia it has been found also in oak forests fQuerco-Carpinetttm orientoÜs, Querceium trojftTme, Quer-cetum c arris moesiocw-m, QusrcHwm /rame^to-cerrisj and the more frequently so in their degradation stadia (rocky places, live-hedges).
Most of the habitats of D , Tl. are termo p hi lic and 'xerophilous. This is also true of some highlands with a high quantity of precipitation, most of them being carstic, which means that the meteoric water quickly flows off to the underground. Moist and cool habitats are an exception, at least as far as Yugoslavia is conccrncd (e. g. Pohorje).
Fig. 11.—J9, HaW'tats of Dryomu« nitetiiila lin Yuig<JsJavia; fig. IJ. Jul■i^ln Alj>s, Planica, 1000 m (Foto B. KryStuEek); Pig, 12. Poh-ca-je, Velika Kapa, 1530 m (Foto D, Sare); iig. 13. Snežnili. SvinS&aikn, 1242 m {Fato J. Go-egari); fig. H. Mwnteine©ro, Bjelasica, JeJavica, 14S(}m (F<)to B. Kri-Stufek); fig. 15. Kom Vasojevlč, 1000 m (Foto B. Kryätufeik); fi^. 16. Ourmitör, Crepuljrta poljana, 1700 m fFoto N. Mršii}; «g. 17. Žljeib, Kula, 1750 m {Foto 3. Kryitufek); fig. 19, Orjen. Crkvice, 1050 m (Foto B. ICrrštufek); fig. 10. PpMpa laike, Kiirbiinovo, 920 m (Foto B. Kryitufek).
SI, 11.—19, Haibiitati drevesnega polha v JugoslanMji

B, Kryätufek: Forest DonriMJuse Dryomys nitedula [PALLAS, 1778),.. 33
B. Ktyätufek: Forest Dormouse I>Tyc>myä nitcdula (PALLAS, 17TB) ... 27
fig, 30. Dryomi/s nitedul® iwterfraedius (carcass). Adult female caught on Pohorje,
VeJika Kopa (1520 m), M 4tli July 1983. (Fatol. Sivec) SI, KO. Dryomys mtedttla intermedius (mritva živaJ). Odrasla samica, ujeta 4. juJija Z983 na VeiiJci Kopi, Pohorje, (FotO I. Sivec)
7. Dominance
Among more than 10.000 specimens of small mammals caught all over Yugoslavia there are only 51 specimens of D. n. In Yugoslavia as a whole D, n. is n rare species. In single habitats where caught its dominance varies between 1,01 and 27,7 "A (Tab. 7), which means that it is a subrecedent and recGdent, and only exceptionally a subdominant species.
Due to a high dominance of D.n. certain habitats destrve special attention.
— Planica (10ÜO m)
In Planica small mammals w<3re being caught during 9 nights. The density of their population has always been low, so all we caught were 39 specimens, arnong them G D. n. In the same habitat we found also Sorea; aratieus, alpinus, Clethrionomys glareolus, Chionotnys nivalis, Pitymys liechtensteini and Apodemus
" Maglič (1700—1300 m)
Into 50 traps laid far one night at Pinetum mugi and the neighbouring rocky areas, 11 small mammals (Sorer omneu^, Clethrionomys plareolus, Dina-
B. K■^yätuf£^^^ F«rest Dmmouse Drvamyi nitedula (PALLAS, 1773)...
29
Locality	Altitude	Haihiitat	N	Nd	D m
Julijske Alpe, Planica	1000	Hhodothamno-Rhododendretnim PredtMmnaiiUy Toclty area		6	15.SR
PohiiTic, Vfelika Kopa	isao	Liizulo silvaiticao-Piceetumi With numerows clearings	43	1	2,33
Snežnik, Svlnäeaki	1242	Abiöti-Fagetüin dinar i cum	80	3	3AQ
Hi&njaik	1450—1500	Pinetum mug!	5B	2	3,45
Veleblt, Zaviizan	1500—1650	Pagetum subalpinMm	33	1	3,03
Saitor Mts,	1550	Piinetum niugi predcflnlnaTitly rooky area	33	1	3,03
Kllpreäka vrata	B80	Abieti-Faüertuin	SO	1	1.01
Zelengora	1500	flooky area With Fagus sylvatica	27	l	3,70
Maglifi	1600—1800	Pinetum raugi	II	3	27.27
Durinitor	1450—1700	Piceetuin excelsae croaticum with immeroiis clearings	53	4	7,55
Bjela&ica	1350—1500	Fagffbum montainum	270	3	l.ll
Kamovi	1800	Fagstaim Ettbalpinum and rodky area	SO	1	3.00
2lje&	1750—ISOt)	Riceetam exoelsae Bubaipi^m	25	2	B.00
Prespan, Aaamati		Quercetnun frainetto — cenfis	62	4	
Prilep	700	Rocky area	30	0	20.00
Valendcivo	65	Sdäx, Popultls	2B	3	7,14
Gewgeli	75	Rocky area ovorgrown with shtiubbery	12	2	16,67
Tab 7. Dominamce of Dryomi^st nitedula in certain Yugwlav locaJitias N — total number of small mammals; Nd — Tnimber of forest -dormice; D — domJ-
nance of forest dormice Tab. 7, Doretimimtnoet -drpvesne^a pdha tia nekaterih nahajališčih v Jugoslajviji N — ^upno Iteviliäfl malte seselcev; Kd — Število drevesnih polhov; D — ckmilnantno&t
drevesnega polha
Tomys bogdanovi, Chionomys nivalis, Apodemus sylvdticus and A, flavicoUis) were caught, among then:i 3 D, n.
— Prilep (7ffO—SOOmj
Into 100 traps set in a rocky area, 31 small mammals (Apodemua sylvatictis and A. mysiacinus) were caught, e of them being D. n.
Living syntopically with D. ti. are no less than 20 species of small mammals, namely: So rex uraneiis, S. alpinus, S. miiiutus, Neomys fodiem, N. anomalus, Croddura leucodon, C. suaveolens, Cl^thrwno'tnys glareolus, Dinaromys bog-daTvouij AficTof-us agrestis, Cfiionomj/a niualis, Pitymys subteTToneus, P. liecft-
Unsteini, Apodcmuj sylvaticus, A. jlavicollis, A. viystacinus, Mus hortulanus, Mus domssticus, Glis glis, and Muscardint« avellanarius.
In no locality has D. n. been found together with Eliomys quercinns.
8. Period of Activity
All specimens of D. n. in Yugoslavia were caught in the period between 7th April and Octobcr. The specimens caught in April aU originate frotn Macedonia. In the period from October-November till the end of March (in mountainous regions probably till the end of May) D. n. hibernates.
Month	N. D. n.	N. r. f.	N. juv.
April	11	—	—
May		—	—
■Tune	n	2	—
July	10	6	3
August	21	—	C
September	20	—	3
October	1	—	—
Total	77	8	11
Tab. 8. Demographic charaotarisbiCE of Dryomys niteduta. from Yiogosiavia N. D. n. -- total number of ftureat dormice; N.i f, — number -of fernales showing signs of reproduction (ipragnancy, 'laotaition); N. juv. — number of juvenile foi-est
dormice
Tab. 8, Demografiske Ma&ÜnoEti drevesnega polha tz Jugoslavije N. D. JI. — sikiipno število drsvesnih polhov; N. r. f. — feteviJo seinic, ki so Itaiale Enalie brejosti, laktacije; N. juv. — žteivilo juvenilnih drevesnih rpolhofv
8. L Reproduction
Females showing signs of reproduction (lactation, pregnancy) were found in June and July. Only three of them were pregnant:
Locality	Date	Embryos Number	Dimensions
Dur/nitor	gill June	5	3 X 2,5 mm
Pohorje	4tli July	2	4 X 3,5 mm
Snežnik	Iflth July	i	5 mm
The first young animals of the same year were caught in July (on July on Snežnik and July on Orjeri),
The testes of sexually mature males are from 7 to IS,5 mm long and 3 to 7,7 mm broad. In young ones they amount to 5 X 3 mm.
B. Kryätitfök: Forest Darmouse Bryomys niteävXa (PALLAS, 1770)... 31
2. Conclusjions gad Discussion
Our results have considerably changed the present knowledge on D.n. in Yugoslavia, STORCH (1&78: 223) mapped this species for the entire territory of Yugoslavia, with the exception of Istria and western Slpvcnia. Now we know that D. Ji, does not live in the Pannpnian lowland to the north of the Sgva and the Danube, being absent also from the mapr part of the sub-Pannonian and hiUy region ta the south of the Sava. It is not an inhabitant üf Istria, a large part of the Adriatic coast and the islands.
MATVEJEV (1961; 97J considers D. n. a species characteristic of the mediterranean mountain rocky woods- This group of elements distinguishes itself by ils tertiary age. However, KEATOCHVIL, (1967} believes that D-n- came to Europe from the East through A$ia Minor as late as in the Holocene. According to SPITZENBERGER (1933: 19) it arrived "into the Eastern Alps together with the spruce from a glacial refuge of mixed oak forest SE of the Alps in the Boreal". STORCH (1978; 232) considers D, n, "ein sehr junger holozäner Immigrant aus SE bzw. E.",
The oldest fossil findings of D. h. in Europe date from the lower and the upper Pleistocene in Hungary (JANOSSY 1962, 1964, according to STORCH 197B}. No data are available on fossils of D. n. from Yugoslavia and the Balcan Peninsula as a whole. With respect to the findings of JANOSSY it seems possible that D. n. inhabited the Balcan Peninsula as a glacial refuge as early as in the Pleistocene.
For the territory of Yugoslavia authors have stated a considerable number of subspecies (up to 5). We, however, distinguish only two but admit the possibihty of identifying the morphologically mixed populations in the transitional area (Herzegovina, Montenegro) as a third subspecies n. diamesusj. Such an arrangement is not likely to be final. With the intermedius as defined the specimens of the Northern Dinaric Alps deviate from the others by their large bullae. When more material is available, the status of the population from this region (Risnjak in particular) will become much clearer. Quite heterogeneous is the population of the Southern Dinaric Alps. Here a disjunctive distribution is observed of two basic colour types, whereat their appsarance cannot be explained by the ecologic conditions of the environment. If paleon-tologic investigations prove that to D.n. the Balcan Peninsula olfered a glacial refuge, it might become more realistic to recognize a greater number of subspecies than recognized by ourselves. Irrespective of the above, the subspecies so far cited for the territory of Yugoslavia are contrary to facts.
Ackiiawled jfcmen ts
The present study could not have been nrealUieid wRbaut the genenoius help of Dr. BORIS M. PETROV (Belgrade) who (put at my disposal his unpubUshed data and material, as well as data on the oollecWon of V. MARTINO in the ZIN-L. His aid consisted also in critiicaJ advices and suggestiOTis. 1 hereby warmly thamk him (or his help,
1 am ithauicful also .to Dr. TSOLO HR, PESEV (Sofia) who kindly .provided me with the pertinent literaituTo and oammuniicffited his views on the subspeoific status q( the forest dormouse in Btilgania.
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Povzetek
Drevesni polh Dryomys nitedula (odslej D, n.) je eden slabäe poznanih sesalcev jugoslovanske favne. Zaradi svojega zanimivega videza in redkosti je vedno zanimal teriologe in zato niso redke kratke notice o njegovih nahajališčih ali taksonomskih vprašanjih. Vedno pa upoštevajo ftiajhno Število primerkov, pogosto le enega samega.
Drevesni polh v Jugoslaviji v glavnem naseljuje glavne gorske masive {Alpe, Dinaride, garsko-Pindske in Karpatsko-balkanske gorske pokrajine). V Makedoniji, kjer je splošno razširjen, naseljuje tudi nizko Makedonijo. Na jadranski obali in otokih D. n. ne živi. Omembne vredno je, da D.n. ni bil najden na dalmatinskih planinah, kjer živi Elicmys queTciiiHs (Mosor, Din ar a).
Severno in vzhodno od gorskega sveta, proti subpanonskim pokrajinam, D. TI. verjetno ne Sivi več. Prav tako ga verjetno ni nikjer v panonskem niža vju.
Vsa znana nahajališča D, ji, v Jugoslaviji lože med 60 in 2000m nadmorske višine. Najnižja nadmorska višina nahajaliääa je v Makedonskem nižavju, najvišja pa v Črni Gori (Durmitor), Večina habitatov je termofilnih in kserofilnih. To velja tudi za nekatere planine, kjer je sicer veliko padavin {npr. Orjen), a ker so v glavnem kraSke, padavine hitro odtečejo v podzemlje. Vlažni in hladni habi ta ti so, vsaj v Jugoslaviji izjemni. Gledano kot celota, je D. n. redka vrsta. V posameznih habitatih, kjer je bil ujet, variira njegova dominantnost med 1,01 in 27,7 Večinoma je torej subrecedentna in recedentna ter le izjemoma subdominantna vrsta.
Vsi primerki D, n. v Jugoslaviji so bili ujeti v času od 7. aprila do 11. oktobra. Samice, ki so kazale znake razmnoževanja, smo dobili v juniju in juliju, prve mlade živali, povržene v istem letu, pa so bile ujete v juliju.
Na ozemlju Jugoslavije razlikujemo dve geografski rasi (podvrsti) drevesnega polha. Ena je omejena na severozahodni del države, druga pa na Makedonijo. Vmesne populacije kažejo mešane {prehodne} znake, zato jih oznafiujemo kot mešani (vmesni) takson. Ce bi tudi njemu priznali status podvrste, bi ga morali poimenovati D. n. diamesus LEHMANN, 1959.
Subspecifična klasifikacija D.n. iz Jugoslavije je takšnale:
D. n. intermedius (NEHRING, 1902)
Nahajališče tipa: Lienz, Tirolska, Avstrija.
Razširjenost: Alpe in severni Dinaridi (Snežnik, Risnjak, Vele bi t, zahodna Bosna).
Diagnoza; lobanja nizka; hrbet večinoma siv.
D, n. ravijajla (PASPALEV, MARTINO and PECHEV, 1952
Nahajališče tipa: Senečki Suvati, Bistra, Makedonija
Razširjenost: Makedonija
Diagnoza: lobanja višja; hrbet večinoma rdečkastorjav.
D. n. intcrmedius J favijojla
Razširjenost: Bosna, Hercegovina, Črna Gora in Kosovo.
Diagnoza: mešana populacija. V obarvanosti hrbta kaže lastnosti obeh podvrst.
B. Kt-yžtutek: Forest Dormou« Dryomys niiedula (PALLAS, 1778).,. 35
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