ISSN 0351 - 0077 PRIRODOSLOVNI MUZEJ SLOVENIJE MUSEUM HISTORIAE NATURALIS SLOVENIAE SCOPOUA 30 Botanka GeoJogica & Palaeontologica Museologica 19 Zoologica Boris KRYSTUFEK: European Sousliks {Spermophilus citellus; Rodentia, Mammalia) of Macedonia Tekunica {Spermophiius citellm; Rodentia, Mammalia) v Makedoniji SCOPOLIA No 30 pp. 1-39 Ljubljana Nov. 1993 SCOPOLIA Glasilo PrirodoslovnegE muzeja Slovenije. Izdaja Prirodoslovni mu^ej Slovenije, sofinancirali &o: Republiäki sekretariat ia raaiskovairio dejavnost in tehnologijo. Republiški sekretariat za kulturo in Znanstvenoraziskovalni cencer S AZU. Uredniški odbor: Jože BOLE. Ernest FANINGER. Janez GREGORI (urednik), Boris KRVŠTUFEK, Ignac SIVEC, Kazimir TARMAN in Tone WRABER. Lektorja: Cvetana TAVZES (za slovenščino) in Helena SMOLEJ (za angleščino). Naslov uredništva in uprave: Prirodoslovni muzej Slovenije, 61000 Ljubljana, PreSemova 20- Izideta najmanj dve številki letno, naklada 600 izvodov. TekoCi račun pri LB £t. 50100-603-4Ü115. Tisk tiskarna Tone Tomšič, Ljubljana. SCOPOLIA Journal of the Museum of Natural History of Slovenia, Ljubljana, Edited by the Museum of Natural History of Slovenia, subsidized by Republican Secretariate for Rcscarch and Technology, Republican Secretariate for Culture and Centre of Scientific Research of the S AS A. Editorial Staff: Jože BOLE, Ernest FANINGER, Janez GREGORI (Editor), Boris KRVŠTUFEK, Ignac SIVEC, Kazimir TAJiMAN and Tone WRABER. Readers; Cvetana TAVZES (for Slovene) and Heiana SMOLEJ (for English). Address of tlie Editorial Office and Administration: Prirodoslovni muzej Slovenije, YU 61000 Ljubljana, PreSernova 20, The Journal appears at least twice a year, 600 copies per issue. Current account at LB No 50100-603-40115. Printed by tiskarna Tone ToraŠič, Ljubljana. SCOPOLIA Wo 30, pp. 1-3», Nov. 1W3 European Snuslik« (Spermophilus eitel I us; Rodenlia, Mammalia) of Macedonia Boris KRVŠTUFEK Slovene Museum of Natural History SLO. 61001 LjubJjana, P.O.B. 290, Preäemova 20 Received: 27. 10. 1993 UDC(UDK) 599-32(497.17)(045) =20 SpermophHus dieilus ABSTRACT - Twy subspt^cic!^ of ground squirrch are known from Macedonia, living in strikingly different T^abitatü. SpErmophitus cfieZ/itT gmdojivici is linked to tlic hiwlandü along the River Vardar, south of Dcmir fCapija, while S. c. karamani populates mountain pasture!^ in the lakupica and Karadjira Muuntains. Spot distribution maps Viith identifying numbcin, and a list of diagnostic charactci^ ave prcwntcd for each siihspceies. SpermnphUus ciiellus gniiJojivici and S. c. karamani differ ir terms of size, skull ishape, their nasals and foramina incisiva, and in tail coloration. Key words: Spermo{fhdus citellia, Macedonia, taxonomy, distrihution POVZETEiK - TEKU NIC A (SFBRMOPHILÜS CITIlLLUS,- RODENT! A. MAMMALIA) V MAKEDONIJI - Na ozemlju Maki:donijc živita dve podvrsti tek Linie na povsem različnih habitatih. ^Spermophilus citeltus gradoj^vici živi \ nižavju vziJuli Vardaija, južno od De mir Kapi je, 5. c. karatnani pia na planinskih p:iSnikih Jakupicc in Karadžiu:. Za obe pndvrsti sta bili izdelani ar^alni karti in seznam morfiiii^itih znakoi», ki «mogCiüa njvno prepouiavsnje. Spcnnophilijs citellus grafio]fvici ge razlikuj« od S- c. kurumani po velikosti, obliki lobanje, obliki nosnih kosti in nebne Špranje ter barvi repa. Kljkifne besede: SpermopMus cM/us, Makedonija, sistematika, razžitjenost 1, Introduction Since 1921, when the European souslik was reported from Macedonia for the first time (Doflejn 1921), it has continued to attract the attentian of researchers from different points of view! taxonomy, jwogeography, ecology, and plant protection. Consequently, a considerable amount of information has accumulated on this species in comparison with the other rodents living in Macedonia. The aim of the present article is to review the distributional and taxonomic status of the European sou-slik or European ground squirrel in Macedonia. 2. Mulerial mid Melhüdü A list of iipecLmens is given in the Appendix. In the majority of cases both skin;; and skuJls were available. External measurements were recorded from the specimen labels. Their abbreviations are: HB - head and body length, TL - tail length, HP - hind foot length, E - ear lettgth, and W - weight. Ten linear measurements were taken from each of the skulls using a vernier calliper, accurate to the nearest 0,1 mm. The abbreviations used are; CbL - tündylobasai length, DiL - diastema length, MxT - maxillary tooth row length, NaL - nasal length, ZgB • zygomatic breadth. BcB ■ bra incase breadth. loC - interorbital constriction, PoC -postorbital constriction, MdL - mandible length, MdL - mandibular tooth row length (Fig.l). NaL DIL MkT CbL i-MdT-n -PoC MdL- Hg. I. Cyunial and mandibul^ measurements of Spennophilus ciselius used in thiit stud}'- See text for abbri:viiition&. St. 1. Diuintzije lobanje in spodnje feljiL^tnicc, kl so bile meijene pri Spfrmophilus citeUus^ _ü. Kry^tufek: Eurofjeaii Sousllkj {Spvimtiphilu^ cintisa) -, KoJenlia, Mammal in uF Maiadiani^ 3 Variatinm in mensural i;fiaTactern among geographic samples and sexes wctc anaiysed by using the siajidard and multivariate analyses. Standard statistical tests (mean and standard deviation) were applied in :dJ comparisons involving a single cliaraetcr. Only adults, i.e, specimens that h fid hibernated at least once, were incnrparated into statistical tests. Forsiinultantusestiniation nf pheneiic differentiatkin between sexes and populations, the raw skull measurements were subjected to discriminant function analysis. Sexual dimorphism was studied according to subspecies. The data were z-standardized and subjected to Principal Components Analysis, This kind of analysis reduces the number of variables that need to be considered to a small number of principal components; these represent linear combinations of the original measurements (Maklev 1986). 3. Kesuitfj sind Discussion 3.1. Distribulion Two isolated souslik populations ate known from Macedonia, living in strikingly different habitHts. One population is linked tfj the mounttlin pastures of the Jakupica and 22|E Fig, 2. Present distribution (black) oi moLjilain (A) and bwland (B) fxipulatitms of Iht Rurop^ar souslik Spermopbilux iiwWu.t in Miiccdonia. Scu Figs, 16 and 21 for details. SI- Z- RüTäiTjtnost (inm) planinske (A) in ni/inskc (H) piipulacijc tcliunice, UpermophUus lisdtus, v Makedoniji. Za podrobnosti ^lej sliki It) in 21- Karadžica Moumaitis, at altitudes from 2(K)(J to 2200 m, Another population inhabits the icwtands along the lUver Vardar south üf D emir Kapiju, from which its area extends further south into irorthern Greece (Fig. 2). Distributional details are discussed for the individual subspecies- Further three regions have been reported in the literaurc to be populated by soushks. As none of these reports have been confirmed by subsequent field studies, they have been largely ignored (Ru2rt^ 1967, 1978). Recently PtTROV (1992) considered these to represent historical records of the places where sousliks had become extinct, providing a reason for reviewing them here. 1. Doflein (1921), who worked in Macedonia during World War mentioned sousliks in Üsküb {= Skopje). Approximately 15 years after Doflein's stay in Macedonia, Karam an (1931) wrote that sousliks did not live in the Skopje Basin, despite an abundance of suitable habitats there. Karam an. who went to Skopje in 1924, was closely familiar with the Macedonian fauna, and also knew soushks from the Kara£zica-Jakupiea Mts. Consequently, it is highiy unlikely that sousliks, if present around Skopje, would have escaped his attention. Since Doflein's reports are generally reliable, this one most probably results from a mistake. 2. Another dubious report is by GradoJEVIČ (1923) for Mt. Pelister. He claimed to have found sousliks in 1925 around the lake ''Vehko Jezero" (= Gole mo Ezero), altitude 2000 m. in one of his later paper; (GkaijOj^viO 1931; cited in PETROV 1992) additional information was published for Mt, Pelister. For the vicinity of Golemo Ezero, one dead specimen was reported, while in (he foothills of M(. Pel iste r, between the villages of Nižcpole and Bacila (altitude 90(1 m), sousliks were supposed to be already cxtinct before Gradojevie's survey in 1925. Gradojevlč was very familiar with sousliks (he had supervised souslik control measures in southern Macedonia), consequently, his report could hardly result from misidentification. On the other hand, V, Martino, the greatest authority on the mammalian fauna of Macedonia between the two war^, did not mention sousliks at Mt. Pelister in his list of the mammals of "Southern Serbia" (= Macedonia; Martino 1939). It is worth mentioning that Mani no and Gradojevtc were in contact with one another. In 1928 Gradojeviif even collected sousliks in southern Macedonia for Martino, who named them after their collector (S, c, gradojefid). Martino also visited Mt. Peüster in 1937 (MarTJNO 1937). Sousliks were not found in Mt. Pelister by any mammalogist collecting there in later years: A. Ružič in 1967, B, Petrov in 1972 (Petrov 1992), and myself in 1990. The higher altitudes of Mt. Pelister are still grassy, thus no considerate habitat change (a common reason for Lhe extinction of local souslik populations in the recent times), is likely to have occurred in Ml. Pelister during this century. 3. '["he third report is hy TkOMNOVlČ (1931) for Sveti Ni kole in the field OvCe pole (eastern Macedonia). It is based on the observations of burrows which supposedly belonged to souüliks, and thus has little credibihty. Therefore, there is no clear evidence for the extinction of entire local populations of the European souslik in Macedonia. 3.2. Taxonomy Two subspecies have been described from Macedonia: S. c, graäojevici (V, and E. Martino 1929) from the lowlands along the Vardar River, and S. c. karamani (V. and E. MAfiTtNO 1940) for Mt. Karadzica. A', c. gmdajevici was described on the basis of its large and robust skull (CbL 43 - 46 mm, average 44.5 mm; ZgB 29.5 - 32 mm, average 30.4 mm) and a more yellow, uniform colour, fn the original description, S. C. karuftiani was compared only with the nominate race. Its diagnostic characters were as follows: _B. kj)rätiiii:k: European SoJSlik& jSpermopkilia dielluif, Rodenlia. MaillTiriaiia of MaLVdania 5 larger zygomatic breadth, broader brainease, weaker moJars, and relatively shorter Jo wer luDthrüw, V. ami E. Martino (1940) also prepared a key for the determiriatiun of the four races of the European iouslik. The dichotomy of firadojevici versus 1 he other three races (the nominLite one and also karamani atid iaxkatevi were considcd) coticemed coluur (back, belly and taiä> and size. S. c-. ^radojevici v/na larger than any other race, Maetino (1939) explains size differences between lowland and mountain subspecies of the Macedo-[lian sousiifcs by "Hinton's rule", according to which size is greater in animals from regions with a longer vegetation period. Such an explanation is consistent with the hypothesis [hat body size follows the duration of the annual pulse of productivity, and predicts small body sizes at higher altitudes (Gfist 1987). MIRIČ (197Ü) discriurinates the two Macedonian subspecies by the size only; HE is said to permit clear distinction (above of 38.5 mm in gradojevki and below 37.5 mm in karamam). R.UŽ1C (197S^ confirmes the validity of both taxa, but provides very few diagnostic characters. S. c. gradojevici was diagnosed as a large race with a pale pelage. Siniilar conclusions were made by Pf7TROv (1992). Karyoiypc was found to be quite stable amongst the six S. citellus subspecies from south-eastein Europe, being characterised by a diploid number of 2n = 40, and fundamental number NF = Sfl {Soi.datovič et al. 1984). The only interpopulation differences involve the position of the centromere on the X and Y chromosomes. In both Macedonian races, the X chromosome is submetacentric with a submediai centromere, and the Y chromosome is the smallest submetacentric (SaviC et al, 1971). On the basis of the frequency of occurrence of 21 discrete nonmetric cranial traits, 1 (KryäTUFEK 1990) give a higli value of Mean Measure of Divergence index (MMD) between the two Macedonian races, m marked contrast to the low MMD's of between Pannonian populations. As suggested by MMD, divergence between the two of the Macedonian races was of approximately the same duration as was their isolation from Pannonian populations. As a first step in morphomctric analysis, the two raccs were subjected to discriminant analysis. The results were in accordance with the geographic origin of the samples (Fig. 3) and both laces were classified into Eheir actual groups (Table 1), In S.c. gradojevici, 81 % of specimens were allocated to the actual sex, and 83 % in Sx. karamani. The two Fin- 3. Projection or male and female ^iousliks of S. c. jfrarfo/eviti and S. c. karamani on tht first two diM-Timinant fjnctiims. Polygons cnclose üoo-res for al] irdividiials within a group, and aymbols for sex are placcd on grokip ttntriiids SI. 3. Projekcija samcev b samic teku nie S. e. ^radnjerici in S. c, karämani na prvi dve diskrtnii-nfictjski funkciji. Poligoni obkrožajo vm; osebke v skupini. Simboli ^a spol so na skupinskih centroi-dih. DFZ Sr«f(j(»¥(ci lurarnarl / Cf J Y / \ ^ / \ 0f1 races were discriminated by the first discriminant function, which was responsible for S6.6 % of the variance in the original data set, The sexes were discriminated by discriminant function 2, which explained 12.1 % of the variance, (n both races, males had higher Scores for discriminant function 2. It was evident lhat phenetic distances between subspecies strongly exceeded sexual dimorphism within the subspecies, consequently, Ihe recognition of the two subspecies has reasonable grounds. TaLle 1. Qassification tabic for discriminmit nnjilysis of [nale and feniale soiiKltks of 5. c. gratiojevid and S. c. karamnni. Rows are actual groups and colunins are predicted groups. Tabela 1. Kiaiifikacijska label a zi diskriminacijsko analszo samcev b aamic dvch podvrst teku nie iz MakcJDnije. Visticc so dejanske, slolopci pa napiwcdant- skupine. Number Group n 1 2 3 4 1 males b S 1 2 (^radojeviii females 15 3 12 3 /caromani males 15 14 1 4 karatnafii feiraleš 21 2 19 [n the tiext step r tried to extract mniphometric characters that permii simple a ad faithful discrimination between the two subspecies. Arithmetic means of 8 measurements (W, HF, and six crajiial dimetismTis) differ significantly between subspecies iti both sexes (Table 2). Eight characters showed significant differences in males, and 10 in females. T-fests were higliJy significant (p<0,0()lj for the majority of these variables, i.e. six in males ^nd eight in females. In both sexes, t-test values were highest for mandible measurements [MdT in males; MdT and MdL in females). Among external characters, weight and hind fnot length were found to provide the best discrimination between the two subspecies. Only the weight of spring animals before parturition (gradojevki, karamaiii) dr just after it (gradojevici) was available to me, Consequently, the weights of the two raccs were comparable, Hind foot length, although a usefLiI diagnastie character, did not prove to segregate all specimen;; of graäojevid from all karamani, as suggested by MitilČ (1970). Three cranial dimensions (MxT, MdL. MdT) showed the least overlap between the two subspecics. Table 2. Es^limated t-stati&tics with probabilities (p) for inteistibspc(^fic differences in soii^liks from Mucedonia. Sexes are treated separately. n.5. - not significant; * p 8.661 0.000 ««m« NaL 2.033 0.053 n,s. 1.234 0.225 n.s. ZgB 3.399 o.no3 4.831 0.000 **** BcB 1.74J 0.960 n.s. 3.658 0.000 loC 2.J18 0.045 * 2.850 O.OfJ? ++ PoC 0.540 0.594 n.s. 0.910 0.369 n.s. MdL 6.629 0.000 10.94] 0,000 MdT 10,539 0.000 9.463 0.000 B. KiyStiifck: European Suu^liks (Spemopliilus ciiel!us)\ Riwfcntia, Manmalia of MsoL-dunia 7 Better results for the means of discrimination between gradojevid and karamani were obtained by bivariate plots of selected variables (Figs, 4-9), particularly by plotting ItiC against MdT (Fig. 9) and MdT against MxT (Fig. 7). Colour was frequently used to distinguish subspecies of European sousliks, summer pelage mainly being used für ttutunomit purposes. In early spring, just after hibernation and when they appear on the surface, sousliks are of different colors /see Ph.trov (19fKl) for i^rmlojevici, and Plate I for karamani/. I'he summer colour of the upperparts was described as "more yellow and uniform" with "the speeklcs nearly absent" in gradoje^iči and as "more greyish or brownish, with Fig. 4. Bivari&tc ?n;atter plot of weight Etgaini;! heart and Ixidy Jcnglh for Spermofihilus diellas gradojevici (sijuares) and S. c. karamani (circles) from MaoeiJonia. ia. 4. TeJa kot funket ja tclcunc Jol^jnc pn lokunici lipermophilu.1 citeliiis gradojevici (kvadrati) in Sc. karamani {krxjgi) tz Makedonije. o 4 I» I a I 3aS a »radojmlei A ktmnenl HF u a*- 31- P" O .O o HB Flg. 5. Bivariate scatter pint of hind foot length against head anrt btidy length fur S. c- gradojei ici aid S, c. kommani from Macedonia. Explanation as fat Fig, 4, SI. 5. Dolžina ülDpala J:iit /unkcifa dolžine teJe^a pti J- C. gradpjevici in S. c karamani iz Makedonije. Simboli kot n;i sliki 4. A. tiivariatc scatii^r plot of maxillary' tooth row length against coTlrtylohasai length of skull for S- C. gradojevici and S- C- karamani from Macedonia. Explanatloti as for Fig, 4, M. 6. Dolžina gornjega niza zob kot funkcij» Vondilobazalnc dolžine lobanje pri tekunici 5. f. groäojei/ici in c. karamani iz Makedonije. Simlxili tut na sliki 4, MiT ti O o o I v % 411 CbL lighter cinnamon-pinkish buff speckles" in karamii/ti (Martino V. & E. 1940), YeJlüwish summer coat was also taken to be a diagnostic character of gradojevici by Ružtč (1978) and Petrov (1992). The underparts are reported by M art! mo V. & E. (1940) to be buH-yelluw in gradojevici and cream-buff or buff-wliitish in karamani. According to MtRlČ (197Ü), karamani has the same coloration of the belly as t^radojevki^ i-e- pale buff-yellow. As regards the tail, Martovo V. & E. (1940) describe a "dark subterminal band on the tail h air-pencil'* to be "scarcely developed, usually light brownish" in gradojevici and "clearly developed, blackish" in karomani. The subsequent authors did not mention this character- MdT oa 8» .oS.Sso' KiÜ. 7. Bivariate scatter pi o i of mandibular tooth ruw Icngiti against max ill a rj tooth row length for S. c. graäojei'id and S. c. karumutti from Macedonia. Eiplanalion aü Fi^t Fig. 4. SI. 7. Dolžina gomjeiia niza lob Vot funkcija spodnjega niza iiob pri tckunici S- c. gruitajevici in S. C. karamani iz MakcOfjuije, Simhnii kot na i'i m*t üliki 4. MdT Fig. S. Bi vari ate plot of mandlbukr tooth row length against mandible length in S. c. gntdojevici and S. c. karamani from Macedonia. Explanation as for Fig, SI. 8. D»liUrba spodnjega nisa rab kot funkcija Jdižlni: spodnje Seljustnice pri tekunki S. c. gra-dajevki ia 5. c. karamani iz Makedonije. Simboli kot tia sliki 4. \ a o < '"o o o 8° ® MdL loC oe DO oa o o C t a a □ " a □ □ □ □ M);. Bivarlate plot of interorbital constriction again^t mandibular Imtti row length for 5. c. R gradojevici and S. c. karamani from Macedonia. Explanation as for Fig. 4. Medofnifna Sirina lobanje kot fjnkcija dol2ine ^piidnjc felj ustnice pri tek uniči S. c. gra- _ dojevici in S, e. karamani iz Makedonije. Simtroli MdT kot na sliki 4, □ B. KiyStiiEfck: European SouslitJ {Spetmaphilia dfettus)-, Rodenlia, Mamtnalia of Maaadtrnw 9 In a siniukaiieoüs comparison of 20 aduit summer pelts of grudojevid with 5 adult summer pelts of karamani, I failed to find any differences between the two, Not only did the palest specimen belong to gradojevici, but so did the darkest one. A single para type of gradojevici from Martinu's collection (PMS 6029) was amongst the palest sousliks from Macedonia, and the only adult with a buff belly. In the other adults of both subspccies the belly was either yellowish or yellowish white. A buff belly was otreerved also in seven subadults of gradojevici, all collected in July, As regards colour, 1 agree with Mmrč (1970) that there are no significant differences between kammani and gradojevici. The tail, particularly on its ventral side, is paler and more yellow in gradojevici. Also, the subterminal band was usually present in karamani but missing in gradojevici, as statetl by Martino V, & E. (1940). This character is best seen from the ventral side. Martino v. & E. (1940) also mentioned two ratios in which karamani is said to differ from Pannonian sousliks: Ratio 1 = CbL r ZyB Ratio 2 = CbL : MdT The mandibular tooth row is said to be relatively shorter in karamarii (hlglier vaJues of Ratio 2). In all 26 adults of gradojevici Ratio 2 was below 4.7, whilst one quarter 25 %) of 36 adults of karamani had Ratio 2 equal to 4.7 or lower. In this character S. c. karamani appears to be more variable than gradojevici- Scores for Ratio 1 broadly overlap between the two subspecies. S. c. karamani is also more variable in this respect (Fig. 10). Martino V. E. (1940) describe the lower b{)rder of the zygomatic arch of karamani as usually without a convex lobe. In the specimens available to me, the zygomatic arch is frequently thinner and with a straight ventral margin in karaTnani, and heavier with a convex margin in gradojevici, but the overlap between the two extremes is considerable (Fig. 11). When comparing karamani with the nominate subspecies of citetiwt, Martino V. & E. (1940) note that the "furrows on the surface of the upper molars /were/ longer, occupying larger part of a transverse diameter of the tooth" in the former. This character is strongly affected by tooth wear, i.e. the furrows become shallower as wear advances. Consequently, 1 compared only specimens with unworn or moderately worn teeth, TTie furrows of the first and second upper molars appear to be deep in the majority of specimens of both subspecies (Table 3). In this particular point I was unable to distinguish karamani from the Pannonian sousliks. The few skulls from eastern Serbia that 1 examined also had deep furrows. Fig. 10. Blvariale plot of two ratios for S. c. gradojevict and Ä. č. karafnani from Macedonia. See text for further i:xplanalian. Symliols as for Fig. 4. SI. 10. Odnos med dvema indck^aitia pn tekunici i'. t. gtadofevici in 5. c. karamani iz Makedonije Razlaga je v besedilu. Simboli lot na sliki 4, Cbi 1* o o o cbLhwr Hg. 11. Shape of left zygomatic acüh in 5. c. gradojevici (left) and 5, c. karamani (right). An-tcriqr is to ihi left, PMS numtwrs. I^ft columnr 705,*!, 6100, 70SS, 7053,6111. Right column: 709«, 7089, 7fm, 7074. 7(W5, SI. 11. Oblika levega liirega loka prt tckunici S. C- gradojevici (k^o) in S. c. kaiamarti (desno). AnlCTioma stran je le>o. l able 3. Occurrence of the two niorphotypes, based on the development of the furrows on M' and M' ill the two sjbspccies of sousliks from Macedonia, Furrow deep: deeper than one hall' of the transverse diameter of the molar; furrow shaliow: shallower than one half of the transverse diameter t)f tlie molar. Tjihela 3-. Pojavljanje dveh mcjrfodpuv gjedc na globino zajede na povriiili M' in M^ pri dveh podvrstah te kuni C iz Makedonije. S.c. gradojevici S.c. karamani M^/M^ MVM= Furrow deep 22J21 19/19 Furrow shallow 212 4/5 Among 25 norimecric (epigeiietic) cranial i;haracters listed by me (KftYäTlJFEK 1990) from five souslik populations from Serbia and Macedonia, the foramen on the maxillofron-ta] suture was found only in 5. c. karamam. Three other epigenetic characters also indicate divergence between gradojevici and karamani {Table 4). lable 4. Ftcqüenciei of oecurrcncc of ftiyr ironmetric Cranial traits ifl two subipecies of sousliks fronn MACEtXJNtA faea>rding to KryStufek 1990). TabI,; 4. Frequencies of occurrence of four nonmetriL; cranial traits in t^o ^ubspeci« of souiliks from Macedonia (according to Kryätufek 1990). S. C. gradojevici S. C. karamani n 19 27 Foramen on niaxilloliontal suture 0 2Ü.4 Lacrymal foramen 0 40.7 Anterior frontal foramen 2.6 75.9 Foramen sphenoidale laterale 32.4 fl _B. Kryjtufek: European Soiisliks {lipermtjphHua f.i>flafJj^R«clt:nlia, ManiTnalia qf Macudcmia LI Several new characters could be introduced to assist in dicrimination of the two Ma;;£:doiiiaa subspecies: Skull shape - Skull of };radojcvici is more angular; supratemporal ridges fused into a short posterior crest in more than one half of ali adult skulls. In karamani the crest was found in only one of 27 skulls. i'ig. 12. Ventral side of rostrum in (left) .S. c. gradojevki (PMS 7054) and (right) S, c. karamani (PMS 70W). SI. 12. Spudnjü Ätran ruütruina pri lekunici (levo) S. C. grad/tjevici in (dcEno) .V. c. karamiini. Hg. 13. Nasal bones in 5, c, gmdojevid (upper now) and S. c. karamani (lower row}, PMS numbers (from left to right). Upper row: 7048; 7058; 7053 ; 7047, Lower ruwj 7066; 706^; 7077; 700], SJ. 1.1. Oblika nosnice pri Lekunici .S. c. grudoje-zki (2giirnja vnita) in J. c. karamani (spodnja vnta). Foramina tncisiva - Longer in karamani. Distance between the anterior margin of for. jncisiva and the posterior alveolar TTiargin of the upper indsor shorter than for. incisiva in karamani, longer in half of specimens of gradojevici examined (Fig, 12). Nasals - Broader posteriorly in ^radojevici (Fig. 13). 3.3. Sexual climiirphL«im Secondary sexual dimorphism has been reported in European sonslik, with the males being larger. According to Ružič (1978), the differences are most obvions in CbL, HF, HB and W. In my material, two external and six skull dimensions differ significantly (p<0.05) between the sexes in graäojevici, and one external and seven cranial measurements in karamani (Table 5). The externüJ characters were less affected by sex than the cranial ones. Head and body length only showed significant intersexual differences in karamani, and HF and W only in gradojevid. Of the skull dimen^iionii, CbL Dil, Zgb, BcB, loC, and MdL differed significantly in botii subspecies, while there were no intersexual differences in MxT, MtiT and PoC. Tabic 5. E»Uinatcd t-statistics with prubdbilitics (p) for inEcrsexual diffcienccs in süusliks trum Macedonia. The subspecie? are treatfd separately. Explanation as in Table 2. 'I'äbel;) 5. Vtcdfiüstl t-testa pri pdmerjanjiL dveh podvrst tckuni^ Makedonije, glede na spol. C. gradojevici t-tcst p S, c, karamani t-icsi HB 0,929 0.365 n.s. 2.0S9 0.044 + TL 0.8(18 0.429 n.s. 0,512 0-612 n,s. HF 3.103 0.006 0.079 ns. E 1.391 o.iso n.s. 1.466 0,151 n,s. W 2.151 0.046 * 1.404 0.173 n,s. CbL 4.714 0.000 3.811 0,000 DiL 2.630 0.014 * 2.540 0,015 It MjiT 0.640 0.528 n.s. 1.555 o,m n.s. NaL 0.519 0.609 n.s. 3.774 0,000 **** ZgB 3.967 0.000 3.670 0.001 HcB 2.521 0.019 * 4.404 0.000 ioC 3.77T 0.004 3.306 0,002 PoC 0.603 0.552 n.s. 1.587 0,121 n,s. MdL 3.643 0.001 5.152 0.000 MdT 1.898 0.069 n.s. 0.929 0,358 U.S. Skull dimensions were subjected to Principal Components Analysis, separately for i;ach Subspecies. The variance explained by the fit:it two principal components amounted to 65.3 % in ^radojevici and 68.2 % in karamani (Table 6). These were relatively low values, since approximately one third of the variance in the original data sets remained unexplained by the first two components for both subspecies. In both subspecies, the highest character loadings for the first principa) component were for CbL, DiL, ZgB, BcB, and MdL. The first principal component (PCI) is defined as that vector in hyperspace _Ü. Kryjlufek: Eunriptian ScusUfc-i [Sp^rmaphiliis dlfllus)-, Kudunlia, Mamina lia of Matgdwiia 13 TabJc 6. Chamcter lojujiii^ for Lhe first tiAiü principal components in the two sjbspccic^ cf sugsJik^ from Macedonia. Tab«l» 6, Vrednosli locfieifcnluv la prvi dve giavni komponenti pri dveh podvrälah fekunic i?. Makedonije. S. C. grsdojevici A. C. kiirantani Principal component 1 1 J. 2 Percentage of variance explained 47.9% 17.4% 48.9 % 17.4% CbL 0.920 0,239 0.859 'O.OÖ6 DiL 0.759 0.461 0.697 0.434 MxT 0.422 -0.176 0.299 -0.777 Nal 0.572 0,233 0.699 0.212 ZgB 0.913 0.813 Ü.230 BcB 0.755 -0.179 0.855 -0.115 oC 0.491 -0.613 (1.845 0.286 PüC 0.050 -0.S91 0.24Ü 0,454 MdL 0.S63 0.246 0.888 -0.040 MdT O.ftSS -0,311 0.365 -ti.736 Hg. 14. Projection of 24 specitDens S. c. graäoje^ via on the first two principal components. Closed symhdis - males; open - females, SI. 14. Projcfcdja 24 primcrtov tekunice S- c. gradojevici na prvi dve glavni komponenti. Polni ijimboli - sanici; prami - samice PCJ PC2 Fig. 15. Projection of 35 specimens of i*, e. karamani on Hie first two principa! compoTicnts, Explanation as for Fig. 14, SI. 15, Projekcija 35 primerkov teku nice S. c. karamani na prvi dve glavni kompioncilti. Simboli kot na sliki 14. □ a □ □ □ ■ □ C ■ ■ ■ ■ □ PCI 0 • • o" • * • • ^ S o o o > o o o •o* ' DO pci which explains the muximuiti püäsible variaiiun oS the dutti (Lemen Since highest character loadißgj for PCI were by CbL and MdL, this component was highly size-correlated. Consequently, it is explained as a "size factor '. The first two principal components for the sous I i ks are shown in Figs. 14, and Males had lii^cr scores tor the firet component in both subspecies, resulting from iheir greater size. The overlap between the sexes was much more evident in kamtnanu with the smallest male approaching the size of the smallest females, i.e. specimens with the highest negative scores for PCl. With regard to the second principal component there was no segregation between the sexes, suggesting that the differences between the sexes were primarily those of size > not of shape. 3.4. Subspecies Spermopkitu-s ciielluM gradojevici (V, & E. Mnrtino, 1929) 1929. Citdlus citellus grudojevici V. & E, Marli no Hololypc. - An adult male. Zoological Institute, St. Petersburg No. 33,844, skin and skull, obtained 3U May 192«, Type not seen. Type locality. - Gevgelija ('Djevdjelija' in older transliteration), Macedonia. En the original description, the type locality was misspelled as "Djerdjelija", Geographic distribution. - south-eastern Macedonia; northern Greece along the Axios River as far south as Pieria (Fraouedakis-Tsoles &. Ondrias 1985, Vohra].ik & sofiantdou 1987); Petrič in south-western Bulgaria {PeSev 1955. Makkov 1957). In Macedonia, this subspecics is known along the River Vardar in the triangle Udovo - Oevgelija - Lake Dojran, and in the valley of the River Strumica near the town of Strumiea (Fig. 16). Along the Vardar River, soushks are associated with dry, warm, open places with deep soil, which are regularly grazed by domestic animals (Figs. 17-19); wh i It they are absent from stony, ertided slopes of hills, as well as lowlands with a high groundwater level. Diagnosis. - Size larger; maxillary tix^th row ^9.8 mm; mandibular tooth row ^9.3 nim; mandible length 3=29,2 mm. Ratio of mandibular tooth row to condylobasal length ^4,7. SktiU more angular, frequently with supra temporal ridges fused posteriorly into a crest. Nasals broader posteriorly. Foramina incisiva shorter, placed more posteriorly. Ventral side of tail uniformly buff-yellow, Measurements, - Listed in Table 7. Remarks. - First specimens were collected in 1917 by the German herpetologist Lorenz Mliller. Two skulls, housed in the Zoologische StaatssamnJung München, are labeled "Kaiukorva" (Fig. 20). "file name of the locality is Turkish, and if the letters "rv" are spelled as "w", we get "Kalukowa", This place is probably identical with "Kaluckova", as indicated by Doflein (li>21, Fig. 52, page 97). Ka lue kova was still to be found in the Rečnik-Imenik naseljenih mesta u Kraljevini Jugoslaviji" of 1930, bt)t it is absent from the imenik naseljenih mesta u SFRJ" of 1985 (M. MlLElfliOVlC, personal communication). It was situated east of Udovo, i.e. within the known current distributional area of sousliks In southern Macedonia, Doflein (1921) reportes sousliks around "Hudova" (= Udovo). His statement is probably based on the two specimens collected by Müller at Kaluckova. Doflhin (1921) B. Kry^tufck: European Soysliks {üpermopiillus citellus): KodeuLla. MnmiriFilia of Matedoiiia 15 l'1g, 16. Distribution of S. c. graiiojevici in Macedonia, Sliaded - lowlands; triangles - mountain peaks. List of localiites: 1 - Udovo; 2 - Kalucfcova; 3 - Mimvci; 4 - Anska Reka: 5 - 1,5 km SE Rabrovü, 6 Smokvicc; 7 - GreiStc; fi - Prdejci: 9 - Gevgelija; JU Bogdartci; IJ - Stoja kovo; J 2 - Bogorodjca; U - 3 km F- Bugonjdka; 14 - Star Dojran: 15 - Nov Uojcan: 16 - ACiko:; 17 - Nikolid; 18 - 4.5 km NW Stnimicii (records "bei Strumica. nordwestlich dsr Stadt, am Fusse des SmrdeS-Gehir^cs" /Kužt«^ "Stnimica" /RUŽttf 1i at Dojranskt} H^era Lake on Müy yih m^ by D. Strc. PriUtga I. Odrasi;! primerek .Spermt/pftilus diclius graiiojevici. Fotografiran v Ačikotu nü nojranskcin jcztru, miijii iWJ (Foto D, Sorc). Ftij. 17. High dqnsily hjbilat of c. fimäojemi at Atikoi, lakt Dojratisko Ezcro, SI. 17. HubiWt I viBoku populucijükü gostuio tcktiiii« .V c- grailnji'vidpr'i Aiikotu. na obsili Dojranskega Fi|ij, 18, Emnitcc lo ihL- dt-n of S. c. firmiojevici with faML's in from uf it. Detail liniii iJii. liHtiilm in Bg. 17. M. IN. VIidJ V rov De kun ice S. r. grtitinjeviü. De taji s ^like 17. U. Ki ySlufekj Lüiopean Stiuslik^ {ü/tennfiphilnx dtelliay, RociL'nliii. Mammalia uf MaCKiitiniLt W Fig. 19. Low density huhiltll of S. k. jfmtlnieviä ncili" Bogdanci. Sl. ly. Habirat z riiko ixjptilacrjsko ^Oülutd tvkunlt-' S, c. s^nuitifevici. Okirtica Bogilarccv, GfSfJiltchi: lljj- Lubiils uf two suusliks. collccled by Lore/ij; Müller in 'Kalutkova', Maecdunia (Zutiio^isclic Slaatssammlung Miiilirhcn). SI. 20, Ftikeli teku nie, Vi ju je Lijreiii Müller ujel na lokacij! 'Kaluzktiva' v Makediiniji. Plate II. Spermophilux tiw/Zui graciojirviLi. Variability in coloration cjf ihc summer coal, l^ronl; PMS 6tl2y, Lpack: FMS Pri]»)^ 11. Speimi/philus cilcllus gradojevici. Variabilnci^t v barvi poletne dlake. Spredaj: PMS, zadaj: PMS 705S. B. Kry^tufek; European Sousliks {Sperinophiiu)! riMuj); kwlentia, M;!tnnialiä of Mscedonia 21 Plate nr. Spenmipkilus ciieUia gradojtuki. Left; aduSt female t PMS 60^8), right: young (PMS 6104). Note thi: white üput on ihc friint of (he young animal. Pillug^ III. Spermaphiluü citfäus gradojevici Levo: odrasla samički (PMS 6WH), dcsnci: inlaid if (PMS 6104). MUdif ima na ätriu belo pego states that sousliks were rare iiroLtnd Udovo. GradojiiVič (1928) reports that ibcy became II pest in the same area "in the last years", and their numher was controlled by fumi^arini' their dens. In the period between 1947-49, Rl:ži:č (197^) estimated population densities around Gevjielija ai 6.1 sousliks per hectare, and aronnd Flogdanci at .I/ha; the highest densities were on pasture^;. Twenty ye;irs later [1965-1%8). lower densities were recorded, njumely, 3,2/ha in Gevgelija and 1-2/ha in Bagdanci [Ružlč i.e.). Such detreases were the result of abondoning pastures and their cultivation, tuminti most of them into vineyards and tleJds. In the subsequent decade, the trend for population declines continued, (n 1977, densities were estimated at 0.9/ha in Gevgelija 0,4/ha in Bogdiinei, [n three decades, the popu i a ti on decreased hy 3-H times fRUŽlC I.e.). Towards the end of the 19Hl)'s. I found only a single large population (over 1(K> sous I i ks) at the tourist set t lenient of Ačikot at Dojransko Ezero Lake. Small colonies (about 10 itnimals each) were spread thinly iüong the northern shores of the iake, and along ihe Vardar, They populated small clcarings uninng.ilawiens. SaUgetieTkund, Mit!., 8(1/2); 1-12. + FRAGLiEDAKtS-iSOLlS, S. E., W77: An immunochemical study of three pc^pulations of the ground squirrel, Ciieilus ciieilus, in Greece. Mam m ah a, 41(1): 61-66, praguedakis^tsolis, S, E.. J. C, Ondrias. l*)«"?: Geographic variation of the ground squirrel Citelius citellus (Mammalia: Rodentiü) in Greece with a description of a new subspecies, Saiigeticrkund. Mitt., 32: 185-198. Geist, V. (1987) Bergmann's rule is mvalict. Can. J. Zool., 65: 1035-1038. GradojeviČ, m,, 1928: Najezda tekunica u Juinoj Srbiji. Priroda i nauka, 1(6): 133-137, Beograd* GradojeviČ, M., 1929: Borba proti v tekunioa u Južnoj Srbiji. Prakt. poljopr, pouke i kalend, za 1929 g. Beograd (not seen). GradojfvjC, m., 1931; o naSiim teku niča m a i izvršenim ogledi ma za njihovo uništavanje. Beograd, pp. 1-14 (not seen). * CiPADOJEVič. M., 1936: Suzbijanje tekunica t hrčkova. Novi Sad. pp. 1-29, Kar AMAN, S., 193 h Le bassin de Skoplje au point de vue zoologique. Glasnik Skopskog nauCnog druätva, 10(4): 214-241. Skoplje (In Serbian with Freneh summary). Karam aiv, s., 1937: Fauna Južne Srbije. Spomenica dvadesetpetogodi^njice oslobodenja Južne Srbije 1912-1937, pp.: 161479. Skopje. * Kranjčev, R.. 198S: Na Jakupici. ■ Priroda, 7(i: 204-2118. Zagreb, _B. KryiiufeJt: üurofeaii ftcmsük,': (.S/jg^up/ifj^i ciielhis)., RthJentia. Mammaiia of Mitctdoniii 29 KRYSTVJPtK, B., 1990; Nonmetric cranial variation and divergence of European sousJiks (Citellus cilellus) from Yugoslavia (Rodenlia, Sciuridae), Boll, Zool., 57; 351-355. KryStUFEK, B-, S. PETKOVSKI, 1990: New records of mammals frnm Mucedonia. Fragm, bale. Mus. mactd. sei. nal., 14, 13/306: 117-129. Skopje. LemeN, C.L., 1983: The effectiveness of methods of shape analysis. Fieldiana Zoul., IS: 1-17. M AN LEY, BFJ., I'JSb: Multivariate statistical methcxis. Chapman and Hall, London, pp. 1-159. Markov, G., 1957: Untersuchungen über die Systematik von Cirellux c.itellus L. - Izvestja na zool. institut, 6: 453-490. Sofia (In Bulgarian with German sumraar>). * Martino, v, B-, 1930; Notes on the ecology of some mammaJs from Jugoslavia. Zapiski Russkago NauCnago Instituta v Bjelgradc, 2: 53-65. Bjelgrad. Marti ivo. v., 1937; Novosti u zbirci. Lovac, H/12; 257-258. Beograd. Martino, V., 1939: Materials for the ecology and zoogeography of the mammals of S. Serbia. Zapiski Russkago Naufnago Instituta v Bjelgrade, 14; 85-106- Bjalgrad (In Russian with trigl. Summajy). Martino V. and E., 1929; A new souslik from Macedonia. J. Mammalogy. lOf I); 76-77. Martino, V. and E., 1940: Note on the Yugoslavian ground-squirrels (Sousliks), Ann. & Mag. N. Hist., 11(5); 465-471. London. * MiriČ, D., 1962: Angaben über Erstbeschreibungen und Typus-exemplare von Saügetier- fornien die (bis Ende 1961) vom Territorium Jugoslawiens beschreiben wurden. Glasnik Primdnja^küg muzeja. Ser. B. 18: 159-192. MilUČ, D,, 1970; Ključi za določevanje živali, Sesalci - Mammalia. Inštitut za biol. Univerze v Ljubljani, pp. 1-132, * Niethammer, J. 1986; Ueber griechische Nager in Museum A. Koenig in Bonn. Ann. Naturhist. Mus. Wien. 88/»9(ß): 245-256. * Ondrias, J, C., 1966; The taxonomy and geographical distribution of the rodents of Greece. Saügetierkunii. Mitt,, 14; 1-136. Pe.SHEV, Z., 1955; Investigations in systematics and biology of Ci tell us cifeWüv L. in Bulgaria. - Izvestja na zool. institut, 4^5; 277-325, Sofia (In Bulgarian with Eng. summary). Pbtrov, B- M,, 1941): Zamjetki po sistema t ike i ekologiji mljekopitajuSčih Juiiioj Serbiji, Zapiski Russkago NauCnago Instituta v Bjelgradje, 16: 57-64. Bjelgrad (in Russian with Eng, summary). * Petrov, B., 1979; Some questions of the ^oogeographical division of the western Palaearctic in the light of the distribution of mammals in Yugoslavia, Folia topI,. 28(1); 13-24. PErRov, B. M.. 1992; Mammals of Yugoslavia, Insect i vores and Rodents. Natural History Museum in Belgrade, Special Issues, 37: 1-186. Beograd. * Ružič, A., 1965: Sistematika, rasprostranjenje, ektjlogija i privre dni značaj teku nice Citcilus citellus L. u Jugoslaviji. Unpublished PH. D. Thesis, Ljubljana, pp. 1-145. RuŽJČ, A., I96fi: Odre diva nje uz rasni h kategorija u populaciji teku nice Citellux citeUus L, Arhiv biol. nauka, 1S(1): 65-70, Ružrč, A., 1967; Die Verbreitung des Ziesels. CileUux citellus (Linne, 1758), in Jugoslawien. Saugetier k und. Mitt., 15(2): HI9-1I2. Ružlč, A., 1969: Neue IJaten über die Verbreitung des Ziesels, Uiellus ciieilus (Linne, 1758), in Jugoslawien. Saügetierkund. Mitt., 17(4): 369-370. Rltžič, a., 1978: Citdlus dtelins (Linnaeus, J766) - Der oder das Europäische Ziesel. In: ,). NitTHAMMEK & F. Kraf? (eds.) Handbuch der Saügetiere Europas, Akademische VcTlagsgesellsehaft Wiesbaden, pp, 123-144, RUŽeC, a1^179: Decreasing number of the ground SijUirrel (Cileiius citcllus L,) populaiiuns in Yugoslavia in the period 1947 to 1977. Ekologija, 14(2): 185-194. Beograd (in Serbian with Eng. summary). RuŽIČ-PEtkOV, a., 1950; Prilog poznavanju ekologije ickuniee Citelius chellus L. Zbornik radova Instituta za ekologiju i biogeografiju, 1: 97-140, Beograd. SavjČ, i., jvj. mll-OŠEVlt, S. ZJVKOVIC, 1971: ChromnSDmes of ground squirrel (Cäeilux citclius Linnaeus, 1766) from Yugoslavia. Arhiv biol. nauka, 23(1/2): 35-37, Beograd. soldatovlc, [}., D. zimonjtc. 1. Savi^, E. GjaüIA, 19ji4: Comparative cytogenetic analysis of the populations of European ground squirrel {Cirellus lilcilus L,) on the Balkan peninsula. Bull. T. LXXXVJ de I'Acad. Serbe des Sei. et Arts, Sei. nat., 25: 47-56. TrojanoviC, S., 1931: Tekunica (Citellus dieHus). Priroda, 21: 53-61, Zagreb. VOHRAl.!K, V., T. SOFIANIDOU, 1987: Small mammals (Insectivora, Rodentia) of Macedonia, Greece. ' Acta Universitatis Carolinae - Biologica, 1985: 3L9-354. * Not mentioned in the text J'ov/eiek V Makedoniji živita dve podvi^sti tekuniee. Podvrsta SpermophiliLs citellwc gmdajevici je poznani! iz trikotnika Udovo - Gevgelija - Dojransko jezero in iz doline reke Srrumice. Najdemo jo v odprtih habitatih (paJnlki) z globokim slojem prsti, kjer se redno pase živina, medtem ko je ni na e rod i rani h pobočjih gričev in na obmoCjih z visokim nivojem talne vode. Podvrsto prepoznamo po ve^ih telesnih dimenzijah, bolj oglati lobanji z izrazitim puSCičnim grebenom, nosnih kosteh, ki so v zadnjem delu äiräe, po kr^jäi nebni äpranji in po enakomerno rumeno obarvani spodnji strani repa. Podvrsta Spermophiius dtetlus karnmani živi na planinskih pašnikih Ja kupice in KaradŽice, poznana pa je s treh nahajališč. Ta podvrsta je manjäa, puščiini greben je le izjemoma razvit, nosne kosti so v zadnjem delu ožje, nebna Špranja datjSa, spodnja stran repa pa je bolj siva s temnim pasom pred koncem repa. Pri obeh podvrstah se kaže sekundami spolni dimorfizem. Samci so večji od samic. [Jimorfizem je bolj viden v lobanjskih dimenzijah, kot pa v telesnih. Ocitnejäi je pri |30dvrsti S. c. gradojevici. B. Kryitufc:k: E-urc)pean Sousliks {S^mmphilifs citetius)-, RtxigJlij. Matnriuli^ ot Macgdonia 3i Appendix The following is ü tabular presentation of the meaiuremetits of sousliks from Macedonia examined in this study, This includes ill specimens from the Slovene Museum nf Niitural Histtiry^ the British Museum (Natural History), the National Museum Prague, and the Zoologische Staatssammlung München, The column ■'Identificution" gives collection, catalogue number, locality statement ami date of collection. Five external (HB, TL, HF. E, W) and ten cranial (CbL, DiL, MxT, NaL, ZgB, BcB, ToC, PoC, MdL, MdT) measurements are listed. AM measurements are given in millimetres, weight in grams. See the "Material and Methods" section for the abbreviations of measurements. Abbreviations Collections: PMS - Slovene Museum of Natural History, Ljubljana BMNH - British Mu.seum (Natural History), London NMP - National Museum, Prague ZSM - Zoologische Staatssamnilung München, Munich Sei: M - male F - female Growth groups: Determination was ba.sed on the degree of wear of tooth enamels as examined tmder 10 tiities magnification. Growth groups are aecording to Ružič (1966). juvl (juveniles 1): age I - l.S month juv2 (juveniles 2); age 1.5-2.5 months sad (subadulls): age 3-5 months adl (iidults 1): before and just after the fli^t hibernation ad2 (adults 2); second calendar year and spring following the second hibernation ad3 (adults 3); third calendar ye sir and spring following the third hibernation sett (senes): autumn before fourth hibernation and following months lilcncificiljiMi Sei HB TI E offiftu fnijfftin PMSlSt NtnCkujiiii H 211 n iS.7 S,1 !?? NwDcj™ a jDO 71 m 11 ISS NorUcifjti H a J115 (6 m 7,5 IIS Ndv Dt^iiii 22 hl)'im F 2DJ S» 17.7 7,9 11» NuvDc^m H ZlO TI 38.9 22D NovDf^D M a; » m S,S 321 N(n ry^ao 41 M m 3S.I !S m Eiti^.AäVDt »JW^ISS? F ■ r. „ ™ m Dojraa.ACilm F 1Ü n ao m Dojra«,Atilo« F m m «,S 300 tlite Diijnt.ACiluii F m 54 J7.() 5,7 m Sim Düjmi.AEils.« f m 17.7 (J «Jin Do|raii,Afilf(il M ■m jj 3S.J S,5 300 ftiii IX^iii, Aatnt M m « 3S7 9.S 290 MM Dujiia. Aäbl SMijlS««) F 2tQ 36.(1 te.4 2JS ? Mayings M :i7 65 lOJt 7(151 Dujnii.Aeäcil U 225 a M MG ?«} Dojnn. U 2i2 » 41.0 9.5 iJO m Dnjr^D.AEibH M S3 10.3 300 m Dcjrar.Afitu - F m (a (.S 300 7Q5C F in Iš u MO ■mi lltMijiJSl F 23« a 11 27S Dojim, ^cn • V 231 a ri.4 91 260 NMPIStif Sur Do] mi tJMa/lii68 M m SUrDiqjnii F .... i8es SUrDojnn M I8W SU;[X)j'nn f i»yv SutUk^ii F [SK SurDojrar F Tl pwsam Otvjtlijii 2kf«m M B5 S3 J7,5 (.5 ßwfnijj-ijj Cevjciii F 1« « 33J (J PMSfflSS Ijiitijh MHayJütB F 23 t5 J5J t? 25 «m □tüIIi: M IS U m U 75 dim) Ciääc M Iii a J2,5 U KI HtlJ Gtcac y M m 41) 9.0 fS 6i(e (jifiäf F ijj 12,7 JO WD) CiiStt u F 115 « S.6 (D 6104 Ciäitc F m 40 J3.; 9.D 70 MB Cinüi: F 134 J2,S 1.1 jn HHT 1.3 IcmSe Itahmv ! May im F 3M 5) 39.7 1(3.0 280 imii Oc^mlu f m U J9.D 250 7*1« 3tmE Bi^iHiidica 4 F 211 äft 37,7 9.4 2ftQ Udmo u F 190 JI.II!l.Ji U4m M -r- 11 II.IIJ« Utmi u M 56 J-O }|.III1J1 Uibw M JiiniM UJam F 11.11,11 J! Uibw) tl F 213 9.D }|.[!.I1.«J lUom y F ii lfl.0 PMS «47 ItiiEStraiiiö 23111*1%? F m 30.1 i.9 U «08 4 M nil H S.6 IJ 96» 4biE Sdl 41.4 m 9.? [S.4 2ej 2.1 9.0 11.9 39.5 9.4 31.11.1 üdl 41.4 9,i 9.4 [4.? 2!.4 20.« «.4 11.7 28.(1 «.S adl 4U 1(14 lÜJ 15.1 29.2 m B.8 IIJI m 9.2 31.11.11« adl 4]J m.« 91 ISJl 29.4 ns 9.i lU 39.9 9.8 PM5W? ji-vl SJl 11.4 2i.ä IS.8 8.0 12J m ««J J5J 7A .... [1.) 2]J IS.3 7.8 Iii 24J m juv] 3(.i (1.4 .... ll.l 22.1 S.3 UJ 15.0 ÖMI m 41.1 lOJ 10.3 14.7 21J rP, U.9 S.5 I jiß 4].t) n lOlS 14j 22.3 [0.0 IIJI 2K.« it. KiyStufck: Eurupcan S^jiks [SpirrmophUns cheHu.';)\ Rpdeiitia, Mammalia of Mitcectonia 35 Iderlilicatinn oc DiL MsT KaL Z?B BcB luC PtiC MdL MJT S^rmopftrtuj rfwWiür hirmnani PMSTffiS sen 415 11.4 9.1 15J 19.j a-i 10.2 12.6 3Si ii TO Mil il.i> [D.b Lljf 331.4 21.« 9.9 12.6 M (.4 IDbl mli 4i.li la.s ^1.4 16.1 S.tt 9.4 m 2S.S B.9 Jll« ict2 4t.D 11.2 I4.1f 27.4 21.7 9J 11.4 TD« «lE 41.& id.b 9.} ISJI m 9.7 12.J 2S.J m adL »JI M 1^1.4 17.2 21.4 S.S 12.J (.6 m »11 415.} in.i 9.4 lU m 21.4 9.4 11.7 I7.1t 9.1 m aJt 4(l.fl ffi.i 9.) 27 Ji 21J S.S 12.2 I7.lt (IJ mi sdl 9.4 S.; 1S.4 21.1 E.7 12.1 17.4 9.0 m adl 45.4 L!.l 9.1 11.4 iU m [1.} 129 314 9.1 m sdl 451 m 211 lO.i 13.4 28.1 ILS a(G 41.4 in.s 9.1) M H.2 21J 9.1 11.9 27.1 Ll rau] ad] 41.? ll.l 9.3 iS.^ 3tl.7 21.4 9.i 11.4 2S.4 S.i im adl USA l(P.O 152 2T.i 21 .J R.ü 126 TT.fi 8.9 ■m afl2 411 U.S 9.5 1W 30.4 JIS 9.6 il4 S.J 9.0 m adl ll.i 9.3 17.1 B.9 23.1 las ll.J 29.5 WS ud2 112 S,1 UJ 3«.9 21J 9.9 L3.6 JfcO S.3 um «.1 10.! ».3 14.3 nt 32.3 94 211.4 8.Ü W! aiE «JS UJ 9.4 19.9 BI m 125 29.1 8.7 ■m adl 11.4 10.; 9.4 14J 18.7 217 9.4 12.4 39.1 Ä.7 um adl 59.8 IOjD 15.4 37.2 21.1 t..: m JU 9.6 II.« 27.! 6 ■11..1 lt.2 9.1 Ii.» S.6 21.4 1D.J 12J .... S.7 BMTiHfl.lUH idl 11.; 9.3 ll.l „„ 1Q.I -H-- 30.1 16.395S »11 44.il ici.7 !i.i 1S.(i SA hi lOJ 11€ 91 47. um jevl 34.1 .... .... .... .... 47,111» adl «.J 11.0 9.6 lli.7 2S.1 il.9 9.B lU S.2 J.0 4?.lil» adl 41.5 9.4 Ib.} 29.7 11.5 IS.O U.4 BJ 9J1 M.W57 43.9 itj 9.3 h.4 21.6 10.2 12.3 9.2 (I6.3ÄS8 üdl 4s.: Mi 9.J 147 32.3 9.; 12.1 28.1 9.6 ja. 11.37.1 adl J3.3 |D.i 9.4 h.4 U.3 1(1.1 11.3 39.(1 9.0 (7.11U5 juvl I-... ..rt .-rr .... .,.. W.395? jBVl S7A ,„. 47.11® M 43.6 17.4 M U,f M 9.0 iull 4J.1 lU 9.4 110 a.9 2i.i 9Ji 12.5 211-1 9.3 tt.396: idl 41.4 !l.fl m 16.3 ffi.7 11.4 9.S 12.4 a.l M.mi juvl JJ.1 „^ „„ .,„ H... 4U 1Q.7 9.5 16.7 m 22.6 U 12D 19.4 9.2 65JIW ml 43.7 9.7 9.3 li.3 36J 21.2 12.3 14.9 8.9 17.1110 »d m 10.« 9J8 !6.0 Zl.l IlJS 9.1 12.4 27.4 9J0 B, Kry^hifet; Evmpenji Soiisliks (SpemophiliLi cilellw,}'. RiHlL;ntia. MiimmEiliH of MiKsdonia J7 2A. Skull of Sperm/jphilax chetlux driidojevici {uJulL mülc, PMS 71^.1) in (ä) latcml, (b) dcin^l and (ü) ventral view. SI, [^hanja tckunice Spermopliiiun diellusgrctiUifevicU^.) z boka, (b) s hrtunc in ermophilus cUeltuf i^radojevici i(ihI (b) 5- c. kar