CODEN SCPLEK ISSN 03S1 - 0077 PRIRODOSLOVNI MUZEJ SLOVENUE MUSEUM HISTORIAE NATURALIS SLOVENIAE SCOPOLIA 38 Bobinica Geologies & Palaeontologica Museologica 24 Zoologica Jan ZIMA, Miloš MACHOLÄN, Boris KRVŠTUFEK, Svetozar PETKOVSKI: Karyotypes of certain small mammals (Insectivora, Rodentia) from Macedonia Kariotipi nekaterih malih sesalcev (Insectivora, Rodentia) Makedonije SCOPOLIA No 38 pp. 1-15 Ljubljana Dec. 1997 SCOPOLIA Glasilo [*rirodüslovnega muzeja Slovenije. Izdaja EVirodoslovni muzej Slovenije, sofinancirala Jta Ministrstvo za kulturo in Ministr&ivo za znanost in tehnologijo. Uredniški odbor: Breda ČINČ-JUHANT, Janez GREGOR! (urednik), Huw ORIFHTHS (GB), Boris KRVŠTUFEK, Lojze MARINČEK, Jgnat SP/EC. Kazimir TARMAN. Nikola TVtrTKOVIČ (HRJ in Tone WRABER. Lektorja: Cvetana TAVZES (zaslovenäfiiiü) in HelenaSMOLEJ (ja angleSčino). Nas!ovuredniätvain uprave: Ptixodoslovm muzej Slovenije, 1000 Ljubljana, Prešernova 20. Izideta najmanj dve Številki letno, naklada 600 izvodov. Tekoči račun pri LB ät, 50 100-603-401 Ii. T^sk. tiskarna Tone TomSiC d.d., Ljubljana. SCOPOLIA Journal cf ihe Slovenian Museum of Natural History, Ljubljana, Ekiited by the Slovenian Museum of Natural Hisiürv, subsidised by the Ministry of Culture and Ministry of Science and Technology. Editorial Staff: Breda CINČ-JUHANT, Janez GREGORI (Editor), Huw GRIFFITHS (GB). Boris KRYSTUFEK. Lojze MARJNČEK. Ignac SIVEC, Kazimir TARMAN, Nikola TVRTKOVIČ (HR) and Tone WRABER. Readers: Cvetana TAVZES (for Slovene) and Heiana SMOLEJ (for English). Address of the Editorial Office and AdministiBdon: Slovenian Museum of Natural History. SI 1000 Ljubljana, Preäemava 2fl. The Journal appears at least twice a year, 600 copies per issue. Current account at LB No 50100-&0340115. Printed by tiskarna Tone TomSiC d.d.. Ljubljana. SCOPOUA Nd3»>pp. MS. P«. Karyotypes of certain small mammals (Insectivora, Rtxlentia) from Macedonia Jan Zima', MiloS Macholan', Boris Kry^lufek^ and Svelozar Petkovski^ ' Atiulftmy of Sciences; of ihe CzE^h Republic, Instituic of Animal Physiology ami Geneiics, Vevcri 97, CZ'602 00 Bmo 2, Czech Republic - Slovenian Museiiin of Natural History, P(5 Bo* 29€, SLO-10()I Ljubljaoti, Slovenia ' Macedonian Nalucal History Museum. Bulkvar [linden MAC-91000 Skopje, Macedüniu Rcccivcd: 18.11.19^7 UDC(UDK) 599.331599.32:575.1 K497.17) (ABSTRACT - Slafldard and hanckd karyotypes are repüncd for 14 üpecies of irseusivoiieji and roderLs ReNullJi are based on ^ i^peciinejis coliecU;)! üL 5 localiüeü in wEKtern Mat^ediinid:. Hcuvx aratieus, S. minutuj/. AT^omV-f/ft/tMj', Tn/frrt ifßittfli'ici, DMiifvrtiys hagdnncn t, C/ft/iimiiamyj gla/vrtlu.t, C/itun/jiny.ttii\vliji, Aficrotuf arvatis, W. subrerraneus. h'aniufipaltui leuaidijn, A/nulemus .fylvalicutA flovicoltis.A. myxladitus, andDi^'f?«fiv.T ttitedula- Key words: karyotypes, small TnaTnniaJsi. Insetitiviira, Rodenn;!, Matcdunia IZVLEČEK - KARIOTJPI NEKATBKIH M Aim SESALCEV flNSECTIVORA, RODENTJ.J^) tZ MAKEDONIJE - PiutlüUivljcni so Mandardnl in pdsasto obdiMiai kuriotipi 14 vrst žuaojedt»' in gloda]i.i:v. RczLltaii so iusnovani na W priniertih, r brani h v zahodni Miikedaniji: Sorex unineui. S. «ninHru.v. Neitmys fodieni, Talpa stankovicl D'maromys hogämv^i, Cietknonomyi ^lureotus, Chioiumiys Micwius armlli, M. /luhlerraneus, Nannnspalax leuci}don, Api'firmus .iyh'aüeu.i,A. j]avicf)lti.l, mymacinux in Dryomrsilitedaln. Kijutirtc besede: kariuiipi. maJi sesaici, glodaioi, Makedonija InlroductLüri During ü mammal survey of Macedonia we also made an attempt lo karyolype insecCivones and rodents. We supposed that chromosomal data wojid be of help in evaluating taxonomic positions of certaivi taxi as well as in understanding phyletic relations among them. Thus, we karyoEyped 99 individuals of 14 speeits of inseetivores and rodents: three taxa were analysed from their type localities. The aim of this report is to summarise synoplitally all the data obtained until now. Matertyl and \felhud$ The maicrial originates from live localities (Fig. 1): 1 - Ml Pellster, Magarevo (lOOO m above sea levei); 2 - Mt, Pelister, Rotinska reka stream (1590 m a.s.l.}; 3 - Mt. Pelister, Lake Golemo Ezero (22IS m a.s.l,); 4 - Mt. Galičiea. bttweßn LeskoCLr and Trpejca (1600 m a.s.l.); 5 -Ml. Bistra, Carcva eeima, i.e. between Mavrovo and Galifnik (t580 m a.s.l,). Small i:iammals were collected by live-traps or by diggnig (moles and mole rats). The animals examined auf listed in Table 1, ChroTTiosome preparations Irom somatic cells of bone marrow and spleen were made in the field by a modified flame-drying technique (Ford & Hamenon 195tj). G- and C-banded karyotypes were prepared in selected specimens by modified methods of Skabrioht (1971) and SumnER (19721. In a male mole rat. meiotic chromosomiei were prepared from testes by a standard lethnique. SCÜPOLlA,No3R T^ble I. Survey of üpecies and indivicJualK examinetl. See wxt und Fig. I fw idsnüticalion nunibeni of c{}llecting site». Specits Sorex anineux 5(?HEy[ ininitius Neomys fodkm Tulpa stmkvvki Oirtammyn bogtianm'i Ctethj-iaitOmys glareolus Micmttis arvalis Micrvlus sublerraiieas 00. indiviclunJs Ifl locality no, 2 2 2 2 S 5 2 5 2 Chianotnys nivalis Nannospalax teucodon Apodetnus sylvaticus Apodcmus flavicoRis Apodemus myslac'mus Dtyomyn nUedula 7 2 2 1 4 2 36 2 I J ] 3 4 1 3 5 5 2 3 4 4 2 Specimens are preserved as .tiandard skijiü and skulls; ±ey are deposited in the collect Ion of B. Kry^tufek. Fig. 1: Map of Macedonia with collecting silen. S« text for locality identifications. ___Jan ZL\< A, MiloS MACHOLÄN, Boris KKV^TUFEK, SvtilQmr FETK0V5K]: Küfiutipi...__3 Results and Duxuiision Sörejr araneus Linnabljs. J758 In 3II animals from Mt. PeÜSler, autosomes and Ihe XY^Y^/XX sex chrtumosome sjfilem were t'ound. The arm to mW nation in the bianned autosomes were ab. he. jl, and lu fae& Seariü at al- 1991 for the chromosome nomenclature). The Y^ was a small metacentric. The autosomal sei of shrewsr from Mt. Pell ste r is similar to that of populations from the vs-estern Alps (AcTocemric or Cordon race, Haujseret aL 1991), However, the metacentric Y, is a unique feature of the Pel ister population (Macholan et aL 1994). iSsrej; miltulus Linnaeus, 1760 The chromosomal complement of a female from Mt, Pclister included Mi autosomes and two acrocentric X ehromu.Homes, Tllere iVere live lai^e and one small biarmcd aulosomal pairs, and 14 acrocentric autosomal pairs fFig. 2). 11 II 11 IIII 11 It U II *A II X X Fig. J. Kjiiyuiype of Sorei mrnutuif from Mt, Pelister. This karyotype is the same ;is described from Mt. Pel ister by PktkOV et al. (1983) as well as from most other European localities (ZtMA & Khal 19^4). Neomys fodiens (PENNAsrr, 1771) The karyotype included 52 chromosomes. The sis« aid centromeric position in the autosomes Wis similar 10 fmdings reported frtim tri her regions of Europe {Zima & Krai. 19544). The X chromosomc was metacentric and the Y chromosome acrocentric. Tiilpa stankorici V. Maftinu and E, Martino. 1931 The diploid number of 34 chTomosomes was found in two males; the specimen from Magarevo originated from the type locality of tiK species, All autosomes were biarmed, mosi of them being mcta- and submetacentric. A large pair was subtelocentric, und a medfum^sized pair was aero- or subteloeentric. The X chromosome was metacentric, and the Y chromosome was dotlike (Pig. This karyotype corresponds with those of T xtanlccvid described in other sites of western MaeedtJnia (Totxjßovic' et al. 1972, 19S7), Specimens from NW Greece differ in the position of the centromere on 14th and I6th chromosome pair {Soldato^^h? et at. I9K6J. iü n rm^ n «it r, it tf. H HA t X y Fig. 3. Karvtrtype of Talpa itankuvici from Magült^ü. Dinam/wys bo^danovi Martino, 1922 Specimens were collccted from the lypt locajily of D. k grtbenscihfvi Martko, 1943. The diploid number of 54 chromosomes was found in both individuals studied. All autosomal pairs bul one were acrot^entric and the smallest pair was metacentrii:. Distinct short arms were visible in most of the acrocentnc autosomes. Compared to the other autosomes, the largesit acrocentric pair was distincily bigger. The presumable X chromusomes were large subtelocentrics (Fig. 4), G- u H flO u nil fn AH Itn rtA yn »A AA AA AA Aft OA /•O OA A A *A H ( Di vw X X Flg. 4. Kfliyulypfi üf /iitiummy.'! k/gdanui t from Ml, Bistra. _Janjpv^. Miloš MACHOLÄN. Boris KRYSTUFEK, S vetozar PETKOV SKI; KariOtipi..._3 bitiuJing help to identify most of ihi homologous chromosomes (Fig. 5). C-banding showed the presence of dark centromeric bands in all acrocentric autosomes, with the exception of the lai]gest pair. The short arms of acrocenttics stained apparently darkly in certain pairs. A distinct centromeric C-positive band was observed in the small metacentric autosomes. The proximal region of the long arm of the X chromosome was C-positivt This heterochromatic block comprised approximately one th i cd of the long arm; the large size of the X chromosoirve i s thu s probably due to heterochromatic area (Fig. 6). ■ Hi »I nm It 11 u u Hif ft^ ft« M ft» M ft« «A »K ft? Fig. S. G-baiuied fcar>'Qtype (Jf Ditiammvi btfgdanovi. !f 1? 3f U f. it 1 • )) X X a V If »f I' Ii It M II 11 1» X X FIr. (t. C-barded karyotype of Dinarmtys bogdtinofi. The convcntiunajly stained karyotype appears similar to those described in D. bogdanovi from Macedonia (Sar plan in ei), teniial Büstiia and Monrtnegro (Matchey 1956. SavicJ et al. 1966. 1967, Djluc! et al. 1971) Available data radicale a conservative character of the karyotype in this relict vole. Chrornosfimal banding pattern in D. bogdamvi is described here for the first time. Clethrionomys giareoius (ScHREBEt!, ] 760) The material originates from the type locality of C g- macedonicux Felh-n and Storch, 1965. The karyotype with 56 L'hromoaomcs was ascertained in all individuals studied. In Ihe autosomal complement, 26 acrocentric and one small metacetttric pair was present. The X chromosome was acruccntric and (lie Y chroniosofiie inetacentric (Fig. 7), IS ti n in n Al n QC H 1« to 00 id M 1« (•1 SA 1» AI II 1« A« * ft I « X r Fig. Karyotype of Clethrionamys gtfiivoias frotu Mt. Pelister. The appearance of the autosomes i& the same as described previously in various regions of former Yugoslavia (SAVitJ et al. 1%8, Živkovt(5 et al. 1975a, Raimsavuevic et al, 1990). Variations, however, were reptirtedin the centromeric position in the Ychrommome. Živkotič et al, [1975a) tound acroccntric and metacentric Y chromosomes in populations from former Yugoslavia. The two chromosomal races were supposed to be separated by the Danube River, and tbe acrocentric Y chromosome was found only in the populations occurring on the left, i.e. .southern riverside, Raixjsavuevi^ et al. (1988) reported a case of intrapopulation variability of Y chromosome in a population from Mt, Jastrcbac. Serbia. Our results from Mt. PcJister revealed the Y chromosome to be metacentric in all males studied. Chionomys nivalis {MAErrtNS, 1842) The diploid number of 54 ehromosomes was found in all individuals studied. All the autosomes and the Y chromosome were acrocentric, rhe X chromosome was a large submetacentric (Fig. 8}. The same karyotype was described in most other iocalities (Zima & Kral 1984). including those in former Yugoslavia (T0noRo\'ic et al. 1971, Rad(3SAVLIEVIC et al. 1990). Microtus ormlh (pallas, 1779) The diploid number was 46 chromosomes in four females studied. The autosomal complement consisted of four large and 13 small meta- and submetacentric pairs, one large subtelocentric pair, and four small acrocentric pairs. The X chromosomes were meditim-sized metacentrics (Pig. 9), Jan ZIMA, MiloS MACHOLA.N, Boris KRYSTUFEK. Sveiozar PETKOVSKT: Karitnipi... u u u m II OD «0 •• •• «0 It U XX Flg. S. Karyotype of Cftißiromyr flivd/ii from GoJemo Ewro, Ml. ftSisler. The karyotype is of standard type found in imny areas in Europe (Zima & Kral I9S4), including former Yugoslavia (SavtiJ et al. 1971, ŽrvKCfvrc et aj. 1975b, Radcisavuevic et al, 1990). A higher proportion of small aero- and subteloceniric autoiomes was recorded in Popova Šapka, Sar planina Mts. (Živkovi^ et al, 1975b). Both, the population of Seit planina Mts. and the population studied by us (Mt. Bistra), are at the very margins of the species distribution range (Petrov 1992), Macedonia lowlands are poputaiecl by a sibling species Micmius rossiaemeridionath Ornev, 1924 (Žmtovnf & Petkov 1974. Ružidet al, 1975, ŽivKovitf et al, 1975b). Fig. 9. Kaiyotype of Micmtas arvatis ftom Mt, Bistra, Microttts subtermneus (de Selvs-Longchamfs, 1836) The karyotype of the Pelister population was idfinncal with tbat of 52-chromosoin^l race of M. xubierroneus in Europe. The Y chroiEosome was relatively large, because of the beteroehromatin SCOt^LU, No 38 ainpiification (KitvSnmiK et ai. 1994}. This result is in agreement with the previous study by Pethov & živkovitf (1979), Nannospatax ieucodon (Nordm^ntj, 1840) The kaiyotypes from all sites studiKl comprised 52 chromosomes. The auiosomal coniplemem consisted of two metacentric, seven submetacentric, seven subtelocentric, and iiine acrocentric pairs. TTie X ciiromosome was a medium-sized submetacentric, the Y chromosome wai one of small acrncenlrics (Figs. 10. 11). The G banded karyotype of the Mt. Bistra population is shown in Fig. 12. We found it difficuit to differenüuTe exactly between the medium sized submcta- and suhtelocentiic autosomal pairs, so we cannot exclude possibilities of intra- or interpopulation variation within this group of chromosomes. However, no clear differences of this type were observed. t -.. IPII^ tj if r. ? 1 ill, 1 I M «Eftn «m« * ft Fig, 10. Karyotype of a male Nancttpa/ax ifuLodttn frum Mt, Bistra, The karyotypes observed are identical with those described by Savič & Soldatovei; (1974, 19S4) in the chromosomal form macetJonicus. This form is distributed in Western Macedonia and adjacent parts of Greeee {Fig. 13). Two more chromosomal fonns were described from Macedonia, both having 2n=34. One of these, a form with the fundainEntal number of 88 chromosome arms {jfrumicifrifii) is restricted to the field strumifiko pole in extreme south-eastern Macedonia; the other form with NF=94 (ovcfKpalen.iis) has also a vejy restricted range, since it was established only in field Ovče pole (SoLD/jiovid & Savič 1973. SAvtd & SotJJATOVič 1984). apitilemus syiyaricits (Liknaels, !75S) A standard karyotype with 48 chromosomes was ascertained in both animals; no B chromosomes were observed. Jar ZIMA. MitüS MACHOUN.Borii (ÜtYSTCFEK, Svelozar PETKOVSKI: Kariolipi.. H Ii M j»if It ii Hü «A r* CO Ii H nft sft «ji « A u XX Flg, 11. Karyotype of » femiüa Nonaspalax /euewc/an from Ml. Öistrj, Ü II u It il 11 II fl •• il H If II II II 40 , 1« tt tl m « Flg. 12. G-banded karyotype of \anospalax leucodon from Mt. Bistra. StJt cliroiuosomes not jdentified. Fig, 13, Distribution of the macedonieui- chromosomal Fprm of NaTwospaiax leacoitoa Macedonia: 1 - Mt. Jakjpica, beiow Solunska giava (aJlitude 2200 m); 2 - between Bitola Mid Kukyrezani (500-600 ni); 3 -Magarevo (1000 mi; 4 - Mt, Pel ister, Golemo fizcro (2218 m); 5 - betwwn Ohrid and Stmga (700 m); 6 - Mt, Bisffa. between Mavroivoand GaliSiiiit (ISSOm). Greece: 7-Araissa. Localities nos, 1,2,5, and 7 are fromSAVi^ & SOLEWTOVU? (1^84). A similar kaiyoLype was reported froni Matcdonia and the neighbouring areas (Ževkovic ci al. 1969, Kral 1972, Soldatovfč et al. 1972, Vuj(^evic et al. 19S4, Gtagia et al. 19S5, VujoSevii: & ŽrvKCJVfč 1987). Apodemus flavicoUis {lAsLOiiO'k, 1834) The reprtsentative sample of this species from Macedonia revealed high incidence of the supernumerary or B chromosomes (46.2% of individuals are wiih B's), This frequency is fairly high compared to other populations Studied in south-eastern Europe {Giagia et al. 1985, VujoSevic? & ŽiVMJVid 19R7, Zima & Machoij^n J995). One or two B chromosomes were found in our specimens. Supernumerary chromosomes were either acrocentric or dot-1 ike, C-banding pattern showed all the acrocenlric chromosomes of the standard set to po&üe^is dark centromeric bands but the B chromosomes stained rather uniform. The X chromosome had a laigc C-positive segmem in the proxiinal area of the long arm (Fig, 14), Apodemus mystacinus (Danford and Alston, 1877) The chromosome complement comprised 48 chromosomes, which were mostly acrocentric. Two smaller autosomal pairs were biarmed (Fig. 15}. Jan 7.IMA, MiloS MACHOLÄN, Boris KRVStUFEK, Svetozar PETKOVSKl: Kariofipi... Fte. 14. C-banded karyotype af AfxaitumfiavicoVii fmni Mi, Peltster Two B fhiwiosornes ara prasem (luwer row). "TE^ II Ii II n Ii H II t'i II • * •• •• «A At tl. Ii XX Fig. IS. Karyotype of Apitdemusmyslac'mui fiuiH Ml. GaJifjca, Identical karyotype was reported from various regions of former Yugoslavia (Soldatovič et al. I%9, 1975). A population with only one biarmed autosomal pair was descnbed trom Greece fGiAfiiA el al. 19S5), ond a B chromosome was found in an individual from Bulgaria (Belcheva el aL. 1988). Dryamys nitedula (Pallas, 1779) The karyotype of a iingle male had 48 chromosomes. The poor quality of preparation did not allow detailed description of throt^iosome morphology, but no apparent differtnces were observed in comparison with the karyotypes described in other tegions (Žima et al. 1996). CanclusioDS Thä area of Macwioma comprises rich mammalian fauna, consisting of species of various Zoogeographie origin. Many mammals reach the border of their distribution in this area. Beside.';, a high degree of endemisni is Biso characteristic (Petrov 1992), On the other hand, Itaryotypic variation seems to he Mmited in Macedonian niammals, ard only few chromosomally divergent farms were found. Apart from the findings reported in this study, numerous data On karyotypes of small mammals of Macedonia were published, e.g. for Talp(i eumpaea and T. caeca (ToixjhovjcJ et al. 1972), Microtus gu^nfftfri (živkovic & Petrov 1975J, M. mssiaemeridiana!i.i (Ružit; et al. 1975, Živsnvič et al. 1975b). M. feUeni (živkoviti & Petrov 1974, žjvkovid et al, i975c). Mits domesTicus (DjlliS et al. 1980), and Apodemui agrarius fSoLDATUvic et al, 1969}. A specifically distinct karyotype was foiind only in MicrolUS frlteni, a species eudemic to Macedonia and the adjacent regions (Petbjdv Živ tov sd 1979), Populations with karyotypes different from those found in the other regions were reported in araneas (MaCHOLAH et al. 1994), ^ifannoxptilax ieutvdon (fonns macedonicus, vvchepolensis, smimiciensis-, Savic & SpLOA-njvic 19S4; diis paper), and Muj; äomeslicus (Robertsonian populations with 2n=36 in Katlanovo and Stnimica; DjL'Ltc et al, 1980), Tlie karyotypes of all the.se species are extremely variable through their ranges, so the presence of local forms in Macedonia is not surprising. The incidence of B chromosomes in the Pelister population afApodemusßavicollis is relatively high in comparison to other populations from the Balkans, The large V chromosonre found in the Pclister population of Microtus subrerraneus is also rather exceptional. Since the last two distinctive features of the karyotypes resulted from heterochromatin changes, they are not likely to include any phyletic information. An exceplional karyotype was further described in an individual of Mictvtus iii!u.v je potrjen kariotip, kakrSnega so ugotovili Peihov et all (1983): 4tl avtosomov in dva akrOL-entrična X kromo.soiiia. Popu lat ij a vrsle Neotfiys fodiens s Pelistra ima Standardno kromo^msko garnituro i 52 kromosomi. Od dveh primerkov vrste Talpa vittnkovki je eden dobljen na tipski lokal i te ti. Kari oi i p (2n=34), ki je ugotovljen pri iraSih krtih, je enak kot v literaturnih podatkih, ki se nanašajo na zahodno Makedonijo. V tem pogledu se ta vrsta iz severozahodne Grčije razlikuje od nominatne oblike T. Stanko vici iz Makedonije. Endemie na voluharica Dinarotttyi; (w^doflflvije bila dobljena na tipsketn nahaJaliSču 7Si pom\.\ bift^ikinovi), ugotovljeni pri vrstah, ki so tudi SirSe po/naue po variabilnem kromosumskcin kompletu {Sutvxaraneus, NumiOipaSaxieucudon. Microtus subterraneus}. V delu .so poudarjena odprta vprašanja, ki zahtevajo nadaljnja raziskovanja, References Belcheva, R. g., tofwshxa-ANCHEVA, M. N., Ata.v.«ov, N.l., 19BS: Karyological studies of five species of mammals from Bulgaria's fauna. Compt, rend. Acad, bulg, Sei. 42; 125-128, Diulič, B,. Sav]<5, 1, stti.natiovic;, B., 1971: The chromosomes of two rodent spceics, Dvlomys hni^danovi (V. and E. Maistino 1922) and Giia gHn (Linnaeus 1766) (Mammalia, Roden tla). Caryologia 24; 299-305, Diulic, B., Sold.^toveJ, B,, Dunderski, Z., 1980: Distnbution of karyotypes in Mus musciiius Linnaeus, 1758 (Roden t la, Muridae) in some region of Yugoslavia, Biosislematika 6: 203-210, Ford, C. E, &. Hamrhton. I. 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