I GEOL. CROAT. I 50/2 1225 - 230 I 6 Figs. 3 Tabs. ZAGREB 1997 Review of the Results of Morphometric and Morphogenetic Analyses of Early Pleistocene Micromammals and Upper Pleistocene Cave Bears in Croatia Maja PAUNOVIC and OOI'dana JAMBRESIC Key words: Morphometry, Morphogenesis, Microcvo­ lulion , Stratigraphy, Cave bear, Micromammal s, Pleistocene, Croat ia. Abstract Due 10 lhe large number and visible morphological variab ility of skeletal paris of Ihe Early Pleistocene micromammals and Upper Pleistocene ursids we utilised morphometric, morphogenetic and sta­ tistical analyses in order 10 determine and process the material. The results indicate rapid microcvolulionary processes which arc rcllcC!cd both in variations of the characteristic parameters of particu lar skele­ tal parts , and their morphology. The Early Pleistocene age of the bone-breccias discovered along the Adriatic coast was confirmed by the morphogenetic analy ses of micromammals, with reference to geochronolog ically important arvi­ col ids: the fauna fro m Razvode ncar Knin comprises arvieolids char­ acteristic of the Lower Early Pleistocene, the fauna from Tatinja dra­ ga near Karlobag comprises species typical for the Middle Early Pl eistocene, while the fauna from Podumci ncar UncSic is of La te Early Pleistocene age. The results of the analyses of teeth and metapodial bones of the cave bears from (he Upper Pleistocene deposits in Vindija cave, Veli­ ka peCina and Veternica cave (NW Croatia) are expressed as morpho­ dynamic indices, namely, the frequency of the morpho types occur­ rence indicates the existence of an iJl(ermediate type (?subspecies) within the phyletic lineage UrslI.l· denillgeri - Ur.l"IIs ,1"{Jefaells, which appeared during the Riss glacial and Riss/Wlirm interglacial, and a lack of deposits of that particular age in Vcternica cave and Velika peCinil. 1. INTRODUCTION For large quantities and variability of fossil material the multifarious options for data processing provided by computer technology has helped to make morphometry a very commonly used method in thc palaeontological analysis of vertebrates. Additional morphogenetic stud­ ies which also enable the determination of relative geo­ logical age of the material and microevolutionary trends, have been carried out for many assemblages, namely families and/or taxa. In Croat ia, such investiga­ tions have intensified in the 1980 's, when processing of Croatian Academy of Sciences and Arts, Institute for Quaternary Palaeontology and Geology , Ante KovaCica 5, HR-IOOOO Zagreb, Croatia. Kljucne r1Jecl: morfomctrija, morfogencza, lllikro­ evolucija, stratigrafija, spilj ski medvjed, mikrosisa­ vei, pleistocen, Hrvalska. Saietak Zbog brojnosti i uocene morfoloske varijabilnosti skeletnih dijclova donjopleisLocenskih mikrosisavaca, odnosno gornjopleisto­ ccnskih ursida, prilikom dctenninacijc i obrade matcrijala koristene su morfometrijske, mor fogenetske i stalisticke analize. Rezullati su ukazali na brze mikroevolucijske procese koji su se odrazili kako u promjenama mjerenih karakterislicnih parametara pojedinih skeletn ih dije lova tako i na njihovu morfologiju. Pretpostavljena donjopleistocenska stmost kostanih breea knje su nadene duz obale Jadrana potvrdella je morfogenetskim anal izama fau na mikrosisavaca u kojima dominiraju geokronolosk i va7.ni arvi ­ kolidi: fauna iz Razvoda kod Knina sadrzi a rvikolide karaktcristicnc za rani donji pleislocen , ona iz Tatinje drage kod Karlobaga vrste tipicne za srednji donji pleistocen, dok je fauna iz Podumaca kod UncSica kasno donjopicistocenske starosti. Rezultati analiza zuba i metapodija spiljskih medvjeda iz gornjo­ pleistocenskih naslaga sp ilja Vindija, Velika peCina i Veternica (SZ Hrvatska) izrazeni morfodinamskim indeksima odnosno uce s!a lo­ stima pojava morfotipova ukazuju na postojanje prijelaznog varijeteta (?podv rste) u razvojnom nizu Ur,\"lIs dellillgeri - Urslls spe!aells koji se javlja lijekom glacijala R iss i inlerglacijala Ri ss/Wlirm, odnosno nedostatak naslaga ove starosti u spiljama Veternica i Velika peCina. the Upper Pleistocene bears from Vindija cave, then Veternica cave and Velika peCina (NW Croatia) began as well as analysis of the Early Pleistocene bone-brec­ cias at Razvodc, Tatinja draga and Podumci (S Croatia). The large quantity and variability of the studied material has so far enab led the establishment or Illor­ phogenelical schemes for particular taxa (e.g. Ursus spelaelfs, Micro/us), thus also allowing a relative dating of the rest of the vertebrates from deposits of undefined or questionable age with respect to the method "count­ ing from the top". This paper was presented at the scientific meeting dedicated to the 80th anniversary of the life of Professor Milan Herak, held on March 5th, 1997 in Zagreb 2. LOCALITIES 2. I. EARLY PLEISTOCENE The Early Ple istocene bone-breccias arc located in karst ic fissures along the Adriatic coast: at Tatinja dra­ ga near Karlobag, Razvode near Knin, and Podumci ncar Uncsic (Fig. 1), The bone-breccia matrix from Tatinja draga and Podumci is reddish-brown in colour, reflecting a tem­ perate climate during the deposition, while the yellow­ ish-green sandy clay matrix from Razvode was depo­ sited under colder climate conditions . The faunal asse­ mblages arc typical for the Early Pleistocene and there arc no signs of sed imentary recording of fossils trans­ portation (see: MALEZ & RABEDER, 1984; PAUNO­ VIC & RABEDER 1996). 2.2. UPPER PLEISTOCENE The Vindija cave is located near Varazdin (Fig. 1) in the sem i-mountainous terrain of Hrvatsko Zagorje. The cave entrance lies at an altitude of 275 m above sea leve l on the slopes of Mt. Ravna gora, and is a single large chamber, 50 m long, 28 m wide, and more than 20 m high. Approximately 12 m of stratified deposits were inves ti ga ted and divided into 13 distinct stratigraphic units designated A (youngest) Lo M (oldest). The layers M and L appear to date to the Riss Glacial, unit K cor­ relates with the Riss Inte rglacial (Eemian), layers J-D encompass the Last Glacia l (Wi..irm=Weichselian), while the upper three layers (C-A) are post-Pleistocene (sec: PAUNOVIC, 1988). All Pleistocene faunal asse­ mblages found at Vindija cave are dominated by cave bear remains. The Velika pecina near Varazdin (Fig. I) is dividcd into three separate chambers. Of these the first chamber is the largest measuring 18 m in brcadth, and approxi­ mately 3 m in high, while the other two chambers are much smaller. The Quatcrnary sediments are over 12 m thick and separated into 16 levels (a-p). After MALEZ ( 1986) the layers 0-1 are correlated to the Riss/Wlirm Interglacial (Eem ian ), layers k-e encompass the Last Glacial (Wlirm - Weichselian), and the upper four (d-a) are post-Plcistocene. All the strata contain skeletal rem­ ains of Quaternary mammals, with a predominance of eave bear. The Vetcrniea cave near Zagreb (Fig. 1) is the lon­ gest cave in NW Croatia with over 10 km of passage­ ways and canals. The entrance chamber is about 15 In long, 7 m w ide and 5-8 m high. Beginning from the NW corner of the entrance chamber, the " left hall " extends for 14 m and is 3-7 m wide. Quaternary sedi­ ments were excavated in the cntrance chamber and "left hall". In the most complete profiles, II geological lay­ ers were identified (a-k) which were deposited from the Riss Glacial period until the Holocene. After MALEZ (1965) the lowest layer k did not contain any palaeonto­ logica l or archaeological remains. The layer j was dated Fig . I Location map. Upper Plei slocene localities: I) Vindi ja; 2) Velika peCina; 3) Veternica. Early Plei stocene locali1ies: 4) Tati­ nja draga; 5) Razvode; 6) Podumci. as Riss/WUrm Interglacial, the layers i-e as Glacial Wi..irm, and the upper four layers (d-a) as post-Glacial. All Pleistocene faunal assemblages found at the Veter­ nica cave are dominated by cave bears. 3. METHODS In the process we app lied morphometric and mor­ phogenetic methods described in the literature (e.g. CHALINE, 1972; RABEDER, J983; MALEZ & RAB­ EDER, 1984; PAUNOVIC, 1988; KRKLEC, 1995) in accordance with the basic condi tions necessary for the reconstruction of functional-morphological changes in mieroevolutionary trends (CARROLL, 1988; BOULA­ RD,1993). 4. RESULTS 4.1. MICROMAMMALS FROM EARL Y PLEISTOCENE BONE-BRECCIAS The determination of micromamrnals from Early Ple istocene bone-breccias that were considered not to be synchronous, yielded a large number of endem ic ta­ xa in the fauna of southern Croatia (Tatinja draga, Raz­ vode, Podumci) and also geochronologically important arvicolids (Tables 1-3). From Podumci near UneSic, a site known since 1930 (sec: MALEZ & RABEDER, 1984), extens ive material discovered by Malez led to new knowledge of the genera Episoriculus, Microtus, Pliomys, Lagurlls and DiJlaromys, as well as to the stratigraphic classifi­ cation of the Podumci fauna as being of late Lower Pleistocene age (op.ciL). Four new taxa were estab­ lished: Ep isoriculus then II, Rete/lomys podumcensis, Microtus then!! and Lagurl/s arankae podumcensis , / / / Palillovic & Jambrcsic: Review of the Results of Morphometric and Morphogenetic Analyses ... 227 Localities Mammalia Razvode Tatinja draga Podumci Talpa minor + + Talpa europaea + Sorex runtonensis + + Episoriculus thenii + + Beremendia fissidens + + Crocidura kornfefdi + + Rhinolophus ferrumequinum Myotis bechsteini Myotis blythi Hypo/agus brachygnathus Lepus sp. + + Marmora sp . + Sciurus sp. + GUs sackdillingensis + + Reteliomys podumcensis Allocricetus croaticus bursae Microtus praehintoni ratticeps Mimomys malezi + + Mimomys savini + + Mimomys pusillus + + Dinaromys da/matinus + + P/iomyscf. schemfeldensis + Pliomys simplicior bolkayi Pliomys d. hollitzeri Lagurus arankae + Apodemus d . sylvaticus + Apodemus mystacinus epimelas + (MA LEZ & RABEDER, 1984) . According to CHA­ LINE (1972), RAB EDER (1981), KOWALSKI (1992) and the presence of successive stages of the evolutio­ nary lineage of the Microtus-species can be in terpreted as local evolutionary trends and therefore the new estab lished Montepeg lian Microtus-thenii-zone is also of loca[ significance (MALEZ & RABEDER, [984). + + + + + + + + + + + + +? + + + + + bursae thenii + + + + bolkayi + podumcensis + Table I List of Lower Pleistocene + fos sil mammals (aner PAUNO- V IC & RADEDER, 1996). Taxa Razvode Tatinja draga Podumci Allocricetus 31 3 4 Mimomys 3 4 1 Pliomys 14 5 36 Dinaromys 0 1 6 Lagurus 0.1 76 21 Microtus 31 7 29 Apodemus 20 4 3 others 1 0 0 Total 166 877 868 The fauna of Razvode ncar Knin contains arvicol ids (Microllfs cf. jJraeliil/(olli, Pliomys simpliciorlhollirzeri, lvlimomys pusilllls) which ind icate an Early Pleistocene age. A new species of hamster (A//ocricetlls croat/clls) is to be emph as ized (PA UNOVIC & RABEDER, 1996). The fauna of Tatinja draga near KarJobag con­ tains many more ind ividua ls and is also dominated by arv icolids (Lagl/rl/s arallkae, PliolllYs bolkayi, PlionlYs liolfilzeri, Dil/aromys da/matillus, Mimomys savilli, M. pl/silllls). Among these remains a new species Microtus eoml/iceps was determined (PAUNOVIC & RABED­ ER,1996) . T abl e 2 Frequenc ies of rodent taxa (mostly