Preliminary Report on the Late Pleistocene Small Mammal Fauna from the Loutraki Bear-Cave (Pella, Macedonia, Greece)

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Cadernos Lab. Xeolóxico de Laxe ISSN: 0213-4497 Coruña. 2001. Vol. 26, pp. 485-495

Preliminary report on the Late Pleistocene small mammal fauna from the Loutraki Bear-cave (Pella, Macedonia, Greece) Estudio preliminar de micromamíferos del Pleistoceno Superior de la cueva de Loutraki (Pella, Macedonia, Grecia)

CHATZOPOULOU, K.1; VASILEIADOU, A.1; KOLIADIMOU, K.1; TSOUKALA, E.1; RABEDER, G.2 & NAGEL, D.2

AB S T R A C T

The Loutraki Bear-cave (Northern Greece) yielded a rich Pleistocene fauna including mammals, amphibians and reptiles. In the present study the small mammal fauna associated with cave-bear remains is studied. The material comes from a long-time excavation project, which is still in progress. This study allows us to propose a Late Pleistocene age for the Loutraki fauna. The composition of the LAC-micromammalin fauna suggests a complex environment.

Key words: Rodents, Late Pleistocene, Greece, Macedonia, Loutraki, Paleoecology (1) School of Geology, Aristotle University, 54006 Thessaloniki, Macedonia, GREECE (2) Institute of Paleontology, University of Vienna, Althanstraße 14, A-1090, Vienna, AUSTRIA 486 CHATZOPOULOU et al. CAD. LAB. XEOL. LAXE 26 (2001)

INTRODUCTION-METHODOLOGY in the Aristotle University of Thessaloniki and the local Physiographical Museum of The cave-site of Loutraki (LAC: Almopia. There are three excavation- Loutraki Almopia Caves) is located in blocks of squares in three chambers (LAC North Greece on the slopes of the Voras I, LAC II and LAC Ib) of the cave. During mountain, very close to the former the excavation in 2000 a new square- Jugoslavian border, about 120km north- trench was opened in the chamber LAC III west of Thessaloniki (figure 1). The inves- in order to add data to the study. The aim tigation of this area started in 1990 due to of the research is the sediments as well as the great palaeontological interest. Five the palaeontological material of all cham- systematic excavation circles took place in bers to be correlated. Two dating projects 1993, 1994, 1996, 1999 and 2000 by of the sediments are in progress, one in School of Geology of Aristotle University cooperation with Laboratory of in cooperation with Ephoria of Archaeometry of "Demokritos", Athens Palaeoanthropology and Speleology, (ESR method) by Bassiakos and the Ministry of Culture and Vi e n n a second in cooperation with Vi e n n a University. All the material is stored both University (U/Th method). About 10.000

Figure 1. Loutraki Bear-cave, location and ground-plan. On the ground-plan the excavated trench squares are shown. CAD. LAB. XEOL. LAXE 26 (2001) Preliminary report on the small mammal fauna 487 ursid remains, which are described and clastic and chemical sediments is evident analysed from three blocks of squares, are in both columns. The calc-crust layers determined as Ursus spelaeus (TSOUKALA (oblique stripes) were deposited during E., 1994, 1996; TSOUKALA et al. , warm and humid intervals, while the clas- 1998). Other large mammalian fauna tic sediments (sand, clay, silt) were accu- remains found in association with cave- mulated during colder periods. The study bears, confer to: Panthera pardus, Crocuta of the small grain size of the clastic sedi- spelaea, Capra ibex, Dama sp. mentation of the floor of the Bear-cave is In order to study the micromammalian an evidence of slow water flow in the fauna from the chosen trench squares D10 deposition site. This is the result of the (LAC II chamber) and N10 (LAC I cham- increase of water mass surface flowing ber) (figure1), the sediments were first put inside the cave, as well as of probable cli- into water and perhydrol (H2O2) and then mate change from wet to dry (TSIRAM- all the material was washed through two BIDIS, 1998). The micro and macromam- double system of sieves, one for micro- mal remains were found in brown clay, mammals (1mm) and the other (3mm) for between the two first calc-crust layers larger, mainly ursid remains. Then the (D10). The ESR-absolute dating of the material was dried, packed up and trans- second crust indicated an age more recent ported to the Aristotle University’s labs than 50.000 years (KAMBOUROGLOU for further processing, such as dry sieving & CHATZITHEODOROU, 1999). The through a set of sieves with diameters of U/Th absolute age of the fossiliferous layer 1.6mm, 1.0mm, 0.63mm and 0.35mm ranges from 35.000 to 30.000 (dating of for easier sorting. Then the teeth were pla- the two first crust layers). ced in special plates and they were num- bered. The measurements of the teeth PALAEONTOLOGY were taken at University of Vienna, with Video Measuring System (VIA-100) Among rodents, the abundance of microscope. bone remains is remarkable, as well as the The teeth were figured in the number of isolated teeth, especially from University of Vienna as well as in the square D10. In N10 the number of teeth Aristotle University. and bones was reduced. In contrary no complete mandibles were found, only very STRATIGRAPHY few broken parts of them, with few teeth. The teeth of arvicolids (plate I) are in Two stratigraphical columns are pre- remarkable abundance. For the determi- 3 sented from the two square-trenches D10 nation only the M1 were used, but M and N10 of the Chambers LAC II and have also been described. The murids LAC I respectively (figure 2). The excava- (plate II) are following in abundance and tion in the third trench is in progress. The all the teeth from both jaws were found accumulation of sediments in Bear-cave and used for the determination of the spe- was in cyclic intervals. The alternation of cies. The same thing applies to the crice- 488 CHATZOPOULOU et al. CAD. LAB. XEOL. LAXE 26 (2001)

Figure 2. Stratigraphical sections of trench square D10 and N10.

tids (plate II) as well. The glirids (plate III) are relatively few but well representative. The spalacids (plate III) are even Vespertilionidae fewer but also well representative. Only Myotis sp. one tooth was found from the family of sciurids (plate I) (CHATZOPOULOU & INSECTIVORA VASILEIADOU, 1998). The chiropters Soricidae and insectivors are under preliminary Sorex sp. study. In the microfauna were also found few remains of amphibians, reptiles and RODENTIA fish, which could not be interpreted in Sciuridae this paper. The following taxa were found Spermophilus sp. from Bear-cave: Arvicolidae Arvicola terrestris (LINNAEUS, 1758) CHIROPTERA Microtus arvalis (PALLAS, 1778) Rhinolophidae Microtus nivalis (MARTINS, 1842) Rhinolophus sp. Pitymys cf. subterraneus Miniopterus sp. Clethrionomys sp. CAD. LAB. XEOL. LAXE 26 (2001) Preliminary report on the small mammal fauna 489

Muridae Chios (STORCH, 1975) and Emirkaya-2 Apodemus mystacinus (DANFORD & (MONTUIRE et al., 1994) is composed ALSTON, 1877) by species that show a much more advan- Apodemus sylvaticus/flavicollis ced evolutionary stage. Cricetidae The provenance of the microfauna fos- Cricetulus migratorius (PALLAS, 1773) sils is being discussed. Micromammals Mesocricetus cf. newtoni probably got into the cave from the surfa- Gliridae ce through fissures with claysilt material. Dryomys nitedula (PALLAS, 1778) Some teeth and bones show traces of trans- Muscardinus sp. portation by water. This observation rela- Glis sp. ted to the taphonomy of the macromam- Spalacidae mals could be the result of the increase of Spalax cf. leucodon water mass surface flowing inside the cave. Finally, they could be the remains of meals of various predators-owls and others rap- DISCUSSION-CONCLUSIONS tors that primarily feed on small mammals hunting over a variety of habitats The faunal list shows a remarkable and returning to the cave to digest and abundance of the small mammals: 19 spe- regurgitate their meals, and mammal car- cies belonging to 9 families. The LAC nivores which carried carcasse of their pray assemblage, only for rodents, shows a into the cave. Among the numerous bones diversity of 14 species belonging to 6 and teeth, there were sometimes more or families. Such a variable diversity is also less complete mandibles, but no complete common in recent faunas. The material skull is found, which is characteristic for from the Vraona cave near Athens does not owl pellets. This source of origin is also show such diversity (RABEDER, 1995). supported by the presence of erosion on The LAC microfauna has a very similar the enamel and dentine of many of the composition to that from the Smolucka teeth due to digestion by predatory birds. cave in Southwest Serbia (DIMITRIJE- Bats inhabited the cave. A small curved VIC, 1991). The LAC assemblage is com- drain found in B11 indicates that some pared to the material from the fissure- species of small mammals, dwellers of filling near Arnissa (MAYHEW, 1977) forests and rocky slopes could have visited and the absence of Sisista sutilis, Lagurus seasonally the cave on their own. l a g u r u s, Microtus guentheri and O c h o t o n a The composition of the micromammal pusilla is observed. The absence of Sisista, fauna concludes a Late Pleistocene age. Allactaga and Ochotona shows a less steppe Most of the species show an advanced evo- environment in view of its presence in the lutionary stage. This conclusion is confir- fossil fauna from the Bacho Kiro cave in med by the age that macromammal fauna Bulgaria (KOWALSKI & NADA- indicates as well as by the absolute dating CHOWSKI, 1982). The LAC-material of ESR (KAMBOUROGLOU & CHAT- compared to Middle Pleistocene faunas of 490 CHATZOPOULOU et al. CAD. LAB. XEOL. LAXE 26 (2001)

ZITHEODOROU, 1999) and U/Th In the surroundings of the cave and on methods. the nearby mountains, deciduous and The surroundings of the Bear-cave are coniferous forest and bushy vegetation was geomorphologicaly very complex-forest developed, which was inhabited by nume- peaks, rocky slopes, vast fields and moun- rous species such as the wood mouse tain plateaus alternate in a small area (Apodemus sylvaticus/flavicollis), the pine (figure 3). Just like today, in the Late vole (Pitymys cf. subterraneus), three species Pleistocene geomorphological density had of glirids (Dryomys nitedula, Muscardinus to result in vegetational and faunal sp. and G l i s sp.) and the bank vole variety. (Clethrionomys sp.). On the rocky slopes and neighboring In the mountain plateaus and fields mountain peaks lived high mountain spe- some species lived related to drier climate cies such as the snow vole (Microtus niva - and grassy vegetational cover, like the lis). The common vole (Microtus arvalis) souslic (Spermophilus sp.), the common vole inhabited mountain meadows, while the (Microtus arvalis), the golden hamster rock mouse (Apodemus mystacinus) lived in (Mesocricetus cf. newtoni) as well as the mole dry woodland and calcareous rocky rat (S p a l a x cf. l e u c o d o n) (ONDRIAS, scrubby hillsides. In the vicinity of rivers 1966). the water vole (Arvicola terrestris) could have been occasionally found.

Figure 3. Composition of the LAC assemblage according to ecological indicators. CAD. LAB. XEOL. LAXE 26 (2001) Preliminary report on the small mammal fauna 491

PLATE I

LOUTRAKI BEAR CAVE

Arvicola terrestris 1. M1dex. 3 3 Microtus arvalis 2. M1dex. 3. M1dex. 4. M1sin. 5. M1sin. 6. M dex. 7. M sin. 3 3 Microtus nivalis 8. M1dex. 9. M1dex. 10. M1dex. 11. M1sin. 12. M sin. 13. M sin. 3 3 Microtus (Pitymys) cf. subterraneus 14. M1dex. 15. M1sin 16. M1sin 17. M1sin. 18. M sin 19. M sin. 2 Clethrionomys sp. 20a. M2dex. 20b. labial view. 21a. M dex. 21b. labial view. Spermophilus sp. 22a. M3sin. 22b. posterior view.

PLATE II

LOUTRAKI BEAR CAVE 1 1 1 2 3 Apodemus mystacinus 1. M sin. 2. M sin, 3. M dex. 4. M dex. 5. M sin. 6. M1sin. 7. M1dex. 8. M2sin. 9. M3dex 1 1 2 3 Apodemus sylvaticus/flavicollis 10. M sin. 11. M sin. 12. M sin. 13. M dex. 14. M1dex, 15. M1dex. 16. M2sin. 17. M3dex. 1 2 3 Cricetulus migratorius 18. M sin. 19. M , M sin. 20. M1sin. 21. M2sin. 22. M3sin. 3 Mesocricetus cf. newtoni 23. M sin. 24a. M1dex. 24b. labial view. 25. M2sin. 26. M3sin.

PLATE III

LOUTRAKI BEAR CAVE 4 4 1 2 Dryomys nitedula 1. P dex, 2. P dex, 3. M sin, 4. M sin. 5. M1dex. 6. M2 sin, 7 M3sin, 8. M3sin. 2 Muscardinus sp. 9. P4dex. 10. M dex Glis sp. 11. M3dex. 1 3 Spalax cf. leucodon 12a. M dex. 12b. labial view. 12c. lingual view. 13. M sin. 14a. M1dex. 14b. labial view. 14c. lingual view. 15. M3dex. 492 CHATZOPOULOU et al. CAD. LAB. XEOL. LAXE 26 (2001)

PLATE I CAD. LAB. XEOL. LAXE 26 (2001) Preliminary report on the small mammal fauna 493

PLATE II 494 CHATZOPOULOU et al. CAD. LAB. XEOL. LAXE 26 (2001)

PLATE III CAD. LAB. XEOL. LAXE 26 (2001) Preliminary report on the small mammal fauna 495

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