Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe

Quaternary International xxx (2013) 1e14

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Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe Roman Croitor a, *, Krzysztof Stefaniak b, Kamilla Paw1owska c, Bogdan Ridush d, Piotr Wojtal e, Ma1gorzata Stach f a

Centre for Archaeology, Institute of Cultural Heritage, Academy of Sciences of Moldova, 1, Bd. Stefan Cel Mare, Kishinau, MD 2001, Republic of Moldova Divison of Palaeozoology, Department of Evolutionary Biology and Ecology, University of Wrocław, Sienkiewicza 21, Wrocław 50-335, Poland  , Poland Institute of Geology, Adam Mickiewicz University, Maków Polnych 16, 61-606 Poznan d Department of Physical Geography and Natural Management, Geographical Faculty, Chernivtsi Yuriy Fedkovych National University, Kotsubynskogo 2, Chernivtsi 58012, Ukraine e Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31-016 Kraków, Poland f Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Przybyszewskiego 63/77, 51-148 Wrocław, Poland b c

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

The article describes morphology and chronological distribution of late Middle e Late Pleistocene fossil remains of giant deer Megaloceros giganteus from the area adjoining the northern and eastern foothills of the Sudeten and Carpathian Mountains and some other regions of Eastern Europe. At list two forms of giant deer are reported from the Late Pleistocene of Eastern Europe. The morphological study of the comparatively small sized M. giganteus from Bisnik Cave (Poland) revealed some archaic cranial and dental characters. Another Late Pleistocene form of giant deer from Eastern Europe is characterized by larger size, moderately short limbs, and relatively long premolar series. The described forms of giant deer differ from a larger long-limbed type of giant deer with short premolar series from Sapozhok (Russia) and some other sites of Eastern Europe. Possible taphonomic biases in morphological study of giant deer from Eastern Europe are also discussed. Ó 2013 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

designate a species level taxon unit type. Blumenbach (1799, page 697) applied the species name Alce gigantean, mentioning the fossil remains from Ireland and quoting the immense size of giant deer skull and antlers (the reported distance between summits of antlers is 14 ft, 4.3 m), however, without any explicit reference to a certain publication or collection of fossils. Apparently, it is difficult or even impossible now to identify the fossil material that Blumenbach (1799) referred to. The first good scientific description of the giant deer from Ireland before Blumenbach belongs to Molyneux (1697), who provided measurements and a figure of an antlered skull from Dardistown near Drogheda (Ireland), but apparently, this is not the specimen that Blumenbach had in mind. The span of antlers of the specimen from Dardistown reported by Molyneux (1697) is only 10 ft 1000 (3.3 m), distinctly less that the antler span mentioned in the brief Blumenbach (1799) description. Therefore, the question of type material of the nominotypical subspecies of Megaloceros giganteus remains open. Nonetheless, several continental forms of giant deer described from Western Europe and mostly distinguished by the antler morphology, are based on explicitly designated type specimens. Nehring (1891) erected a new subspecies Megaloceros giganteus

The giant deer is one of the fossil species best known to the public. Paradoxically, it is still imperfectly known by specialists. Problems are revealed immediately as soon as one tries to get into taxonomical basics of the species. Van der Made (2006) points out that the taxonomy of giant deer is still unclear and its type material and type locality remain unknown. Recently, Vislobokova (2012a,b) published a detailed account on giant deer taxonomy. She regards the giant deer form from Ireland as a nominotypic subspecies Megaloceros giganteus giganteus and reports the fossil material from Ireland as a “type population of subspecies”. This expression sounds rather doubtful from the taxonomical point of view, and the International Zoological Code does not provide such a possibility to

* Corresponding author. E-mail addresses: [email protected] (R. Croitor), [email protected]. wroc.pl (K. Stefaniak), [email protected] (K. Paw1owska), [email protected] (B. Ridush), [email protected] (P. Wojtal), [email protected] (M. Stach). 1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.10.068

Please cite this article in press as: Croitor, R., et al., Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.10.068

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ruffii from the Kottbus area (Germany) characterized by smaller antler span, broader palmation and its crown tines inserted on the distal edge of the palmation, not on the anterior side as in Irish giant deer. Pohlig (1892) also described a new subspecies Megaloceros giganteus germaniae characterized by a more compact antler crown, and frontal orientation of palmations, which apparently become more visible during the frontal visual contact of their bearer with a rival. Its antler crowns are strongly bent medially and toward the posterior, the middle tine normally is separated from the palmation. Another of Pohlig’s subspecies, Megaloceros giganteus italiae described in the same publication (Pohlig, 1892), is generally characterized by the same antler shape, as M. giganteus ruffii and M. giganteus germaniae. However, its medial tine commonly adjoins the distal palmation. According to Vislobokova (2012a), subspecies names ruffii, germaniae and italiae are synonymous. Megaloceros giganteus antecedens (Berckhemer, 1941) from the Middle Pleistocene of Germany is regarded as a primitive form of giant deer and possible forerunner of M. giganteus ruffii and M. giganteus giganteus (Azzaroli, 1953; van der Made, 2003; Vislobokova, 2012b). Azzaroli (1953) granted to Berckhemer’s giant deer full specific rank. In our opinion, antlers of M. giganteus antecedens seems to be very specialized and most deviant from the initial morphological condition: the basal tine is transformed into a broad plate-shaped palmation, the middle tine adjoined the distal antler palmation, the crown tines are inserted on anterior and distal sides of palmation, and the posterior tine is very large and flat. Therefore, the general shape of distal part of antler is rather elk-like. Kahlke (1999) noticed a conspicuous morphological affinity of M. giganteus antecedens with Sinomegaceros pachyosteus from Zhoukoudian and explained this affinity as a morphological parallelism in similar ecological circumstances. Caloi (1973) reported some differences in cranial morphology and proportions between subspecies Megaloceros giganteus latifrons (Raven) from Holland (characterized by very large size, large distance between supraorbital orifices, and relatively long premolar series), Megaloceros giganteus hiberniae from Ireland, and M. giganteus germaniae (¼M. giganteus italiae fide Caloi, 1973) from Germany, Italy, and Eastern Europe. However, Lister (1994) regarded the differences in cranial proportions reported by Caloi as statistically insignificant. Van der Made (2006) analyzed the proportions of metacarpals of M. giganteus from Rhine Basin and distinguished two morphological forms: the older M. giganteus “antecedents/germaniae” with slender metacarpals, and younger Eemian M. giganteus ssp. with robust metacarpals. Both Van der Made (2006) and Lister (1994) regarded the “typical” giant deer from Ireland as one of the most long-limbed forms. According to Van der Made (2006), the relatively slender Irish form of giant deer arrived in Western Europe 10e13 ka from Eastern Europe. However, Van der Made also acknowledged that data on giant deer from Eastern Europe remained unavailable for the broad scientific community and he could not test his hypothesis in 2006. All remains of giant deer from the vast area of Eastern Europe and Northern Asia were ascribed to two Western European subspecies, M. giganteus giganteus and M. giganteus ruffii. According to Scheglova (1950, 1958), the archaic woodland form of giant deer M. giganteus ruffii (¼M. giganteus germaniae) was present in Eastern Europe and Northern Asia during Mindel-Riss and Riss. During Late Pleistocene, it was replaced by the more advanced M. giganteus giganteus (¼M. giganteus hiberniae), showing adaptations to open landscapes. This point of view was uncritically accepted by the majority of authors who studied giant deer from Eastern Europe and Northern Asia (Svistun, 1968; Kozhamkulova,

1969; David, 1980; Shpansky, 2011; Vislobokova, 2012a). However, Alekseeva (1980) suggested that the high individual variability of antlers in giant deer and lack of exact dating of fossil material makes such separation of subspecies impossible. Alekseeva (1980, p. 67) also mentioned that in some specimens (without, however, indicating any particular specimen) one antler corresponds to the ruffii type and another bears characters of the giganteus form. Despite the fact that fossil remains of giant deer are often represented by fine complete skulls and articulated skeletons (Shpansky, 2011; Vislobokova, 2012b), the attention of researchers was focused only upon very few characteristics (antlers and metapodials) that do not give a complete picture of evolution of this remarkable cervid species. Therefore, our knowledge on morphology of various forms of M. giganteus remain incomplete and fragmentary. Such systematically important characters, as cranial morphology and proportions of postcranial skeleton, generally remain unclear, especially for the mainland forms of giant deer. Unfortunately, within the limits of the present paper we cannot solve these numerous taxonomical and systematical questions. We present descriptions of recent and some older findings of giant deer from Eastern Europe, and try to determine the precise chronological provenance of some earlier described specimens in order to fill the gaps in the paleontological record of this species. 2. Material and methods The major part of material described in this article comes from the area of Eastern Europe adjacent to the north and northeast of the Sudeten Mountains and North and East Carpathians (Fig. 1). The fossil material is stored in several scientific institutions of Eastern Europe: Bisnik Cave: the material is stored in the Divison of Palaeozoology, Department of Evolutionary Biology and Ecology of the University of Wroclaw (ZPALUWr), Poland. Wiercica Cave: the material is a part of J. Czarnocki’s collection in the Geological Museum of the Polish Geological Institute, National Research Institute (PIG-PIB), Warsaw (Poland) and the collection in the Educational-Scientific Centre of the Landscape  near Pilica; Parks Complex of Silesian Voivodeship, Smolen _ Barycz, Zabikowo: the material is a part of the Collection of the  Institute of Geology, Adam Mickiewicz University (IGAMU), Poznan (Poland); Krosinko: the material is a part of a private collection (Poland);

Fig. 1. Location of the fossiliferous sites discussed in the present paper: 1, Krosinko; 2, _  ); 3, Wiercica Cave; 4, Bisnik Cave; 5, Barycz; 6, Burshtyn; 7, BukoZabikowo (Lubon vynka Cave; 8, Hadikfalva (Dornes¸ti); 9, Brînzeni; 10, Duruitoarea Veche; 11, Ofatint¸i; 12, Mizyn.

Please cite this article in press as: Croitor, R., et al., Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.10.068

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Burshtyn, Bukovynka Cave, Hadikfalva (¼Dornesti): the material is stored in the Geological-geographical Museum of Chernivtsi “Yurii Fedkovych” National University (GMCU), Chernivtsi (Ukraine); Mizyn: The Archaeological Museum of the Institute of Archaeology of the National Academy of Sciences of Ukraine (AMIA), Kiev (Ukraine); Brînzeni-1, Duruitoarea Veche: the material is stored in the Institute of Zoology, Academy of Sciences of Moldova (IZAM), Kishinau (Moldova); Ofatint¸i: the material is stored in the Museum of Ethnography and Natural History (MENH), Kishinau (Moldova). The present study also involves fossil material stored in the Natural History Museum of London (NHML), the National Museum of Natural History in Paris (NMNH), the Natural History Museum “Grigore Antipa” in Bucharest (NHMB), the Museum of Natural History of the University of Wroclaw (MNUW), The Paleontological Museum of Munich (PMM), and the Natural History Museum of the Florence University (NHMUF). The measurements of cranial material are taken according to Driesh (1976) and Vislobokova (1990); measurements of postcranial material are taken according to Driesh (1976) and van der Made and Tong (2008); the measurements of antlers and pedicles are taken according to van der Made and Tong (2008). Abbreviations used in the text: DLM, lateromedial diameter (measurement); DAP, antero-posterior diameter (measurement); L, length; dist, distal; prox, proximal; H, height; D, the largest thickness of horizontal ramus of mandible or transversal diameter. The radiocarbon studies were conducted at both the Radio and the Oxford Radiocarbon Accelcarbon Laboratory in Poznan erator Unit Research Laboratory for Archaeology in Oxford. 3. Systematic description Family Cervidae Goldfuss, 1820 Subfamily Cervinae Goldfuss, 1820 Genus Megaloceros Brookes, 1828 Megaloceros giganteus (Blumenbach, 1799)

3.1. Bisnik Cave (50 230 N, 19 400 E) (Poland) Cave Bisnik is located along the Woda˛ ca Valley near the village  in the Cze˛ stochowa Upland. The cave has the form of of Smolen several interconnected chambers and rock shelters, originally almost completely filled by deposits. Systematic archaeological and palaeontological studies started in 1991 and continue (Cyrek, 2002; Wiszniowska et al., 2002; Cyrek, 2009, 2010; Stefaniak and Marciszak, 2009; Stefaniak et al., 2009a, b). The profile of the deposits includes 20 layers with bone remains and tools of Palaeolithic humans (except for Layer 20). Cave Bisnik is the oldest cave locality in Poland with traces of occupancy by Palaeolithic humans. A total of 23 settlement levels, from the Middle Palaeolithic to the Middle Ages, were distinguished. The stratigraphy of Bisnik Cave is still debatable. The ”old stratigraphy” (Cyrek et al., 2010) assumes continuity of most strata. The new approach (Krajcarz et al., in this volume) assumes that the sequence of deposits is divided by hiatuses, and redeposition was important. Layers 1a and 1b are related to Holocene (MIS 1). Layers 1 and 2 are related to Late Vistulian and LGM (MIS 2), and 3e6 are related to several phases of the Interplenivistulian (MIS 3). The sandy layer 7 is related to the transition between the Interplenivistulian and Older Plenivistulian (MIS 3/4). Layer 8 is related to Older Plenivistulian (MIS 4). Layers 9e12 represent different phases of the Early Vistulian (9, 10 a e MIS 5a; 10 e MIS 5b; 11 e MIS 5c; 12

3

e MIS 5b), and layers 13, 13a, 14 and part of the layer 15 are related to the Eemian (MIS 5e). Layers 15e18 are related to the Drenthe (Odra) Glacial (MIS 6). Layers 19 and 19a are correlated to the Röpersdorf e Schöningen (Lublinian) Interglacial (MIS 7), and layers 19bc, 19d, and 20 are related to the boundary between the Röpersdorf e Schöningen (Lublinian) Interglacial and Saalian Glacial (Krzna Glacial) (MIS 7/8). Radiocarbon dates of layers 5e6 (Poz-46807: 40 000  1000; cal. BP (68%) 44 690 e 42 949; Poz-46  809: 45 000  2000 cal. BP (68%) 51 280 e 46 132; Poznan  , Poland) indicate InterRadiocarbon Laboratory, Poznan plenivistulian (MIS 3). Layer 7 is dated with thermoluminescence (Or-5301 TL 64.6  6.5 ka; Or-5303 TL 71.0  7.4 ka, Faculty of Earth Sciences and Spatial Management, University of Maria CurieSk1odowska, Lublin, Poland) which revealed the layer’s age as Older Plenivistulian (MIS 4) (Cyrek et al., 2010; Krajcarz et al., in this volume). The animal remains represented more than 160 taxa, from the Lublinian Interglacial (Röpersdorf e Schöningen Interglacial) to the Holocene (Wiszniowska et al., 2002; Socha, 2009; Stefaniak and Marciszak, 2009; Cyrek et al., 2010; Socha et al., 2010; van Asperen and Stefaniak, 2011; Tomek et al., 2012). The described material comes from the IV sediment series, layer 7 (70e60 ka; MIS 4 and MIS 3: Krajcarz et al., in this volume). The skull of female IV-74/1995 (the data on stratigraphic provenance are not available, possibly, the specimen also belongs to the IVth faunal complex, layer 7) is missing its rostral part and braincase and preserved frontal bones and maxillae with complete tooth rows. The breadth of skull above the orbits amounts to 187.5 mm; the breadth of braincase is about 100 mm. The measurements of tooth rows are quoted in Table 1. The profile of frontal suture is convex in its posterior part behind the orbits, and concave in its anterior part between the orbits. Nasal bones are not preserved; however, it is clear that they do not reach the imaginary line connecting the anterior edges of orbits. The anterior edges of orbits are situated above the middle part of M3 (if the tooth row is oriented horizontally). Preorbital pits (fossae) are shallow (their depth is 5.5 mm), but clearly outlined. The cingulum in upper molars is weak (Fig. 2), hardly visible on some molars. Its development is not symmetric on right and left sides and it is strongest on the right M3 where it completely encloses the lingual side of anterior part of the molar. The additional enamel folds are not particularly well expressed: only enamel spur of metaconule is present. The protocone and the metaconule of P2 are separated by a vertical sharp groove on the lingual wall of the tooth crown (Fig. 2). On the grinding surface of P2, the protocone and the metaconule are detached, with an isthmus. The inner side of metaconule is supplemented with complicate enamel folding. In P3, the protocone and the metaconule are not detached with an isthmus, and the groove on the lingual side of the tooth is less expressed. The inner side of the metaconule is supplemented with a large enamel fold. In P4, there is no sign of separation of the protocone from the metaconule, and the groove

Fig. 2. Left upper tooth row of the skull of female IV-74/1995 of Megaloceros giganteus from Bisnik Cave.

Please cite this article in press as: Croitor, R., et al., Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.10.068

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on lingual wall of tooth is missing. As in previous cases, the metaconule of P4 also is supplemented with a large inner enamel fold. The upper tooth series lengths and proportions of the giant deer from Bisnik Cave are quite similar to those of M. giganteus from Italy reported by Caloi (1973). A very weak development of the cingulum, which is hardly visible in the majority of molars, is another peculiar character of the giant deer dentition from Bisnik. Only a few specimens show a moderate development of cingulum.

Table 1 Measurements of upper tooth row of giant deer; *) the length of upper tooth series of the specimen from Sapozhok (Russia) are calculated from the figure in Flerov (1962). Site/country, collection number, source

L P2eM3

L P2eP4

Bisnik, ZPALUWr/W/97/IP-74/1995dx Bisnik, ZPALUWr/W/97/IP-74/1995sin Bisnik, ZPALUWr/W/97/1684dx Bisnik, ZPALUWr/W/97/1684sin Bisnik, ZPALUWr/W/99/2123dx Bisnik, ZPALUWr/W/99/2123sin Brînzeni, IZAM, BR-A Tuscany, NHMUF, IGF11630 Rome, Italy (Caloi, 1973) Italy (Caloi, 1973) Italy (Caloi, 1973) Italy (Caloi, 1973) Italy (Caloi, 1973) Germany (Caloi, 1973) Germany (Caloi, 1973) Germany (Caloi, 1973) Germany (Caloi, 1973) The Netherlands (Caloi, 1973) The Netherlands (Caloi, 1973) The Netherlands (Caloi, 1973) The Netherlands (Caloi, 1973) The Netherlands (Caloi, 1973) Naul, Ireland, NHML, M2326, \ Ireland, NHML, no number, \ Ireland, NHML, M28968, _ Ireland, NHML, M48833, _ Ireland, NHML, no number Ireland, NHML, no number Ireland, NHMB, no number, _ Ireland, MNHUW, no number, _ Ireland, MNHN, 1856-25, _ Ireland, MNHN, 8-VI-80, \ Dunshauglin, Leinster, Ireland, PMM, SNSB-BSPG 1956 I 494, _ Sapozhok, Russia, PIN337, _ (Flerov, 1962)* Siberia, PGPI-582 (Shpansky, 2011) Siberia, PGPI-1175 (Shpansky, 2011) Siberia, PGPI-nn (Shpansky, 2011) Siberia, KKM-51/129 (Shpansky, 2011)

141.8 143.3 140.5 140.5 140.7 140.4 151.7 140.5 139.0 140.0 154.0 140.0 141.0 156.0 153.5 153.0

59.2 60.9 59.0 59.3 61.3 59.4 66.0 61.2 60.0 57.0

145.0 149.0 146.0 145.5 149.0 141.6 143.3 150.0 153.3 153.5 148.0 147.0 150.0 150.0 147.0 144.5 163.5 178.0 174.0 150.0 174.0

L M1eM3 79.4 79.9 79.6 81.0 85.7 84.2 85.0 83.2 81.0 73.0

64.5 65.5 68.5 60.6 61.3 60.0 64.2 65.0 64.0 63.2 67.0 61.5 66.0 63.0

83.0 84.5 89.6 92.0 93.0 86.0 86.0 87.0 86.0 86.5 85.0 83.3 88.7 90.0 86.8 91.0 90.5 89.0 88.0 92.7 89.5 88.7

64.5 68.0 64.0 68.0

102.8 92.0 88.0 85.0

60.0 67.3 66.0 65.5 61.0 66.0

The lower mandible Nr. ZPALUWr/W/97/1489 (dx, Fig. 3, Table 2) is the best preserved specimen. It is somewhat damaged, with processus angularis destroyed. The teeth are on the medium stage of wearing. P3 is partially destroyed. P4 is molarized. Basal pillars on lower molars are developed and have moderate size. Only the anterior basal pillar is present on the third molar (M3). The ascending ramus is slightly deviated backwards; the lower (ventral) side of the horizontal ramus of mandible is convex. The mandible appears rather high at the level of M2eM3. Mandibular pachyostosis is rather weak. The thickness of mandible corresponds to the most gracile mandibles from Ireland that presumably belong to females (Fig. 4). The anterior part of the mandible (before P2) is horizontally oriented and situated on parallel line with respect to the grinding plane of the cheek tooth row.

Table 2 Measurements and tooth row proportions of lower mandibles of giant deer; *) the length of lower tooth series of the specimen from Sapozhok (Russia) are calculated from the figure in Flerov (1962). Site/country, collection number, source

L P2eM3 L P2eP4 L M1eM3 D

PP/MM

Bisnik, ZPALUWr/W/97/1489w Bisnik, ZPALUWr/W/97/1570w Bisnik, ZPALUWr/W/97/1731 Duruitoarea Veche, IZAM, DV280 Duruitoarea Veche, IZAM, DV281 Duruitoarea Veche, IZAM, DV278 Ofatint¸i, OF-5-923 Buco del’Orso (Anfossi et al., 2002) Val di Chiana, NHMUF, no number Naul, Ireland, NHML, M2328 Ireland, NHML, no number Ireland, NHML, no number Ireland, NHML, M26124 Ireland, NHML, no number Ireland, NHML, M2972 Ireland, NHML, no number Ireland, NHML, no number Ireland, NHML, no number Ireland, NHML, M14126 Ireland, NHML, M26124 Ireland, NHML, no number Ireland, NHMB, no number, _ Ireland, MNHN, 1856-25, _ Ireland, MNHN, 8-VI-80, \ Athlone, Ireland, MNHUW, no number Limerick, Ireland, MNHUW, no number Ireland, MNHUW, no number Ireland, MNHUW, no number Dunshauglin, Leinster, Ireland, PMM, SNSB-BSPG 1956 I 494, _ Bukovinka, GMCU, PB45 Romankovo, Ukraine, 34-1019 (Svistun, 1968) Grigorievka, MP PGPI721 (Shpansky, 2011) Dnepr, Ukraine, Nr. 3841 (Alekseeva, 1980) Krasnyi Iar, Siberia, Nr.84 (Alekseeva, 1980) Bachaty, Siberia, Nr. 6/1 (Alekseeva, 1980) Irtysh, Siberia, Nr. 21010 (Alekseeva, 1980) Sapozhok, Russia, PIN337, _ (Flerov, 1962)*

156.3 160.1 161.0 161.0

62.2 60.7 63.3 61.8

65.6 61.9 64.1 61.2

165.0 143.3 164.0 168.6 166.0 162.6 163.6 161.1 166.8 167.3 169.0 173.1 170.1 163.0 168.0 166.0 160.0 169.0

61.0 58.0 61.0 62.1 62.7 61.0 61.4 60.1 61.6 62.7 61.2 64.0 61.1 58.5 63.2 60.8 64.0 67.0

94.8 98.1 98.7 101.0 102.3 103.6 101.5 93.5 94.0 104.0 115.8 102.6 101.7 103.2 99.6 105.0 106.4 106.1 109.5 106.6 104.0 106.3 104.0 99.5 101.2

169.9

62.8

103.4

60.7

172.9 167.1 169.0

63.4 58.8 61.0

105.5 107.0 109.0

60.1 55.0 41.5 56.0

179.0 162.0

67.0 61.0

113.0 100.0

43.0 59.3 39.0 61.0

159.0

61.0

98.0

38.0 62.2

185.0

118.0

179.0

109.0

159.0

101.0

170.0

106.0

164.6

58.4

109.2

33.0 38.4 35.0 35.2 36.3 38.0 35.6 36.0 37.8 39.6 38.0 36.0 35.5 38.1 38.8 38.8 39.1 38.8 38.2 36.7 42.3 35.4

65.2 61.7 58.7 53.6 61.1 60.0 59.5 60.3 58.7 58.9 57.7 58.4 57.3 56.3 59.5 58.5 64.3 66.1

53.4

The fragmented lower mandible Nr. ZPALUWr/W/97/1731 from layer 7 of Bisnik is also characterized by a lesser degree of pachyosis and apparently belongs to a female. The lower mandible Nr. 1570 is more robust, and its thickness corresponds to the specimens from Ireland attributed to males. All the mandibles from Bisnik are characterized by comparatively long premolar series (one of the longest among the compared samples) and rather small size comparable to the smallest specimens from Ireland according to their lower cheek tooth row length (Fig. 4, Table 2). The giant deer from Bisnik seems to be a rather small-sized form compared to the sample from Ireland. Upper and lower tooth rows are generally shorter than in the sample from Ireland, just slightly overlapping with the smallest individuals from Ireland. The lower premolar series is relatively longer than in Irish giant deer, and the available material does not show overlapping of premolar/molar ratio of the compared samples (Fig. 4). According to the size of molars, the giant deer from Bisnik is among the smallest forms of the Eurasian continent.

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Table 3 (continued )

Fig. 3. Right hemimandible Nr. ZPALUWr/W/97/1489 of Megaloceros giganteus from Bisnik Cave. S, the shape of transversal section of mandible below M3.

The postcranial material from Bisnik is also rather small-sized (Table 3). The complete radius Nr. ZPALUWr/W/97/1703 is significantly smaller than the specimen from Schlutup, Germany (according to data from Pfeiffer, 1999) and still smaller if compared to the specimens of giant deer from Ireland stored in various collections. The radius from Bisnik is also characterized by the relatively narrowest upper epiphysis among forms involved in the comparison. The size and proportions of the metacarpal Nr. ZPALUWr/W/ 97/1654 (relatively narrow distal epiphysis and comparatively massive diaphysis) suggest that the giant deer from Bisnik belongs to the group of rather short-limbed forms, such as the sample from Rheinebene (Germany) and Zhana-Aul (Kazakhstan) (Fig. 5). Table 3 Measurements of postcranial bones of giant deer involved in the present study. Site/country, collection number, bibliographic source Metacarpus Bisnik Cave, ZPALUWr/B/W/97/1654 Ireland, NMNH, 1856-25, _ (dx) Ireland, NMNH, 1856-25, _ (sin) Ireland, NMNH, 8-VI-80, \ Dunshauglin, Leinster, Ireland, PMM, SNSB-BSPG 1956 I 494, _ Ireland, NHMB, no number, _ (dx) (Cacoveanu, 1986) Ireland, NHMB, no number, _ (sin) (Cacoveanu, 1986) Ireland, MNHUW, no number Ireland, MNHUW, no number Ireland, Athlone, MNHUW, no number Ireland, Athlone, MNHUW, no number Ireland, Enniscorthy (Reynolds, 1929) Schlutup, Germany (Pfeiffer, 1999) Sapozhok, Russia, PIN-337 (Svistun, 1968) Galkino, Russia, CM-9925 (Shpansky, 2011) Zhana-Aul, 0213-A/1956P (Kozhamkulova, 1969) Zhana-Aul, 0278-A/1960P (Kozhamkulova, 1969) Zhana-Aul,0279-A/1960P (Kozhamkulova, 1969) Kottbus, Germany (Nehring, 1891)

L

DLMp DAPp DLM1/2L DLMd DAPd

311.7

57.8 40.9

32.3

58.6 35.9

342.0 348.0 295.0 351.0

75.4 72.4 61.0 69.2

51.6 53.3 43.8 50.0

47.5 43.4 39.0

78.0 74.3 62.7 72.2

305.0

67.0 50.0

40.0

65.0 45.0

327.0

69.0 52.0

46.0

67.0 42.0

323.7 321.1 332.3

67.3 49.6 68.9 48.7 68.3 46.5

36.3 38.1 39.9

70.8 45.0 67.3 41.9 71.2 42.7

338.5

64.1 46.7

36.3

68.0 42.3

314.0

71.0

74.0

351.0

74.5

75.8

347.0

73.0

47.5

75.0 e

330.0

65.0

41.3

70.0

314.0

62.0

36.0

61.0

332.0

67.0

41.0

66.0

332.0

67.0

40.0

65.0

327.0

56.0

36.0

63.0

49.6 44.8 38.3 43.9

(continued on next page)

Site/country, collection number, bibliographic source

L

DLMp DAPp DLM1/2L DLMd DAPd

Romankovo, 34-1013 (Svistun, 1968) Romankovo, 34-1022 (Svistun, 1968) Radius Bisnik Cave, ZPALUWr/B/W/97/1703 Schlutup, Germany (Pfeiffer, 1999) Ireland, NHMB, no number, _ (dx) (Cacoveanu, 1986) Ireland, NHMB, no number, _ (sin) (Cacoveanu, 1986) Dunshauglin, Leinster, Ireland, PMM, SNSB-BSPG 1956 I 494 _ Ireland, NMNH, 1856-25, _ Ireland, NMNH, 8-VI-80, \ Ireland, MNHUW, no number Ireland, MNHUW, no number Ireland, Athlone, MNHUW, no number Galkino, Russia, CM-9925 (Pavlova, 1908) Metatarsus Brinzeni-1 (III), IZAM, BreB Ireland, NHMB, no number, _ (dx) (Cacoveanu, 1986) Ireland, NHMB, no number, _ (sin) (Cacoveanu, 1986) Ireland, NMNH, 1856-25, _ Ireland, NMNH, 8-VI-80, \ Dunshauglin, Leinster, Ireland, PMM, SNSB-BSPG 1956 I 494 _ Ireland, MNHUW, no number Ireland, MNHUW, no number Sapozhok, Russia, PIN337 (Svistun, 1968) Galkino, Russia, CM-9925 (Pavlova, 1908) Lysmosen, 2 (Aaris-Sørensen and Liljegren, 2004) Talus Brinzeni-1, IZAM, BR-1-63C Ireland, MNHUW, no number Ireland, MNHUW, no number Duruitoarea Veche, IZAM, DV-15 Komarowa, KO/6B/150/3 Mamutowa, Poland, nn Krosinko, KR1 111.F137 Russia, ZIN, 2101(1) (Kozhamkulova, 1969)

344.0

60.0

43.8

78.0 46.0

349.0

61.0

38.6

60.0 44.0

331.4

74.2

68.4

408.0

106.0

96.2

375.0

95.0 40.0

55.0

88.0 69.0

380.0

85.0 51.0

54.0

96.0 71.0

401.0

104.4 52.0

66.0

93.4

362.0 350.0 355.8 355.8 380.0

92.0 87.2 92.1 53.0 92.6 49.9 92.3 48.7

51.1 51.3 56.1

83.5 68.8 57.1 82.6 54.5 81.9 53.7 93.8

410.0

105.0

100.0

330.0 330.0

49.7 51.5 56.0 60.0

35.1 35.0

62.5 38.0 65.0

335.0

61.0 61.5

41.0

69.0 46.0

360.0 330.0 370.0

61.6 64.4 56.3 59.8 63.3 65.7

40.5

68.6 44.5 66.8 40.5 74.0 43.3

323.7 321.1 355.0

67.3 49.6 68.8 48.7 65.0 69.0

38.1 38.1 39.0

67.4 41.9 66.8 40.6 77.0 51.0

360.0

64.0

38.5

70.0

380.0

66.5

43.5

79.5

77.6 89.1 90.5 73.4 81.02 78.7 88.6 107.0

52.0 57.9 59.1 52.6 43.7 52.0 56.1 71.5

3.2. Krosinko (Poland) Some remains of giant deer were discovered in a gravel pit (at  depth of 6e8 m) near Krosinko village (52130 N 16 490 E), Poznan County. The material is presented by a right completely preserved astragalus (specimen number KR1 111.F137) and an incompletely preserved skull with antlers (KR1 111.F139). The 14C dating of the fragment of cranium (KR1 111.F139) from Krosinko yielded a numerical age of >45,400 BP (OxA-26793). The edges of astragalus are slightly rounded and the surface is matt. Color of the specimen is dark gray. This is one of the largest specimens found in Eastern Europe, rivaled by the astragallus 21010 (1) stored in the Zoological Institute of Saint-Petersburg (Table 3). The preserved posterior part of skull from Krosinko includes the occipital bone, temporalis, parietal, frontal, petrosum, and lacrimal, as well as the base of the skull. The anterior part of the skull is missing. The right antler is better preserved than the left, and is

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R. Croitor et al. / Quaternary International xxx (2013) 1e14

represented as a pedicle, burr, part of brow tine, and main beam. The left antler has only a preserved burr with an old fracture. The specimen belongs to a young male, as the cranial sutures (parietale temporalis) are still visible and show the fusing stage. Fractures of the skull are mainly predepositional. The edges are slightly rounded and the surface is shiny. The specimen is brown. The available measurements of the skull (Table 4) suggest that it belongs to a very large individual giant deer, rivaling the largest specimens from Ireland: the frontal breadth of the skull from Krosinko corresponds to the maximal limit of variation range of the sample from Ireland reported by Lister (1994).

Table 4 Measurements of cranial fragments of giant deer. Measurements

Antler Circumference of pedicle under burr Circumference of the burr Circumference above the burr Height of pedicle Antero-posterior diameter of pedicle Antero-posterior diameter of burr Antero-posterior diameter above burr Latero-medial diameter of pedicle Latero-medial diameter of burr Latero-medial diameter above burr Height of basal ramification Horizontal diameter of orbit Greatest mastoid breadth Breadth of occipital condyles Greatest occipital breadth Greatest diameter of foramen magnum Frontal breadth Greatest cranial breadth at orbits Least breadth between orbits Occipital heigth

_ Krosinko Wiercica Cave Zabikowo

Barycz

KR1 MUZ PIG 111.F139 335.II.20

IG-Z_ 14.F2

Right 258.0

Right 259.0

Right Left Left 282.0 281.0 294.0

307.0

294.0

331.0

280.0

292.0

301.0

36.6 85.3

37.0 79.4

99.3

99.3

93.7

109.8

97.9

94.1

101.3

78.6

83.6

92.6

91.8

93.4

99.7

80.1

82.3

77.9

ca 54.2

58.7

42.0

IG-Br 12.F1

56.5

59.1

202.7 121.3

209.3 93.3

186.2 43.4

172.6 ca. 46.5

195.4 ca. 258.3

198.9 ca. 207.4

Fig. 4. Comparison of size and proportions of lower mandibles of Megaloceros giganteus from various sites.

broad groove ranging from basal tine to middle tine was observed by Vislobokova (2012b) in M. giganteus antecedens and was regarded as a peculiar character of that form of fossil deer. However, one of us (R. C.) observed a similar sharp groove on the proximal parts of antlers in the specimen from Ireland exposed in NHMGA,

215.7 262.8 212.8

131.6

3.3. Wiercica Cave, Cze˛ stochowska Upland (Poland) _ Wiercica Cave is situated in the slope of Ostre˛ znik Mountain (50 400 N 19 240 E). The numerical age of fossil material according to radiocarbon dating is 40,500  900 BP (Poz-53605). The material is represented by a cranial fragment with right antler (MUZ PIG 335.II.20). The skull (Fig. 6, Table 4) includes the right parietal and frontal bone. The specimen is light brown. The antler is preserved as pedicle, burr, part of brow tine, and main beam. The pedicle is rather short and robust indicating the advanced age of a male individual. Modern fractures do not allow assessment of the original completeness of the specimen. Its antler is characterized by a broad and sharp groove that ranges from the upper side of basal tine to the anterior side of proximal part of the beam. Such a peculiar

Fig. 5. Comparison of size and proportions of metacarpal bones of Megaloceros giganteus from various sites: data on Bruine Bank and Rheinebene are adapted from Van der Made (2006). The metacarpal from Kottbus was described by Nehring (1891) together with type specimen of M. giganteus ruffii.

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Fig. 6. The skull fragment with right antler MUZ PIG 335.II.20 of Megaloceros giganteus from Wiercica Cave.

Bucharest, and a quite weak, but still visible, groove on the antlers of the giant deer from Ireland exposed in the Paleontological Gallery of NMNH in Paris. 3.4. Barycz (Poland) An incompletely preserved skull with antlers (IG-Br 12.F1) includes a damaged occipital part, fragments of right and left temporal bones, and a frontal bone with basal parts of antlers. The skull belongs to an adult male with fusion of cranial sutures still not completed. The left antler, with total length of preserved fragment of 110 mm, is represented by pedicle, burr, incomplete brow tine and a part of main beam. No part of the right antler further than w290 mm from the burr is preserved. The 14C dating of the specimen from Barycz yielded a numerical age of 39,800  1000 BP (Poz-52083). Apparently, the skull from Barycz represents a rather large form of giant deer (Table 4). Its frontal breadth (215.7 mm) corresponds to larger specimens from the Irish sample: according to Lister (1994), width behind orbits in males from Ireland ranges from 187 to 220 mm, with mean value 198.4 mm.

7

spelaeus, Panthera spelaea), Mammuthus primigenius, Equus caballus, Coelodonta antiquitatis, Bison priscus, Megaloceros gigenteus, Rangifer tarandus, etc. (Sergeyev, 1950). Anisiutkin (2001) carried out the new excavations in the site and revealed the presence of three layers: the upper Mousterian level, the middle Micoquian level, and the lower Riss-Würmian level. The age of the newly described layers of Ofatint¸i was based on combination of archaeological and faunal evidence. According to Anisiutkin (personal communication to Croitor), the upper stratum belongs to the Mousterian due to the presence of reindeer R. tarandus and the tools of Mousterian type, including a side scraper with a slight nucleus-like adzing (similarly adzed tools are known from Butesti, Trinca 1 and 2 from Moldova and Ripiceni-Izvor from Romania). It is not clear if those tools are from the end of Mousterian, and an earlier age cannot be excluded. Anisiutkin (personal communication) confirms that remains of giant deer are not present in the upper cultural level of Ofatint¸i. The middle layer of Ofatint¸i is characterized by typical Eastern Micoquien bifaces and, according to Anisiutkin (2001), by the conspicuous absence of remains of reindeer. This layer may be referred to the end of Riss-Würmian (Mikulian) Interglacial or even to the beginning of Würmian (before the Brerup Interstadial), ca. 115e 110,000 years (Anisiutkin, personal communication). The most probable age of the lower Riss-Würmian layer with Tayacian industry is 120e125,000 years (Anisiutkin, 2001; personal communication). Therefore, if remains of giant deer are yielded only by the middle and lower strata of Ofatint¸i, they represent the oldest paleontological evidence of this species in Moldova. The rather limited very fragmented skeletal remains of giant deer (13 specimens) from Ofatint¸i belong to four individuals (David, 1980). The most interesting and best preserved fossil is a fragment of left lower hemimandible OF-5-923 with M1eM3. This bone fragment is peculiar because it has tool-made notches on its lateral side. The mandible is slightly larger than the lower mandibles from Bisnik and is relatively thick (Table 2, Fig. 4): the ratio of mandible thickness below M2 to height of horizontal ramus at the same level amounts to 73.1%, approaching to the most robust mandibles from Ireland, while in the sample from Bisnik this ratio varies from 66.0% to 67.4%. The other specimens from Ofatint¸i are represented by poorly preserved fragments of antlers and postcranial bones.

_  (Poland) 3.5. Zabikowo, now Lubon The incompletely preserved skull with shed antlers IG-Z_ 14.F2 _ (Mus W. Zabikowo, 52 210 N 16 520 E, Polska) was originally described by Kiernik (1913). The inventory number was assigned to the specimen after the publication of its first description. The skull is represented with frontal bone, partly preserved interorbital, and parietal bone. The base of the skull is missing. Cranial sutures are in fusion stage (frontal-parietal as well as parietaletemporalis). There are no signs of gnawing or ancient cuts. Cut marks on the base of the right antler appears to be a modern cut made using metal tools, so it was ignored. Fractures are old, associated with the deposition stage, during the original preservation of the skull. Some of the edges have a modern worn edge. Specimen is light brown. The _ measurements of cranial remains from Zabikowo (Table 4) are rather close to mean values of the sample from Ireland. 3.6. Ofatint¸ i (¼Vyhvatintsy, Moldova) Sergeyev (1950) was the first who described the site of Ofatint¸i, and revealed a great similarity of the bifaces from Ofatint¸i with analogous tools from the Late Acheulian site La Micoque (France) and Kiik-Koba (Crimea). However, he did not recognize the pluristratigraphic character of the site. The most important faunal elements from Ofatint¸i are the cave carnivores (Crocuta spelaea, Ursus

3.7. Duruitoarea Veche (¼Starye Duruitory, Moldova) The remains of M. giganteus from Duruitoarea Veche were excavated from the lowermost layers III and IV. The silicic artifacts from layers III and IV belongs to the oldest industry discovered in Moldova. The archaic industry of this type called “Duruitorian” is analogous to a certain extent to the “Tayac of the Fonteschwad type” (Chetraru, 1965, 1973; Anisiutkin, 2001). The lower layers from Duruitoarea Veche were formerly attributed to the Riss, as those strata yielded remains of reindeer (David and Chetraru, 1970). The spore-pollen analysis indirectly supports this conclusion. Medianik (unpublished data) has noted pollen of some exotic Neogene holdovers in Layer III, which, normally, did not survive the maximal Riss glaciation. The layer could be dated to the early stage of Riss that is in accordance with the archaeological age. According to Anisiutkin (2001), the numerical age of lower layers of Duruitoarea Veche may be ca. 70,000 years, much younger than the previously assumed Riss Age. Lister and Stuart (personal communication) recently have obtained a younger date of 37,050  450 (OxA-19630) on a Megaloceros specimen (an isolated left upper molar M2 “17U-3”) from layer III of the site. According to the inscription of the specimen, it comes from the 17U square, which is situated before the entrance in the grotto, and might come from the disturbed part of the deposits that may not correspond to layer III

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from the depth of the grotto (Covalenco, personal communication to Croitor). David (1980) reported also some remains of giant deer from the younger layer II of Duruitoarea Veche. The dating possibilities of the archaeological material from this layer are quite limited. The artifacts from the layer II show some analogies in blades and two-side  r worked points with the material from La Sa aturi (Moldova), while the spindle-shaped arrow-heads made of antlers have some analogies with the 6th layer of Molodova-1 (Ukraine), which permitted Borziac (1983) to assume the late Glacial age of the layer. Chetraru (1965) believes that the presence of large Gravettian arrowheads in layer II suggests the early stage of Magdalenian. However, Stuart et al. (2004) revealed a striking absence of giant deer in Western and Central Europe within the time interval around 20e12.5 ka (uncalibrated). Therefore, a question arises about the assertion of David (1980) on the presence of giant deer in the Carpathian area during ethe arly Magdalenian. Chetraru (personal communication to Croitor) believed that the older bones of giant deer could have been admixed during material sorting and the preliminary study of the osteological remains, thereby explaining the chronological contradiction that suggests the presence of the giant deer in the younger layer of Duruitoarea Veche. The lower mandible (Fig. 7) thickness and length of lower molar series from Duruitoarea Veche are similar to the specimen from Ofatint¸i and samples from Ireland and Rhine Basin (Table 2, Fig. 4). However, unlike the sample form from Ireland, the complete lower tooth row from Duruitoarea Veche (specimen DV280), the only one known in Moldova, is characterized by relatively long premolar series, as in the giant deer from Bisnik. Cingulum in upper molars of the giant deer from Moldova is also very weak, as in the sample from Poland. The cingulum of upper molar “17U-3” (the maximal length of its crown amounts to 33.3 mm; the length at tooth crown base e 30.0 mm; the breadth at tooth crown base e 25.0 mm) is rather vestigial and clearly visible only on protocone. The size of talus from Duruitoarea Veche (Table 3) is similar to the larger specimens from Krosinko and Komarowa Cave. Therefore, one can assume that the early form of giant deer from Duruitoarea Veche is characterized by a “normal” large body size; however they still maintain a relatively long premolar series and weak development of cingulum. Lister (1994) reports comparatively small teeth for the synchronous giant deer from Ipswichian/Eemian Interglacial of Western Europe that, apparently, represents a quite different form than the giant deer from Moldova, a smaller Western European form.

Fig. 7. Lateral and occlusion views of the fragment of lower mandible DV280 of Megaloceros giganteus from Duruitoarea Veche.

3.8. Brînzeni-I (Moldova) The archaeological complex from layer III of Brînzeni-I is confidently dated to the early stage of Upper Palaeolithic (Chetraru, 1973). Nonetheless, the radiometric dating yielded a significant dispersion from 26,000 to 18,000 years, and some samples have yielded even younger ages (Rogachev and Anikovich, 1984). This dispersion of dates may occur because the material is possibly mixed. Apparently, the presence of remains of giant deer suggest that the third layer of Brînzeni-I is not younger than 20,000 years, considering the absence of M. giganteus in paleontological record of Western and Central Europe within the time interval 20e12.5 ka (Stuart et al., 2004). However, only direct radiocarbon dating can answer this question. The complete upper tooth row BR-A is rather large (Table 1), slightly exceeding the total length of the largest specimens from Ireland, and falling within the range of size variation of the sample from Rhine Basin of Holland and Germany (according to the data from Caloi, 1973). Unlike the Irish form of giant deer, the specimen from Brînzeni-I is characterized by relatively long premolar series, as are the samples from Bisnik Cave and Rhine Basin. The giant deer from Brînzeni-I also is characterized by very weak development of the cingulum. The single complete metatarsal BR-B from Brînzeni-I is the shortest among the specimens involved in the comparison, and defines the Moldavian giant deer as a particularly short-limbed form (Table 3). 3.9. Hadikfalva, now Dornes¸ti, Southern Bucovina (Suceava district, Romania) The site is situated in Suceava River valley (47 520 N, 25 590 E) (Late Pleistocene). The almost complete right antler attached to pedicle P-0002 (Fig. 8) that belongs to a young male was originally described by Botezat (1912). Details of the find are unknown. Due to the remains of black mud on the antler, the specimen was bedded in swamp deposits. The basal tine, the posterior tine, and the three of four crown tines are damaged. The basal tine is situated very close to the burr. The antler beam is curved and forms an obtuse angle of 137 in the area of posterior tine. Scheglova (1950) regards the angle between axis of beam and palmed part of the antler as a diagnostic characteristic that distinguishes M. giganteus ruffii from M. giganteus giganteus: this angle is less than 145 in M. giganteus ruffii, while in M. giganteus giganteus the angle between antler beam and palmed portion is more obtuse and approaches to 180 . However, there does not appear to be any significant difference in

Fig. 8. The right antler of Megaloceros giganteus P-0002 from Hadikfalva (¼Dornes¸ti).

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this characteristic between the antler from Dornesti, the shed antler of a young individual from Ireland figured by Reynolds (1929: Fig. 5a, p. 20) and the type specimen of M. giganteus ruffii figured by Nehring (1891), which, apparently, also belongs to a young individual (Fig. 9). However, one can assume that the antler bending and palmation orientation is fully expressed only in mature fully grown antlers. Above the posterior tine, the antler expands into a palmation, which reminds the typical for the giant deer from Ireland shape of antlers: the palmation is supplemented with two tines in its distal part and two tines on the anterior side. The posterior side of palmation is straight, slightly concave and has no tines. The middle tine is situated on the anterior side of beam on the middle between antler base and posterior tine. The middle tine is cylindrical, slightly

9

undulated and directed forward. The total length of the preserved part of antler amounts to 1210 mm, and the circumference of the antler base above the burr amounts to 323 mm (see Table 5). The shape of the specimen from Hadikfalva is quite similar to the young antler from Ballybetagh, Ireland (Fig. 9). However, some minor details of antler morphology associate the giant deer from Romania to ruffii/germaniae type: the anterior tine is not included in the palmation and remains separated from the palmed part of antler by a certain space, as in the type specimen of M. giganteus ruffii, and in the larger antler from Ural Region, Yekaterinburg (Vislobokova, 2012a,b: p. 710, Fig. 40); the antler palmation in the specimen from Romania is somewhat broader than in the antler from Ireland, and therefore the shape antler under discussion shows an intermediate condition between M. giganteus giganteus and M. giganteus ruffii. Both types of antler morphology of giant deer are found on the territory of Romania (unfortunately, the exact age of the finds is not known): a complete antler resembling M. giganteus giganteus (figured by Apostol, 1976: Fig. 28) and an antlered skull with compact antler crown as in M. giganteus ruffii (Apostol, 1976: Fig. 30), both discovered in Southern Romania.

Table 5 Measurements of giant deer antlers P-0002 (GMCU) from Hadikfalva (¼Dornes¸ti, Romania) and AM 70/552 (AMIA) from Mezyn (Ukraine). Measurements

P-0002

Total length of antler Circumference of antler base Height of basal ramification Height of middle tine ramification Distance between first and second ramifications Height of posterior tine ramification above burr Breadth of palmation above posterior tine Maximal breadth of palmation Length of palmation from posterior tine Antero-posterior diameter above burr Antero-posterior diameter of burr Antero-posterior diameter of pedicle Latero-medial diameter above burr Latero-medial diameter of burr Latero-medial diameter of pedicle Height of pedicle

1210.0 232.0 73.0 325.0 250.0 580.0 185.0 290.0 540.0 112.0 116.8 95.9 90.5 106.3 87.2 49.2

AM 70/552

36.5

98.1 100.6 76.3 97.9

3.10. Bukovynka Cave (Chernovtsy district, Ukraine)

Fig. 9. Young antlers of giant deer: A, M. giganteus ssp. from Hadikfalva (¼Dornes¸ti), Romania; B, M. giganteus giganteus from Dublin, Ireland (adapted from Reynolds, 1929); C, M. giganteus ruffii from Kottbus, Germany (type specimen, adapted from Nehring, 1891).

Bukovynka Cave (Upper Pleistocene, MIS 3) is located in an inactive gypsum quarry, not far from Stalinesti Village, on the left slope of the Matka River valley, a left tributary of the Prut River. It was discovered in 1976. In 2000, M. Vremir and B. Ridush excavated the deposits, finding animal remains, attributed to a cave hyena den (Vremir et al., 2000). The deposit is composed of loams (Ridush and Kuprich, 2003; Ridush, 2004; Ridush and Levytska, 2005; Korzun and Ridush, 2011). The giant deer was represented by a partly preserved mandible. Cave hyena remains from the same stratigraphical unit were 14C dated to 41,300 (þ1300/1100) BP (VERA2529). The left hemimandible PB-45 (GMCU) belongs to an old male individual (Table 2, Fig. 10). This specimen is interesting: it is very large (larger than any specimen from the sample from Ireland included in this study), characterized by very long lower cheek tooth row and is extremely robust, exceeding the thickness of lower mandibles from Ireland stored in the Natural History Museum of London. The relatively short premolar series of the specimen from Bukovinka Cave is another interesting peculiarity that distinguishes this mandible from the giant deer forms of Eastern Europe and approaches the giant deer from Ireland,

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Fig. 10. The left hemimandible PB-45 of Megaloceros giganteus from Bukovynka Cave: A, occlusion view; B, lateral view; C, occlusion view of P4.

representing an advanced evolutionary condition of this character (Fig. 4). 3.11. Burshtyn (IvanoeFrankovsk region, Ukraine) The site (49150 N, 24 390 E) (Middle Pleistocene, Dnieper, MIS 7 e MIS 6) is situated near Burshtyn. Numerous faunal remains are associated with fluvioglacial sands in the bottom of the Gnyla Lypa River, a left tributary of the Dniester River (Ridush, 2012). Remains of giant deer are represented by a right damaged calcaneus BUR-04 (GMCU, the greatest breadth amounts to 49.0 mm) and a basal fragment of antler BUR-30 (GMCU, the antero-posterior diameter above burr amounts to 57.3 mm). 3.12. Mizyn (¼Mezin, Chernigiv region, Ukraine) The Late Palaeolithic site of Mizyn from northern Ukraine (51480 4100 N 33 401200 E) has yielded a basal fragment of shed antler (AN 70/552, AMIA) of giant deer, which is the only specimen from Mizyn that was ascribed to this species (Fig. 11, Table 5). This is a recent finding which undoubtedly belongs to the giant deer. The antler beam is broken above the basal tine, which is situated close to the burr. The basal tine is flat in its distal portion, slightly undulated and terminated with two small rounded paddles. It is assumed that the antler fragment was used as a hammer for a drum-like instrument made of mammoth bones (Nuzhnyy, 2008). Radiocarbon dates for the site from mammoth teeth are 15,100  200 BP (OxA-719) (Svezhentsev, 1993: 26) and 14,560  90 BP (GrA-22499) (Nuzhnyy, 2008). The finding of a giant deer antler from Mizyn in the context of the obtained radiocarbon dates for the associated mammal remains needs special attention, as the available data do not fit with the absence of giant deer in the Paleontological record of Western and Central Europe during the 20e12.5 ka time span reported by Stuart et al. (2004). Possibly, Mizyn marks the western border of area of distribution of M. giganteus during the late phase of Last (Vistulian) glaciation. 4. Discussion 4.1. Giant deer from Eastern Europe At least two forms of giant deer were present in Eastern Europe during the geological period covered by the material included in

Fig. 11. The basal fragment of shed antler AM 70/552 of Megaloceros giganteus from Mizyn: A, upper view; B, basal view; C, lateral view.

this study. The giant deer found in the Bisnik cave is of great interest. The set of characters of the fossil material from Bisnik suggest a comparatively small primitive form of giant deer with relatively long premolar series, weak cingula of upper molars, and rather short massive metapodials. Its relatively narrower epiphyses of limb bones could be correlated with smaller body size. M. giganteus from Bisnik is smaller than the early form of this species from Holsteinian of Steinheim (ca. 400 or 300 ka), which is reported by Lister (1994) as a giant deer of “normal” body size (as large as the sample from Ireland), with possibly elongate distal limb elements. The metacarpal bone from Bisnik (which presumably belongs to a female) is particularly short and has a rather equivocal position compared to the data of Van der Made (2006), who distinguished in the sample from the Rhine basin and North Sea two morphological forms of giant deer: the older Holsteinian/ Hoxnian and Saalian/Wolstonian M. giganteus “antecedents/germaniae” with slender metacarpals, and younger Eemian M. giganteus ssp. with robust metacarpals (Fig. 5). The metacarpal from Bisnik may be regarded as slender in distal breadth; however its relative mid-shaft breadth is rather significant. Although rather short, the metacarpus from Bisnik is quite close in its dimensions and proportions to the metacarpal from Kottbus (Germany) found together with the holotype of M. giganteus ruffii (regarded by van der Made, 2006; Vislobokova, 2012a; as a synonym of germaniae), but also is close to the sample of giant deer from Zhana-Aul (Kazakhstan) described by Kozhamkulova (1969). Generally, the material from Bisnik, Kottbus, and Zhana-Aul fall within the range of Van der Made’s “slender” type of metapodials. Some other postcranial elements of the giant deer from Bisnik Cave (talus B/04/W/3764, radius Nr. 1703) are quite small as well, and possibly belong to the same small-sized individual. The giant deer from Bisnik shares smaller dentition size with the fossil form from Grigorievka (Northern Kazahstan) described by Shpansky (2011) as M. giganteus ruffii. Lister (1994) reported a rather small-sized form of giant deer from “stage 7” interglacial of Crayford (Kent, England) as a local

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stunted population that evolved in a wooded landscape. The relationship of the giant deer from Bisnik Cave and the stunted M. giganteus from Crayford is not clear at present. The primitive condition of dental proportions and cranial morphology is another noteworthy peculiarity of the giant deer from Bisnik. M. giganteus from Bisnik is characterized by rather short nasal bones that do not reach the imaginary line connecting the anterior edges of orbits. The taxonomic significance of the morphology of nasal bones needs discussion, since various authors assigned different significance to this characteristic. Lydekker (1915) regarded the shape and relative length of nasal bones as a diagnostic character of some subspecies of modern muntiacines. Flerov (1952), Sokolov (1959), and Baryshnikov and Tikhonov (1999) regarded the numerical and relative length of nasals, the position of their posterior edge, and their shape as diagnostic characters at genus and subgenus level in Cervidae. Sokolov (1953) used the relative length of nasal bones (as a ratio to the basal length of skull) in characterization of subfamilies of Bovidae. Vislobokova (1990) regarded the relative length of nasale as a diagnostic character of cervid subfamilies. According to Sokolov (1953: 19), the length of nasal bones results from the morphogenesis of the anterior part of the skull. Flerov (1952) applied the ratio between length of nasal bones and length of upper tooth row (one of the most conservative characters in individual variation and evolutionary change) in order to describe the relative length of nasale. The position of posterior edge of nasal bones in cervids mostly depends on the relative length of nasale and relative length of facial or orbito-frontal parts of skull. The relative length of nasal bones is influenced by sexual dimorphism, being longer in males characterized by longer facial portion of skull, and by the interspecific geographical variation, being shorter in primitive and endemic insular forms with relatively shorter facial part of skull (for instance, in red deer, the subspecies Cervus elaphus barbaricus and Cervus elaphus corsicanus respectively). However, in all those cases, the position of fronto-nasal suture varies very little (Croitor, unpublished data). Generally, a more anterior position of nasofrontal suture in cervids is regarded as primitive (plesiomorphic) character (Flerov, 1952). This character is advanced in the giant deer from Ireland, characterized by long nasal bones reaching far behind the line connecting anterior edges of orbits even in female skulls, for instance in the skull M2326 stored in NHML (Croitor, 2006). In modern fallow deer Dama dama, which represents the closest living relative of the giant deer (Lister et al., 2005), the skull is relatively short, also its facial part is comparatively short, and therefore the nasal bones reach behind the line of anterior ridges of orbits, which, in their turn, are relatively large (Flerov, 1952). The long nasalia in Irish giant deer and modern fallow deer evolved independently in parallel, and cannot be used as an argument for their close phylogenetic relationship, as the nasalia were short in the primitive form of Megaloceros discovered in Bisnik. Unfortunately, this character remains unknown in M. giganteus antecedens and M. giganteus ruffii, so we cannot make a conclusion on its taxonomical significance. The giant deer from Bisnik also is characterized by the relatively very long lower premolar series, compared to samples and specimens involved in the present study. The only specimens that have similarly long premolars as in the sample from Bisnik are the lower mandible from Buco del’Orso (Lombardia, Italy) reported by Anfossi et al. (2002), the mandible from Grigorievka (North Kazahstan), the lower mandible of mounted female skeleton B-VI-80 from the Paleontological Gallery in Paris, and the male specimen from Limerick (Ireland) stored in the Museum of Nature in Wroclaw (Fig. 4). However, the specimens from Buco del’Orso and Limerick are larger than the material from Bisnik Cave, while the specimen stored in Paris is very close to the sample under consideration.

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Lower mandibles from Duruitoarea Veche and Romankovo (Ukraine) are also similar to the sample from Bisnik in size and tooth series proportions, occupying a transitional position between the giant deer from Bisnik and the main part of the sample from Ireland with the short premolar series. The sample of giant deer from Ireland stored in various museums of Europe, apparently, does not represent a single population. Besides the broad variation in tooth series proportions, the material from Ireland shows also a significant and meaningful split in metacarpal length and robustness. The metacarpals of mounted skeletons of the female exposed in the Paleontological Gallery of Paris, the male from the Natural History Museum “Grigore Antipa” of Bucharest, the male from Athlone (Ireland) and the male from unidentified Irish site exposed in the Museum of Nature of Wroclaw, and the skeleton from Enniscorthy (Ireland; stored in the Sedgwick Museum of Cambridge) described by Reynolds (1929) are short and relatively robust, overlapping with the sample from Bruine Bank (Fig. 5). The metacarpals of the mounted male skeletons from the Paleontological Gallery of Paris and the Paleontological Institute of Munich are particularly long, approaching the giant deer from Schlutup (Germany) and Sapozhok (Russia) (Fig. 5). Apparently, this group of specimens corresponds to van der Made’s Late Devensian group of “intermediate type of metapodials”. Regarding the relative length of upper premolar series, the sample from Ireland is divided in two groups: the group of specimens with relatively long premolar series that includes the main part of sample from Rhine basin, North Kazakhstan, and the specimen from Brînzeni-1, and the group with relatively short premolar series that may be approached to the larger specimen from Sapozhok (Fig. 12). It is difficult to affirm the correlation between limb proportions and tooth series proportions taking in consideration only the mounted skeletons of giant deer from Ireland, because those showpieces originally were subjects of commercial sale and it is not known if the skeletons are genuinely articulated, or (what is more probable) compiled. Moreover, according to our observations, it is very difficult to define a “typical Irish” subspecies M. giganteus giganteus, as the material from Ireland mentioned by includes both “robust” and “long-limbed” types of giant deer. The majority of findings from Eastern Europe are characterized by “normal” larger body size. The studied fossil material from Eastern Europe, including the oldest remains of giant deer from Ofatint¸i (110e125 ka) and the latest findings from Brînzeni (20e 26 ka BP, possibly one of the latest presences of giant deer in the region), are characterized by rather similar body size. It seems that the giant deer from Krosinko is a rather large individual, which is close to the largest specimens of the sample from Ireland. According to the data from Duruitoarea Veche and Brînzeni, as well as the literature data on giant deer from Romankovo, Ukraine (Svistun, 1968), and Grigorievka, Northern Kazahstan

Fig. 12. Comparison of lengths of upper tooth series of Megaloceros giganteus from various sites of Eurasia.

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(Kozhamkulova, 1969), Late Pleistocene giant deer from Eastern Europe and Northern Kazahstan possessed relatively long lower premolar series. The form of giant deer represented by the complete metatarsal bone from Brînzeni definitely was rather shortlimbed. Therefore, we can assume that some fossiliferous sites from Poland, Moldova and Ukraine yielded a large short-limbed form of giant deer with relatively long premolar series that distinguish this Eastern European form of giant deer from the latest survivor of this species in Europe, from the Late Devensian of Ireland. The particularly large and robust lower mandible from Bukovynka with very short lower premolar series represents a rather exceptional large and advanced (in its dental proportions) form of giant deer, resembling in this characteristic the main sample of Irish giant deer. The problem of the specimen from Bukovynka requires some comments on the articulated skeleton of giant deer from Sapozhok (Ryazan, Russia) stored in the Paleontological Institute, Moscow (collection number PIN-337). The specimen was originally described by Pavlova (1929), some additional measurements were published by Svistun (1968) and Vislobokova (2012a), and its dentition was figured by Flerov (1962). This is a rather large form characterized by long and little curved antlers with four crown tines situated on the distal and anterior sides of palmation as in M. giganteus giganteus from Ireland, but also resembling the antler of giant deer from Romania figured by Apostol (1976). The middle tine is coalesced with distal palmation of the antler. The angle between antler beam and palmed portion of antler is straight. The typical “Irish” morphology of antlers was regarded as a decisive argument for ascribing the giant deer from Ryazan to M. giganteus giganteus (Vislobokova, 2012b). The cheek teeth of giant deer from Sapozhok figured by Flerov (1962) are characterized by particularly short premolar series, which is shorter than in the majority of specimens from Ireland, but is similar to the Irish specimens with the shortest premolars. Possibly, the giant deer from Bukovinka is similar to Megaloceros from Sapozhok. The giant deer from Sapozhok is also characterized by particularly long metapodials, which relate this deer to the most “long-limbed” giant deer from Western Europe. 4.2. Possible taphonomic biases The mandibular pachyostosis of giant deer is a constant character through time, as the earlier jaws, including the specimens reported in the present study, are as wide for their size as those from Ireland. The mandibles of females are relatively narrower than in males, and therefore the mandibular pachyostosis is exposed to a strong sexual dimorphism (Lister, 1994; Croitor, 2006). Lister (1994) regards the jaw thickening in M. giganteus as a part of a generally enhanced skeletal calcium store, related to the unusually large size of the antlers. According to the mandible thickness, females are more frequent in the samples from the Bisnik Cave and Moldavian grottos, unlike the “reference” sample from Ireland. Such a specific demographic composition of the samples from our study makes comparison with the sample from Ireland dominated by males difficult. Therefore, the results of morphological comparison should be regarded with a certain caution (Croitor, 2006). According to Barnosky (1985), the sex ratio of the sample is altered by the selective entrapping of the males in bogs, but also by strong collection bias. The remains of giant deer from caves and grottos had been accumulated by prehistoric hunters and thus must be affected by a different type of taphonomic bias. Selective hunting by Palaeolithic humans was the main factor influencing the sex ratio of the fossil remains of game mammals. The prey selection of collective hunters to a greater extent depended on the body mass of the prey species (a larger prey is a

difficult prey) and human resources available (more hunters can have more success in getting difficult prey). According to Croitor and Covalenco (2011), Palaeolithic hunters were less selective while hunting smaller herbivores, such as reindeer, and therefore their remains in many cases reflect the demographic structure of natural populations. Selective hunting became more pronounced with the increase of body mass of the hunted mammals. In this case the preference of hunters is often focussed upon females, juveniles, and seniles, which are regarded as more vulnerable and easier prey (Croitor and Covalenco, 2011). Apparently, more frequent presence of females in the samples on giant deer from caves and grottos accumulated by Palaeolithic hunters represents the result of specific prey selection. Sex ratio biases of fossil samples of giant deer may also influence the general picture of morphological characters of a sample. Apparently, the strongly ossified vomer, reduced size of foramen ovale, diminished or even completely closed ethmoidal openings, strong obliteration of cranial sutures, and even development of cingula in upper molars may be caused by pachyostosis (Croitor, 2008). However, this question is very little studied. Cingulum in upper molars is regarded by many authors as a characteristic for Megaloceros morphological trait, but the nature of cingulum remains unclear. The internal cingulum on the upper molars is often considered a diagnostic characteristic of the Tragulidae, but also is retained in primitive pecoran genera such as Gelocus, Eumeryx, and Prodremotherium, and traces of the internal cingulum are retained in the extinct higher pecoran genera Dremotherium, Palaeomeryx, and Dicrocerus (Janis and Scott, 1987). Cingulum is also recorded in few Cervines, including Metacervocerus pardinensis, and its presence is assumed in Arvernoceros ardei (Heinz, 1970). M. giganteus is one of few exceptional examples of cervines with lingual cingulum in upper molars. However, this characteristic is very variable in giant deer and its taxonomical importance is questioned by some authors (Azzaroli and Mazza, 1993). The cingulum is welldeveloped in the sample of giant deer from Ireland, but it is very variable: normally, cingulum is present in upper molars and shows a different degree of expression. In some cases, cingulum is developed in upper premolars and even in lower molars (Croitor, 2008). However, finds from Eastern Europe often show very weak development of cingulum. The cingulum is quite weak in the male skull Nr. 34e1022 from Romankovo (Svistun, 1968: 29, Fig. 15) ascribed to M. giganteus ruffii, where it is clearly present (but still very weak) only on M3. The male individual from Sapozhok has a somewhat stronger cingulum on upper molars: it is well-developed on M3 and clearly visible on M2, although it is not so strong, and is virtually missing on M1 (Flerov, 1962: 376, Fig. 496). Cingulum is quite weak in specimens found in Duruitoarea Veche, Brînzeni, and Bisnik. Possibly, the cingulum represents a sort of dental hypertrophy related to the hyperostosis of the cranium in giant deer (Croitor, 2008). 5. Conclusions Regarding the results of our comparative biometric study, it is difficult to assert how much the results of our research are affected by the taphonomic biases, as the fossil material from each fossiliferous site is not numerous. However, we cannot assume the anthropogenic nature of all sites involved in the present study. Morphological and biometric study revealed the presence in the Bisnik Cave (ca. 70e60 ka; MIS 4 and MIS 3) a comparatively smaller form of giant deer with some primitive cranial characters (short nasal bones) relatively long premolar series, weak cingulum in upper molars, and comparatively short metacarpal. It is difficult to classify the shape of metacarpal from Bisnik as robust or slender, as it is characterized by relatively narrow distal epiphysis and rather

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massive shaft. However, there is a certain similarity of the specimen from Bisnik with the metacarpal from Kottbus found together with the holotype of M. giganteus ruffii, and with a sample of giant deer from North Kazakhstan. Nonetheless, the majority of remains of giant deer from Eastern Europe are characterized by “normal” larger body size. The larger, but still primitive in its weak development of cingulum in upper molars and relatively long lower premoral series specimen of giant deer is recorded in the older deposits of Ofatint¸i (120e130 ka) and a short-limbed type of giant deer with similar morphology of dentition is recorded in the younger site of Brînzeni1 (20e26 ka BP). Perhaps, the majority of findings from Poland, Ukraine, and Moldova could be ascribed to this form. This form of giant deer is characterized by a normal for Megaloceros mandibular pachyostosis, relatively long upper and lower premolar series, very weak cingulum in upper molars, and, apparently, represents the “robust type” of Lister (1994) and Van der Made (2006). Another exceptionally large form of giant deer is found in Bukovynka Cave (Ukraine), but possibly some remains from Russia, such as the well-known articulated skeleton from Sapozhok, also belong to this form. Besides its larger size, the giant deer from Sapozhok is characterized by relatively short premolar series and well developed cingulum, as the assumed “classical” M. giganteus from Ireland. The giant deer from Sapozhok, as well as the specimen from Schlutup and a part of the sample from Ireland, are characterized by comparatively long metacarpal bones and represent the so-called group of “intermediate metacarpal robustness” indicated earlier by Lister (1994) and Van der Made (2006). The presence of large long-limbed giant deer with advanced dentition in Eastern Europe supports the hypothesis of Van der Made (2006) on immigration of “classical Irish” form of M. giganteus from the east around 13 ka. According to the new finding from Mizyn (Ukraine), the presence of giant deer in Eastern Europe, unlike the paleontological evidence from Central and Western Europe, was continuous. Therefore, Eastern Ukraine and European Russia may be regarded as a “foothold” for the Devensian recolonization of northern Western Europe by the long-limbed giant deer with advanced proportions of dentition. Acknowledgments Radiocarbon dating from Krosinko, Wiercica Cave and Barycz sites was supported by a grant from the Polish Ministry of Science and Higher Education (N N307 050139; grant manager Kamilla Paw1owska). We gratefully acknowledge Tatiana WoroncowaMarcinowska (Geological Museum of the Polish Geological Institute, National Research Institute (PIG-PIB) in Warsaw), Professor Jerzy Nawrocki (Director of the Polish Geological Institute, National Research Institute (PIG-PIB) in Warsaw) and Waldemar Wia˛ zek for providing to one of us (Kamilla Paw1owska) the study materials from Wiercica Cave and Krosinko, respectively. Roman Croitor thanks Professor Dumitru Murariu (Director of the National Museum of Natural History “Grigore Antipa”), Gertrud Rössner (Paleontological Museum of Munich), and Christine Argot (National Museum of Natural History of Paris) for access to the giant deer remains stored in the collections under their care. We give deep thanks to Adrian Lister and Jan van der Made for their valuable comments that helped to improve our article. We also thank our colleagues Andrey Shpansky, S¸tefan Vasile and Stephen Poropat for supplying us with bibliographic sources. References Aaris-Sørensen, K., Liljegren, R., 2004. Late Pleistocene remains of giant deer (Megaloceros giganteus Blumenbach) in Scandinavia: chronology and environment. Boreas 33 (1), 61e73.

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Alekseeva, E.V., 1980. Pleistocene Mammals of South-east of Western Siberia (Carnivores, Proboscideans, and Hoofed Mammals). Nauka Publishing House, Moscow, p. 188 (in Russian). Anfossi, G., Rossi, M., Santi, G., 2002. Un’emimandibola destra di Megaloceros giganteus (Blumenbach, 1803) dalla grotta del Buco dell’Orso (Laglio e Como, Lombardia). PIANURA e Scienze e storia dell’ambiente padano 15, 5e13. Anisiutkin, N.K., 2001. Mousterian Age of South-west of Russian Plane. Evropeiskiy Dom Publishing House, Saint-Petersburg, p. 308 (in Russian). Apostol, L., 1976. L’étude morphométrique des mammifères fossiles quaternaires de la plaine Roumaine et leur distribution paléozoogéographique. In: Travaux du Muséum d’histoire naturelle “Grigore Antipa”, vol. 17, pp. 341e375. Azzaroli, A., 1953. The deer of the Weybourn Crag and Forest Bed of Norfolk. Bulletin of the British Museum (Natural History), Geology 2 (1), 3e96. Azzaroli, A., Mazza, P., 1993. 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REkonstrukcja Zasiedlenia Jaskini Na Tle Zmian   , pp. 9e231 (in Polish Srodowiska Przyrodniczego. Wydawnictwo UMK, Torun with English summary). Cyrek, K., Socha, P., Stefaniak, K., Madeyska, T., Miros1aw-Grabowska, J., Sudo1, M., _ Czyzewski, q., 2010b. Palaeolithic of Bisnik Cave (Southern Poland) within the environmental background. Quaternary International 220 (1e2), 5e30. David, A.I., 1980. Pleistocene Theriofauna of Moldova. S¸tiint¸a Publishing House, Kishinau, p. 185 (in Russian). David, A.I., Chetraru, N.A., 1970. Mammal fauna of Paleolithic age of Moldova. In: Fauna of Cenozoic of Moldova. S¸tiint¸a Publishing House, Kishinau, pp. 14e15 (in Russian). Driesh, A. fon den, 1976. A Guide to the Measurements to Animal Bones from Archaeological Sites. Peabody Museum Bulletin (Harward University), pp. 1e137. Flerov, K.K., 1952. Musk deer and deer. In: Novikov, G.A. (Ed.), The Fauna of USSR. Mammals, vol. 1 (2), pp. 1e256. Moscow-Leningrad (in Russian). Flerov, K.K., 1962. 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Please cite this article in press as: Croitor, R., et al., Giant deer Megaloceros giganteus Blumenbach, 1799 (Cervidae, Mammalia) from Palaeolithic of Eastern Europe, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.10.068