Socrates J. Roussiakis & George E.Theodorou National University of Athens

Carnivora from the Late of Kerassiá (Northern Euboea, Greece)

Roussiakis, S.J. & Theodorou, G.E., 2003 – from the Late Miocene of Kerassiá (Northern Euboea, Greece) – in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRA- TION OF TERTIARY IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 469-497 [ISSN 0923-9308] Published 1 December 2003

Excavations carried out in recent years in Kerassiá brought to light many fossiliferous sites. In this study, the carnivores from the three richest sites, Kerassiá 1 (K1), Kerassiá 3 (K3) and Kerassiá 4 (K4), are discussed. K1 includes Metailurus cf. parvulus and possibly Adcrocuta eximia; K3 inclu- des Plioviverrops sp. and cf. Ictitherium pannonicum; and K4 includes Adcrocuta eximia and giganteus. The carnivores as well as the other faunal remains from these localities indicate an early to middle Turolian age.

Correspondence: S.J. Roussiakis and G.E. Theodorou, University of Athens, Department of Historical Geology and Palaeontology, Panepistimiopolis, 157 84, Athens, Greece; e-mail: [email protected]; [email protected]

Keywords: Greece, Euboea, Late Miocene, Turolian, Carnivora

INTRODUCTION and Palaeontology) started systematic exca- The existence of Miocene vertebrates vations in the area. These excavations, finan- in Euboea has been known since 1878 ced by the University of Athens, the (Cordella 1878). Further information is provi- Municipality of Nileas and the General ded in Woodward (1901), Déprat (1904), Secretary of Research and Technology Mitzopoulos (1947), Melentis (1968, 1969), (Project 95 ∑YN107) brought to light more Jacobi (1982), and others. These authors pre- than seven fossiliferous sites. The fauna of sent information on the Limni, Hagia Anna, Kerassiá has been described until now by Rhovies, Achladi, Eria, Prokopi, Palaeovrissi Köhler (1983), Van der Made & Moyà-Solà and Halmyropotamos localities. The presence (1989), Theodorou et al. (1995, 1998, this of in Kerassiá has been known to local volume) and Kostopoulos et al. (2001). people since 1966, when fossil bones were Köhler (1983) mentions the presence of found during the construction of a road close Microstonyx sp., Bovidae, Giraffidae, two to the village. In 1981, Köhler tracked down Hipparion , two species of Probo- the fossil bones of Kerassiá during geological scidea and a large carnivore. Van der Made & fieldwork in the area. The first excavations in Moyà-Solà (1989) briefly describe the suids Kerassiá were carried out in 1982 by Hans de and attribute these to Microstonyx major ery- Bruijn and Albert van der Meulen (University manthius. Moreover, they mention the pre- of Utrecht) and Constantin Doukas (Univer- sence of Deinotherium and Dorcatherium and sity of Athens). The excavations were not give a middle Turolian age for Kerassiá. resumed until 1992, when the University of According to Kostopoulos et al. (2001), the Athens (Department of Historical Geology Kerassiá Microstonyx major is smaller than

469 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

the Pikermi M. major erymanthius and proba- from the same ossiferous block and most pro- bly represents a new subspecies. bably from the same individual. The carnivores of Kerassiá are represented from a small number of specimens, and the Description and comparisons material is not always in a good state of pre- From Tables 1-3 it is clear that the available servation. In the present study, we discuss the postcranial material is similar in its dimen- carnivores of the three richest sites, the sites sions to that of Plioviverrops orbignyi Kerassiá 1 (K1), Kerassiá 3 (K3) and (GAUDRY & LARTET, 1856) from Pikermi, Kerassiá 4 (K4). The material of this study mentioned by Gaudry (1862, pl. 11, figs. 6, comes from the excavations carried out from 10) and Pilgrim (1931). The available fourth 1992 onwards and is temporarily stored in the metatarsals are only slightly larger than the Athens Museum of Palaeontology and specimen mentioned by Pilgrim (1931). In Geology. In the future, this material will be their morphological characters, the present exhibited in a new museum in Kerassiá that specimens also show great resemblance to P. is planned by the local prefecture. orbignyi. The distal articular surface of the calcaneum is flat and narrow as in P. orbignyi Methodology and abbreviations (Pilgrim 1931). The cuboid facet of the Mt The methodology used for the dental meas- IV is not extended posteriorly more than the urements follows Werdelin (1988a). A value tuberosity for the ligament of the peroneus inside parenthesis signifies an approximate longus, something also observed in P. orbig- measurement, "a" signifies a measurement nyi (Pilgrim 1931). The humerus is similar to taken at the alveolus and "r" signifies a meas- that of P. orbignyi from Pikermi, except in its urement taken at the root. The methodology smaller supratrochlear foramen. Otherwise, used for the postcranial material is given with the humerus has an entepicondylar foramen each table of measurements. AMPG: Athens accompanied by a strong bar, as in P. orbig- Museum of Palaeontology and Geology. nyi. MNHNP: Muséum National d’Histoire Except for P. orbignyi, the postcranial Naturelle, Paris. BMNH: The Natural History material of the Miocene small carnivores is Museum, London. poorly known. "Progenetta crassa" (=Protic- titherium crassum) from the Can Llobateres SYSTEMATICS II in Spain (Crusafont-Pairó & Petter 1969, fig. 5, pl. 3, fig. 7) is significantly larger from Family: Hyaenidae GRAY, 1869 the Kerassiá 3 material. Promeles palaeatti- Subfamily: Ictitheriinae TROUESSART, 1897 cus (WEITHOFER, 1888) has a smaller calca- : Plioviverrops KRETZOI, 1938 neum and metatarsals (Pilgrim 1931; Plioviverrops sp. Roussiakis 1996), and its fourth metatarsal is Locality: Kerassiá 3 (K3) shorter relative to the calcaneum. The ratio height calcaneum/length Mt IV is 58.3 in P. Material palaeatticus (data from Roussiakis 1996) but K3/B1/4: left humerus, lacking the proximal- 51.7-52.7 in the Kerassiá Plioviverrops. More- most end. K3/B1/15: distal epiphysis of a over, in P. palaeatticus the cuboid facet of the right femur, juvenile. K3/B1/18: distal epi- calcaneum is concave and oval, with its grea- physis of a left femur, juvenile. K3.192: right ter diameter obliquely oriented. In the Plio- calcaneum, damaged on its sustentaculum viverrops sp. of Kerassiá the cuboid facet is tali. K3.206: proximal epiphysis of a left almost flat and anteroposteriorly elongated, humerus, juvenile. K3.161: left Mt IV. as in P. orbignyi. K3.195: right Mt IV. Of the specimens listed The available postcranial material thus above, those with the indication B1 come shows dimensions similar to P. orbignyi from

470 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Figure 1 Logarithmic ratio diagram comparing the Kerassia 3 ictithere with various species of icitheres. H. wongii from Samos according to Werdelin (1988a), I. viverrinum from Pikermi according to Werdelin (1988b), I. pannonicum from Polgardi and the Black Sea from Table 4.

Pikermi. In most of their morphological char- from the late Turolian (MN13) of Brisighella acters, the Kerassiá specimens also agree in Italy and MN14 of La Gloria 4 in Spain with P. orbignyi. Nevertheless, at the present (Alcalá 1994). P. orbignyi, which shows time we prefer to refer the specimens consi- resemblance to our findings, is reported from dered here to Plioviverrops sp., since the lack the early-middle Turolian (MN11/12) of of more material, especially teeth, does not Ravin des Zouaves 5, Vathylakkos 2, 3, permit us a more precise determination. Prochoma 1 and Perivolaki (De Bonis & The genus Plioviverrops is considered to be Koufos 1991, Koufos et al. 1999, Koufos the oldest known hyaenid, appearing first in 2000), and from the middle Turolian (MN12) MN2a of Laugnac in France, with the species of Pikermi and Samos (Major 1888, Major Plioviverrops collectus (DE BONIS, 1973) 1894, De Beaumont 1969, Bernor et al. 1996) (Ginsburg 1999). Various other species of in Greece. It is interesting to notice that in Plioviverrops are known, such as Pliovi- Vathylakkos 3, P. orbignyi seems to have verrops gervaisi DE BEAUMONT & MEIN, coexisted with another form, listed as P. cf. 1972, from MN7/8 of La Grive (France), guerini by De Bonis & Koufos (1991) and Plioviverrops guerini (VILLALTA & CRUSAFONT, Koufos (2000). 1948) from MN11 of Piera and Viverro de Pinos and MN12 of Cerro de la Garita and Genus: Ictitherium ROTH & WAGNER, 1854 Los Mansuetos in Spain (Ginsburg 1999), cf. Ictitherium pannonicum KRETZOI, 1952 and Plioviverrops faventinus TORRE, 1989 Locality: Kerassiá 3 (K3)

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Material SEMENOV, 1989. K3/204: right mandibular ramus with ci, p3, Ictitherium viverrinum ROTH & WAGNER, p4 and m1. The alveoli of p1, p2 and m2 are 1854 is a well-known species with a wide visible. geographical range. Compared to I. viverri- num from Pikermi, K3.204 has clearly larger Description p3, m1 and m2 (Fig. 1). Thalassictis robusta The available mandible is badly damaged and GERVAIS EX NORDMANN, 1850, from Kishinev, some of its characters cannot be evaluated is also smaller than the Kerassiá ictithere precisely. The mandibular corpus has a men- (Fig. 2). Hyaenictitherium hyaenoides tal foramen below the middle of the alveolus (ZDANSKY, 1924) has larger teeth than the for p2. The lower border of the mandibular Kerassiá specimen and a small m2 (Fig. 3). corpus slopes down from the symphysis to a Moreover, the mandible of that species has a point under m1. The depth of the mandible more "hyaenoid" appearance. It is robust, thus increases from the anterior to the poste- straight, and with a lower border that is rior part and the depth in front of p2 is smal- almost parallel to the alveolar border (Qiu ler than that behind m1 (Table 4). The total 1985). The mandibular corpus of K3.204, on length of the symphysis measures about 40 the other hand, is deeper in its posterior part mm and its posterior border lies under the (Table 4). Palinhyaena reperta QIU, HUANG middle of the alveolus for p2. The alveolus & GUO, 1979, is known from Chinese locali- for p1 is separated from the canine by a dias- ties only (Werdelin & Solounias 1991). This tema of 9.9 mm. The p3 has a posterior species is also larger than the Kerassiá 3 ictit- accessory cusp and most probably a smaller here, but with a smaller m2 (Fig. 3). The anterior one. The p4 is more elongated than mandible of P. reperta is similar to that of H. p3 (Table 4) and has a robust anterior acces- hyaenoides, i.e., ‘hyaenoid’ in appearance. sory cusp. The posterior part of p4 is not very The lower premolars of K3.204 are slightly well preserved but almost certainly had an more robust than those of Hyaenotherium accessory cusp. The most important character wongii (ZDANSKY, 1924) from Samos (Fig. of m1 is the presence of a strong metaconid. 3). This is especially true for p3, while p4 is The protoconid and paraconid are almost badly damaged and its width is only approxi- equal in height. The talonid of m1 is large mately given (Table 4). The most important and wide, but very worn and its morphologi- difference concerns m2. This is not pre- cal details are not visible. The only preserved served in the Kerassiá specimen, but judging cusp is the entoconid, which is high. The m2 from its elongated alveolus that measures is not preserved, but judging from its elonga- about 8.9 mm, it was especially large (Table ted alveolus it was large. 4, Fig. 3). Such a character differentiates our specimen from H. wongii and makes it more Comparisons similar to Ictitherium pannonicum KRETZOI, The group of the ‘ictitheres’ contains many 1952, a species that was first described from species and genera that share numerous char- Polgardi in Hungary, but it was actually igno- acters and sometimes have slight differences red until Semenov (1985, 1989) restudied that make their identification difficult. In this these specimens and attributed some more study, we follow the revision of Werdelin & specimens from the Black Sea area to it. Solounias (1991). The various species of the Werdelin & Solounias (1991), however, note ‘ictitheres’ have been referred mainly to the that we cannot be certain if the Black Sea genera Ictitherium WAGNER, 1848, Thalassictis specimens belong to the same species as the GERVAIS EX NORDMANN, 1850, Hyaenicti- Polgardi specimens. I. pannonicum is slightly therium KRETZOI, 1938, Palinhyaena QIU, larger overall than H. wongii, but in some of HUANG & GUO, 1979, and Hyaenotherium its dimensions is within the range of H. won-

472 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Figure 2 Logarithmic ratio diagram comparing the Kerassia 3 ictithere with various species of icitheres. H. wongii from Samos according to Werdelin (1988a), T. robusta from Kishinev according to Kurtén (1982), I. pannonicum from Polgardi and the Black Sea from Table 4. gii (Fig. 3). The most important distinguis- p4 is larger. Moreover, that specimen lacks hing character of I. pannonicum is the large any teeth posterior to p4. Another species m2 (Figs. 2, 3). As mentioned earlier, such a possibly related to I. pannonicum, is character is also present in our specimen. Ictitherium intuberculatum OZANSOY, 1965, Aside from the large m2, the Kerassiá 3 spe- from the Yassiören in Turkey. Werdelin & cimen resembles I. pannonicum in its m1 but Solounias (1991) emphasize the possibility it has more robust p3 (Table 4, Fig. 2). that this form might be conspecific with I. Unfortunately, the species I. pannonicum is pannonicum, but Ozansoy (1965) does not poorly known. In addition to Polgardi give metric data for the lower dentition and (Kretzoi 1952), which is the type locality, comparison with the Kerassiá ictithere is not specimens referred to that species are also possible. known from Chobruchi in Moldova, and Of the localities mentioned, Chobruchi is Novaja Emetovka 2 and Cherevichnoe in considered to be early Turolian (MN11) Ukraine (Semenov 1985, 1989). A specimen (Semenov 1989), Valdecebro 5, Novaja from Valdecebro 5 in Spain referred to Emetovka 2 and Cherevichnoe middle Thalassictis aff. hipparionum by Adrover et Turolian (MN12) (Adrover et al. 1986, al. (1986), may, according to Werdelin & Semenov 1989) and Polgardi late Turolian Solounias (1991), also belong to I. pannoni- (MN13) (Ginsburg 1999). cum. Alcalá (1994) further mentioned a frag- mentary mandible from the middle Turolian Subfamily: Hyaeninae MIVART, 1882 (MN12) of Cerro de la Garita in Spain under Genus: Adcrocuta KRETZOI, 1938 the name ‘Ictitherium aff. I. pannonicum’. Adcrocuta eximia (ROTH & WAGNER, 1854) The p2 and p3 of that specimen are compara- Locality: Kerassiá 4 (K4) ble in size to those of I. pannonicum, but the

473 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Figure 3 Logarithmic ratio diagram comparing the Kerassia 3 ictithere with various species of icitheres. H. wongii from Samos, P. reperta and H. hyaenoides from China according to Werdelin (1988a), I. pannonicum from Polgardi and the Black Sea from Table 4. 4.

Material posterior half of P3. The maxillopalatine K4.7: fragment with the right P3 and suture is not clearly visible, but most proba- P4. The P3 is badly damaged and the alveo- bly its anterior limit lies opposite the posteri- lus of M1 is visible. K4/∆388/1: skull frag- or part of P3. The anterior palatine foramina ment with the left I1-P3, Cs, P1-P4 and M1, are situated level with the middle of P2. The and the right I1-I3, Cs, and P3-P4. K4/∆388/2: posterior palatine foramina are not visible. left mandibular ramus with the complete den- The I1 and I2 are arranged in an almost tition (i1-i3, ci, p1-p4 and m1). K4/∆388/3: straight line, while I3 is situated slightly pos- right mandibular ramus with the complete teriorly. The I2 is only slightly larger than I1, dentition except p1. The lower mandibles but I3 is significant larger. The I1 and I2 have K4/∆388/2-3 were found connected at their a distal cingulum composed from two cusps, symphysis, and so come from the same indi- with the lingual one slightly larger than the vidual. The specimen K4/∆388/1 was found lateral one. The upper canine has a mesiolin- next to these mandibles, and it is almost cer- gual and a distal crest. The P2 is not well pre- tain that all these specimens belong to the served, but shows a posterior accessory cusp. same individual. This is also indicated from On its anterior part, we cannot clearly obser- their ontogenetic stage, since the teeth are ve an accessory cusp, but only a small bulge almost unworn and neither the upper nor the at the lingual side of the base of the anterior lower canines have fully erupted. crest. The P3 has a posterior accessory cusp and a small anterior one in a mesiolingual Description of the skull and the position. The greatest width is at the anterior upper dentition part of the tooth. The P4 is characterized by a The infraorbital foramen opens above the small protocone, a parastyle that is smaller

474 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

and lower than the paracone, and a metastyle Pikermi. Later it has been found at many fos- that is slightly longer than the paracone silliferous sites in Europe and Asia. The (Table 5). The distal part of the metastyle generic attribution of the species has changed blade is directed labially. M1 is small and its several times, and it is now attributed in metacone projects slightly posteriorly. Adcrocuta KRETZOI, 1938, opinion followed also here. Description of the lower mandible The available specimens from Kerassiá and the lower dentition have all the morphological characters of A. The mandibular corpus has two mental fora- eximia. This species shows some variation in mina. The anterior foramen opens under the the position and the number of the mental middle of p2 and the posterior one under the foramina (Gaudry 1863, Werdelin & contact between p2 and p3. The symphysis Solounias 1991, Koufos 2000). The specimen measures about 52 mm in length and extends figured by Roth & Wagner (1854, pl. 8, fig. posteriorly to the anterior part of p2. The 6) has two mental foramina, almost in the anterior border of the masseteric fossa is situ- same positions as the Kerassiá specimens. ated posterior to m1. The incisors increase in Gaudry (1863) mentioned that of three speci- size from i1 to i3 and they have a lateral mens available to him, two had two mental cusp, which shows an increase in size from i1 foramina, but another specimen had only one to i3. The lower canine has a mesiolingual mental foramen. The same variation is also and a distal crest and is slightly more curved present in the Axios material. The specimen than the upper canine. The cheek teeth are RZO-126 has two mental foramina, while the arranged in a curved line. The posterior part specimen RZ1-4 has one (Koufos 2000). of p3 slightly overlaps the anterior part of p4, Some variation is also observable on the and the posterior part of p4 the anterior part accessory cusps of the upper and lower pre- of m1. The anterior and posterior borders of molars. This variation concerns their presence p3 and p4 are almost straight, giving them a or not, as well as their development. The P2 rectangular outline. The p1 is small and situ- and P3 generally have small accessory cusps. ated lingually relative to p2. The p2 has a The anterior accessory cusp of p2 is small or posterior accessory cusp but not an anterior absent, while that of p3 is present but someti- one. The p3 has a posterior accessory cusp mes small. The m1 most commonly lacks a and a faint anterior one at the lingual side of metaconid. It has been observed, however, in the base of the anterior crest. The p4 has two some cases, as on MNHNP MAR G.12 from accessory cusps, an anterior and a posterior Maragha figured by De Mecquenem (1925, one. The anterior one is more robust and situ- pl. 9, fig. 7) and on RZ1- 4 from Ravin des ated slightly lingually. Behind the posterior Zouaves 1 (Koufos 2000). A small metaconid, accessory cusp, the cingulum is elevated. The appressed to the main body of the protoconid, premolars lack a lingual cingulum. The m1 has also been observed (Roussiakis 1996) on lacks a metaconid, while the talonid is small AMPG P.G. 95/1505 from Pikermi (specimen and two-cusped, retaining the entoconid and mentioned with the number 113 by Howell & the hypoconid, of which the hypoconid is the Petter 1985, tab. 6a). larger. There is a weakly developed cingulum The dimensions of the specimens at the anterior part of the paraconid, both lin- K4/∆388/1 and K4/∆388/2-3 are within the gually and buccally. There is no m2 or an range of variation of A. eximia from Pikermi alveolus for it. (Tables 5-6, figs. 4,5). The observable diffe- rences are not statistically significant. The Comparisons M1 of K4/∆388/3 is slightly large relative to Hyaena eximia was first described by Roth & the premolars, but is damaged at the area Wagner (1854) from the classical locality of between the trigonid and the talonid. The P3

475 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

and P4 of K4.7 are comparable to the largest lower premolars, lies close the minimum specimens from Pikermi (Fig. 4). The meta- values of the A. eximia from Pikermi. The style of the P4 of K4.7 is large (Table 5) but robusticity index for the p2 of DTK-126 that tooth is badly damaged and broken at the (sin.) is also similar to that of the A. eximia limit of the paracone and metastyle. specimen RZ1-4 (sin.) (Fig. 5). The morpho- The present specimens show significant logy of the lower premolars of C. bonisi, differences from Adcrocuta eximia leptoryn- however, differs from those of A. eximia. The cha BONIS & KOUFOS, 1981, from the Valle- lower premolars of C. bonisi have a more sian of Ravin de la Pluie. According to Bonis elliptical outline, especially anteriorly, while & Koufos (1981) and Koufos (2000), this the accessory cusps lie almost on the axis of subspecies differs from the typical A. eximia the teeth. In A. eximia the anterior cusps of in its longer snout, narrower palate and more the lower premolars are in a more lingual slender premolars. With regard to the lower position, and the anterior and posterior bor- premolars, there are no significant differences ders are almost straight, giving an almost rec- from the Pikermi A. eximia, something men- tangular shape to the teeth. tioned also by Howell & Petter (1985). The Most probably A. eximia is also present at p3 and p4 are slightly smaller in some of the Kerassiá 1 (K1) site. This is indicated by their dimensions (Fig. 5) and p4 has an index the specimen K1/∆104 (a left P4). Unfortuna- of compression (Wp4/Lp4) equal to 50.7 on tely, this specimen is so fragmentary that no RPL-15, which is slightly lower than the measurements can be taken. Pikermi range (Table 6). The indices of the The species A. eximia is known mainly upper premolars are within the range of A. from the Turolian (MN11-13) of Europe and eximia from Pikermi (Table 5, fig. 4). The Asia as well as from northern Africa, with its skull characters of A. eximia leptoryncha, maximum in the middle Turolian (MN12; however, distinguish that subspecies from the Howell & Petter 1985, Koufos 2000). Its Pikermi, Samos and Maragha A. eximia. The existence in the Vallesian of Hostalets de skull of A. eximia leptoryncha has a longer Pierola in Spain and Soblay in France, has snout and relatively narrower palate, the been shown to be unconfirmed (Howell & length/width ratio of the palate being 154 Petter 1985). Ravin de la Pluie, Ravin des (Bonis & Koufos 1981). The corresponding Zouaves 1 and Xirochori 1 are the only ratio (estimated following the methodology known late Vallesian localities with Adcro- of Bonis & Koufos 1981) for K4/∆388/1 is cuta eximia remains (De Bonis et al. 1988; 123, a value very close to those given by Koufos 2000). As shown earlier, the Ravin de Bonis & Koufos (1981) for the A. eximia la Pluie subspecies A. e. leptoryncha, is clear- from Pikermi (125) and Maragha (117). On ly different from the Kerassiá A. eximia. The P.A. 490/91 from Pikermi, this ratio is about Ravin des Zouaves 1 A. eximia is also diffe- 127. rent from the Kerassiá specimens. According The species Chasmaporthetes bonisi to Koufos (2000), the late Vallesian A. eximia KOUFOS, 1987 is known from the early from Ravin des Zouaves 1 has strong lingual Turolian locality of Ravin des Zouaves 5 and cingulum on the lower premolars, a character the late Turolian locality of Dytiko 1. This considered to be primitive by Koufos (2000). species differs from A. eximia mainly in the Such a cingulum is absent on the Kerassiá upper dentition. The M1 is larger than that of specimens, indicating that the Kerassiá A. A. eximia (Fig. 4) and the protocone of P4 is eximia is more similar to the typical Turolian more developed and directed slightly anteri- forms. orly. The P2 is also slightly less robust than that of A. eximia from Pikermi, but not signi- ficantly so. In C. bonisi the robusticity of the

476 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Figure 4 Logarithmic ratio diagram comparing the upper dentition of the Kerassia 4 Adcrocuta with A. eximia and C. bonisi from various localities. H. hyaena (standard) according to Howell & Petter (1980). Data for the Axios, Ravin des Zouaves 5 and Ravin de la Pluie hyaenids according to Koufos (2000).The rest from Table 5.

Family: GRAY, 1821 Description Subfamily: Machairodontinae GILL, 1872 The upper C (K4.14) is laterally compressed, Genus: Machairodus KAUP, 1833 has an anterior keel in a slightly lingual posi- Machairodus giganteus (WAGNER, 1848) tion, as well as a posterior one. Both keels Locality: Kerassiá 4 (K4) are poorly preserved, but show crenulations at least near the tip of the canine. The length Material at the base of the canine is about 30.0 mm K4.14: right upper canine. K4/∆69/1: mandi- and the width about 11.6 mm. The resulting ble, with both left and right mandibular rami. index (W x 100/L) Cs is 38.7, showing the The condyles are only partly preserved, while degree of compression of the canine. the mandibular angles and the coronoid pro- The mandible has a strong, downwardly cesses are broken. In the right ramus the expanded and angular mental crest. The i1-i3, p3-p4 and m1 are preserved, but the height of the symphysis measured anteriorly tips of i2 and i3 are broken and the posterior is about 75 mm. The area of the symphysis part of p4 is badly damaged. Only the root of just below the incisors and anteriorly is deep- the right ci is preserved. In the left ramus the ly grooved, while its lower part is smoother. complete dentition (i1-i3, ci, p3-p4 and m1) The upper part of the symphyseal area is situ- is preserved but the main cusp of p3, the ated in front of the mental crests. There are anterior half of p4 and the posterior half of two mental foramina. These foramina are on m1 are broken. K4/∆110/16: Left radius, the same level and closer to the lower margin complete. K4/∆82: Right Mt IV, badly dama- of the mandible than to the upper one. The ged. anterior mental foramen opens under the mid- dle of the postcanine diastema and the poste-

477 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Figure 5 Logarithmic ratio diagram comparing the lower dentition of the Kerassia 4 Adcrocuta with A. eximia and C. bonisi from various localities. H. hyaena (standard) according to Howell & Petter (1980). Data for the Ravin des Zouaves 1 & 5, Dytiko 1 and Ravin de la Pluie hyaenids according to Koufos (2000). A. eximia from Halmyropotamos according to Melentis (1968).The rest

rior one just in front of p3. The masseteric from p4. The mandibular corpus, at the fossa is deep and well defined by an acute region of the diastema and on its upper part, crest on its upper part. It extends anteriorly to is laterally concave and an acute crest is for- the posterior limit of m1 on the right ramus, med dorsally. There is no trace of a p2. The slightly more anteriorly on the left. The con- p3 is elongated (Table 7). The right p3 has a dyle is situated 1.5 cm lower than the level of faint anterior accessory cusp and the left one the cheek teeth, while the incisors and the a larger one. There is also a posterior acces- canine are clearly at a higher level. The man- sory cusp that is larger than the anterior one. dibular foramen opens close to the lower The greatest width of the tooth is in its poste- margin of the mandible and about 65 mm in rior part. In p4, both the anterior and the pos- front of the condyle. terior accessory cusps are strong, but the The lower incisors are arranged in an anterior one is thicker. Behind the posterior almost straight line, are conical in shape, accessory cusp there is also a small supple- increasing in size from i1 to i3 and situated mentary cuspid. Both the posterior accessory clearly in front of the lower canines. Their cusp and the supplementary cuspid are direc- tips are directed upwards. The total width of ted slightly backwards. The greatest width of the incisors (i3-i3) measures about 43 mm. the tooth lies posteriorly. In m1, the paracon- The lower canine is not significantly laterally id is robust while the protoconid is more compressed (Table 7). A large diastema sepa- slender and slightly longer. There is a small rates the lower canine from p3 (Table 7) and metaconid. The greatest width of the tooth a small diastema of 2.5 mm separates p3 lies in the posterior part of the paraconid.

478 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Comparisons known, however, on how many specimens Various species of Machairodus have been this has been observed and how much varia- described from the Late Miocene of Europe bility is involved. Judging from the drawings and Asia, but De Beaumont (1975) retained of some mandibles in lateral view, the arran- only two species, the Vallesian Machairodus gement of the lower incisors varies in M. aphanistus (KAUP, 1833) and the Turolian giganteus. On the mandible from Pikermi Machairodus giganteus (WAGNER, 1848). figured by Roth & Wagner (1854, pl. 9, fig. These two species are of comparable dimen- 1), for example, i2 is only just in front of i3. sions, but differ in many morphological char- On another mandible (Wagner 1857, pl. 5, acters. De Beaumont (1975) considered the fig. 11), i2 is significantly in front of i3. species Machairodus leoninus (ROTH & These observations show that the disposition WAGNER, 1854), Machairodus tarakliensis of the lower incisors varies. The width of the RIABININ, 1929, Machairodus palanderi lower incisor row (i3-i3) of K4/∆69/1 is 43 ZDANSKY, 1924, and possibly Machairodus mm, only slightly different from P.G. 01/100 tingii ZDANSKY, 1924, to be synonyms of M. (estimated at 47 mm). The height of the sym- giganteus. The available Kerassiá 4 speci- physis of P.G. 01/100 is about 67 mm, slight- mens are compared to bibliographic data as ly smaller than that of the Kerassiá mandible. well as to three undescribed specimens of M. De Beaumont (1975) mentioned that the post- giganteus from Pikermi stored in the AMPG canine diastema is generally large in M. (Table 7). giganteus (as also on the Machairodus from As is indicated by De Beaumont (1975) and Kerassiá 4) and shorter in M. aphanistus, but Koufos (2000) the upper canine of M. apha- he also mentioned that there is considerable nistus is less compressed than that of M. variation in this character. On the type speci- giganteus. According to Koufos (2000), the men of M. aphanistus from Eppelsheim, this index of compression (Wx100/L) of the upper diastema is 30-35 mm and in another speci- canine in M. aphanistus ranges from 41.5-52 men from the same locality, it is 35 mm (De and in M. giganteus from 38-41. The canine Beaumont 1975). The length of this diastema from Kerassiá 4 has an index of compression varies significantly in M. giganteus from equal to 38.7, well within the range of M. Pikermi. It is 55 mm in a specimen from giganteus. An upper canine from Pikermi Pikermi described by Roth & Wagner (1854, (AMPG no 1967/7) has an index of compres- pl. 9, fig. 1) and only 24 mm in another spe- sion 38.8. This specimen is attributed by cimen from the same locality described by Melentis (1968) to M. aphanistus, but in our Wagner (1857, pl. 5, fig. 11). On P.G. 01/100 opinion must be attributed in M. giganteus. from Pikermi, this diastema is 48.8 mm, For the Machairodus skull from Halmyro- which is slightly less than that of K4/∆69/1 potamos (Melentis 1968), the compression (Table 7). In P.G. 01/100, there is a minute index of the upper canine is about 40.6. This alveolus for p2 at the middle of the diastema. specimen has been referred to M. aphanistus De Beaumont (1975) mentioned that in M. by Melentis (1968), but was considered to giganteus there is no trace of p2. In our opi- belong to M. giganteus by Petter & Howell nion, however, such a character is not very (1987) and Koufos (2000). important since, as we mention also later, it is In lateral view, the lower incisors of sometimes present in Machairodus alberdiae K4/∆69/1 are clearly in front of the canines. GINSBURG, MORALES & SORIA, 1981, as well They are arranged in an almost straight line, as in species such as Metailurus parvulus while in P.G. 01/100 from Pikermi the incis- (Hensel, 1862). In the Kerassiá 4 Machai- ors are arranged in an arc. De Beaumont rodus, the width of the lower canines is much (1975) observed that in M. giganteus the less than the distance that separates them lower incisors are arranged in an arc. It is not from the symphysis. Such a character is

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always found in M. giganteus while in M. Eppelsheim (Fig. 6). The p3 of P.G. 01/100 aphanistus the opposite is observed (De has a larger anterior accessory cusp than Beaumont 1975). According to the same K4/∆69/1, but De Beaumont (1975) mentio- author, the lower canine of M. giganteus is ned that in M. giganteus the anterior cusp less compressed than that of M. aphanistus. varies from faint to well developed. The In a specimen of M. giganteus from Samos, accessory cusps of p4 in K4/∆69/1 are not the lower canine has dimensions (L x W) at different from those of P.G. 01/100 and P.G. the base of the crown of 18.0 x 14.0 (De 01/101 from Pikermi, the anterior one being Beaumont 1975), giving it an index of com- slightly more robust than the posterior one. In pression of 77.8. According to the measure- K4/∆69/1, the metaconid of M1 is slightly ments given by De Beaumont (1975), the more developed than in P.G. 01/100, P.G. lower canine of M. aphanistus from 01/101 and P.G. 01/102, in which feature it is Eppelsheim described by Kaup (1833, pl. I, more like two specimens of M. giganteus fig. 3) under the name "Agnotherium anti- figured by De Beaumont (1975, figs. 6e, 6f). quum" has an index of 59.6, while in another In M. giganteus, however, the m1 is, in specimen of M. aphanistus from Eppelsheim general, more slender than in M. aphanistus (BMNH 4996a) the corresponding index is (Fig. 6). 58.1. The index of compression of the lower The species M. pseudailuroides SCHMIDT- canine of K4/∆69/1 is 74.7, a value signifi- KITTLER, 1976, from the localities of cantly greater than in M. aphanistus and clo- Akçakoy and Eskihisar in Turkey, differs ser to M. giganteus. In P.G. 01/100 from from the Machairodus from Kerassiá in its Pikermi, the lower incisor has an index of shorter p4 and m1. The upper canine of this compression of 66.9. This value, however, is species is more slender than that of M. gigan- taken at the root of the tooth. Higher on the teus, with an index of compression of 35.9. crown, we would expect a greater value, M. alberdiae GINSBURG, MORALES & SORIA, since the anteroposterior diameter (L) of the 1981, from the Vallesian locality of Los canine decreases more than the transverse Valles de Fuentidueña has been considered as diameter (W). a possible ancestor of M. giganteus (Ginsburg The lower premolars and the lower carnas- 1999). This species has smaller dimensions sial of M. giganteus show no significant dif- than the specimens described here and some- ferences from those of M. aphanistus. As times preserves a p2. Its upper canine is sig- shown in Figure 6, the cheek teeth vary signi- nificantly less compressed ficantly in size, something also mentioned by (W x 100/L=50.0-53.9, n=2) than that of the De Beaumont (1975). Aside from the size Kerassiá 4 machairodont, but its lower canine variation, De Beaumont (1975) also emphasi- is slightly more compressed (W x 100/L=63.4- zed the morphological variation of M. gigan- 71.4, n=4). Most characteristic of M. alber- teus, a variation that concerns the postcanine diae is the large p3 relative to p4 and m1 diastema that we mentioned above, the deve- (Fig. 7). lopment of the anterior accessory cusp of p3, Two machairodonts have been described and the development of the metaconid/talonid from the Late Miocene of China, M. tingii complex of m1. As shown in Figure 6, the p3 and M. palanderi (Zdansky 1924). As we and p4 of M. giganteus are generally more mentioned earlier, M. palanderi has been elongated than they are in M. aphanistus. considered conspecific with M. gignateus by There are, however, specimens referred to M. De Beaumont (1975), and the same may be aphanistus with p3 and p4 equally or almost true also of M. tingii. As we can see (Fig. 7) equally elongated as those of M. giganteus, M. tingii is very similar in dimensions to as for example the M. aphanistus from K4/ x 69/1, but M. palanderi has shorter p3 Charmoille and the type specimen from relative to p4 and m1.

480 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Figure 6 Logarithmic ratio diagram comparing the Kerassia 4 Machairodus with M. giganteus and M. aphanistus from various loca- lities. M. aphanistus from Eppelsheim, Soblay and Charmoille according to Beaumont (1975), M. giganteus from Samos and M. ex gr. giganteus from Baccinello according to Rook et al. (1991), M. giganteus from Ravin X according to Koufos (2000), M. giganteus from Pikermi according to the authors.

Sotnikova (1992) described a new species, form, a compression index of P4 of 38.7 sig- M. kurteni, from the Late Miocene locality of nifies that P4 is not narrower than that of M. Kalmakpai in Kazakhstan. According to giganteus. The M. giganteus from Halmyro- Sotnikova (1992), this species has a propor- potamos, for example, has a compression tionally narrower P4 than other species of index equal to 34.3, which indicates a narro- Machairodus, retains a small P2, the tips of wer P4 compared to the Kalmakpai form. The the lower incisors are directed upwards, the same is true for the Ravin X skull portion of length of the lower premolars is small com- M. giganteus, which has a compression index pared with that of m1, and has completely of P4 equal to 36.4 (Koufos 2000). A similar lost the metaconid/talonid complex on M1. comparison shows that the P3 of the Kalmak- Sotnikova (1992) did not support her state- pai form is not shortened relative to P4, more ments with adequate comparisons, but actual- than it is in M. giganteus. Concerning the ly compared her specimens to ‘M. taraklien- lower teeth, the index Lm1 x 100/Lp3-4, sis’ from Taraklia, a form that has been consi- which shows the relative development of the dered a synonym of M. giganteus by De premolars, varies in the Kalmakpai form from Beaumont (1975) and a subspecies of M. 67.7 to 74.5 (n=3). These values are not dif- giganteus by Sotnikova (1992). Most of the ferent from the Pikermi specimens P.G. above mentioned characters of M. kurteni are 01/100 (73.2) and P.G. 01/101 (70.4). also found in M. giganteus. Concerning for According to Sotnikova (1992), the tips of example the upper teeth of the Kalmakpai the lower incisors of the Kalmakpai form are

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directed upwards, contrary to the Taraklia M. greater. The anteroposterior diameter of the giganteus, where they are directed forwards. proximal part is 29 mm. In P.G. 01/100, however, as well as on the The species M. giganteus is known from mandible figured by Roth & Wagner (1854, the Turolian of Halmyropotamos (possibly pl. 9, fig. 1), the tips of the lower incisors are MN11 or MN12), the early-middle Turolian also directed upwards. Other characters, such (MN11-12) of Ravin X in Greece and from as the presence of P2 and p2 are not reliable the middle Turolian (MN12) of Mont alone, but only in combination with other Leberon (France), Kalimanchi (Bulgaria), characters, since they seem variable. Samos, Pikermi (Greece) and Taraklia Sotnikova (1992) for example figured a man- (Moldavia) (Ginsburg 1999, Koufos 2000). It dible of M. laskarevi from Kalfa, where there is reported also under the name Amphimachai- is a p2 on the right ramus but not on the left. rodus giganteus from various localities in The same tooth varies in development in M. Spain, ranging from MN11 to MN13 (Fraile alberdiae, being absent or rudimentary. The et al. 1997). Rook et al. (1991) also mentio- P2 is in general absent in M. giganteus, but ned some specimens under the name present in M. palanderi and M. tingii. M. kur- Machairodus ex gr. giganteus from the MN12 teni has completely lost its metaconid/talonid locality of Baccinello-V3 in Italy. complex on m1. This character varies signifi- cantly in M. giganteus and can be almost Subfamily: TROUESSART, 1885 absent (De Beaumont 1975). From the above Genus: Metailurus ZDANSKY, 1924 observations is seen that, except for the last Metailurus cf. parvulus (HENSEL, 1862) character, M. kurteni is not clearly different Locality: Kerassiá 1 (K1) from M. giganteus and further comparison has to be made. Material The available radius is comparable in size K1/∆18: left mandible with ci, p3 and p4. to that described by Gaudry (1863, pl. 16, The m1 has fallen out and the tip of the cani- figs. 2, 3). The radial tuberosity is not very ne is broken. prominent. The radius has two wide grooves in its anterodistal part, a medial one for the Description extensors carpi radialis longior and brevior The mandibular corpus has a large mental indicis and a lateral one for the extensor com- foramen situated under the middle of the munis digitorum. These two grooves are postcanine diastema and a very small one separated by a blunt crest. More medially lies under the front end of p4. The lower canine is the groove for the extensor ossis metacarpi in a poor state of preservation, but shows an pollicis and more laterally the triangular arti- antero-lingual keel and most possibly a poste- cular surface for the ulna. A proximal part of rior one. The p3 has only a posterior acces- a radius from Halmyropotamos is also men- sory cusp and its greatest width is in its pos- tioned by Melentis (1969) under the name terior part. The p4 has an anterior and a pos- Machairodus aphanistus. In our opinion it is terior accessory cusp, the posterior one being not certain that this specimen represents a slightly stronger. As in p3, the greatest width Machairodus, since it has smaller dimen- occurs in the posterior part of the tooth. The sions. The maximum diameter of the head of alveolus for m1 is especially large and its that specimen is 23 mm (Melentis 1969), sig- posterior limit lies well up on the ascending nificantly smaller than in the radius of ramus of the mandible. Moreover, the mandi- Kerassiá 4. The present Mt IV has a badly bular corpus shows an abrupt thickening at damaged shaft and its distal epiphysis is the region of the m1. The above characters slightly broken. Its total length is 128 mm, may indicate that our specimen is a patholo- and must be expected to have been slightly gical one.

482 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Figure 7 Logarithmic ratio diagram comparing the Kerassia 4 Machairodus with other species. M. aphanistus from Eppelsheim (standard) according to Beaumont (1975), M. tingii and M. palanderi according to Zdansky (1924), M. alberdiae according to Ginsburg et al. (1981), M. pseudailuroides according to Schmidt-Kittler (1976) and M. kurteni according to Sotnikova (1992).

Discussion and comparisons without a mental crest. The height of the The species M. parvulus (HENSEL, 1862) has mandibular corpus in front of P3 is only a wide range from Spain to China. Following slightly smaller than that of the Chomateri Thenius (1951) and De Beaumont (1961), we specimen (Symeonidis 1978) or P.G. 01/103 include in this species the "Felidae indet., 3rd (Table 9). There is no alveolus for p2 species" (Gaudry 1863), leiodon between the lower canine and the p3. Such an WEITHOFER, 1888, Metailurus minor ZDANSKY, alveolus, with a single, very small root, exists 1924, and other remains referred to under on the specimen from Chomateri, on the spe- various specific or generic names. cimen P.G. 01/103 from Pikermi, as well as The available specimen from Kerassiá 1 in one specimen from China (Zdansky 1924, has been compared with specimens of M. ex. 3). parvulus from Pikermi, Chomateri and China. Unfortunately, the teeth are not very well We have included also in our comparison an preserved and some of their measurements undescribed until now specimen from could not be taken with the appropriate preci- Pikermi, with the catalogue number AMPG sion. The p3 is slightly smaller than other P.G. 01/103 (Table 9). The mandible of specimens of M. parvulus, but the indices of K1/∆18 has the typical form of the Felinae, the teeth are within the range of M. parvulus

483 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Figure 8 Logarithmic ratio diagram comparing the Kerassia 1 Metailurus with M. parvulus from Pikermi, Chomateri and China. Data from Table 9.

from China (Table 9). As we can see (Fig. 8), in the type specimen of P. ogygius the postca- the Greek specimens show a tendency to nine diastema measures about 17 mm (De smaller size than the Chinese specimens. Beaumont 1975), as opposed to 8,5 mm in Some of these specimens also show a narro- the Kerassiá one specimen. wer p4. The Kerassiá one specimen is slightly M. parvulus is known from the Turolian of smaller in most of its dimensions than other Halmyropotamos (possibly MN11 or MN12) specimens referred to M. parvulus, but the (Melentis 1968), the middle Turolian (MN12) observed differences are of little value, as the of Pikermi (Hensel 1862, Thenius 1951) and statistical sample is not adequate. The present Chomateri (Symeonidis 1978) in Greece, Los specimen, however, lacks the m1 and it is Mansuetos in Spain (Morales & Soria 1979, preferable to refer it as Metailurus cf. parvul- Fraile et al. 1997) and the late Turolian of El us. Arquillo in Spain (Morales & Soria 1979, Paramachairodus ogygius (KAUP, 1832) Fraile et al. 1997). has teeth comparable is size with the largest specimens of M. parvulus from China. The GENERAL DISCUSSION AND Kerassiá one specimen, however, differs from CONCLUSIONS P. ogygius. The height of the mandibular cor- The carnivores of K1, K3 and K4 are consis- pus in front of p3 measures about 26 mm tent with a Turolian age for these sites. A. (measurement from the figure) on the speci- eximia is very common in the middle men figured by Kaup (1833, pl. 2, fig. 3), as Turolian, but rarer in the early or late opposed to 17.5 mm on K1/∆18. Moreover, Turolian (Howell & Petter 1985). In Greece,

484 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

A. eximia seems absent from the late Turolian Municipality of Nileas and the General localities, such as those of Dytiko (De Bonis Secretary of Research and Technology et al. 1992, Koufos 2000). The same is true (Project 95∑YN107). Moreover, we would for the rare M. giganteus, which is absent like to thank Dr L. Ginsburg (Muséum from Dytiko. Both species are present in the National d’Histoire Naturelle de Paris) who Main Bone Beds of Samos (Bernor et al. allowed the senior author to study material in 1996), dated to 7.1 Ma (Swisher 1996), the collections of MNHNP and who helped while A. eximia is also present in the older him in various ways. The authors are also White Beds of Samos (Bernor et al. 1996; grateful to the revisers of this study, Swisher 1996). According to the recent calib- Professor Alan Turner (John Moores ration of the MN zones (Steininger 1999), the University, Liverpool) and Dr Lars Werdelin Main Bone Beds of Samos can be placed in (Senior Curator, Swedish Museum of Natural MN12. In addition to the carnivores discus- History, Department of Palaeozoology) for sed here, the K1 faunal list (Theodorou et al. their valuable comments and linguistic this volume) also includes a Tragoportax, improvements on the manuscript. provisionally referred to Tragoportax cf. amalthea. T. amalthea (Roth & Wagner 1854) REFERENCES is a boselaphine that is well known from Adrover, R., Alcalá, L., Mein, P., Moissenet, E. & many Late Miocene localities, with a strati- Orrios, J., 1986 – Mamíferos del Turoliense Medio en graphic range from the early to the middle la Rambla de Valdecebro (Teruel) – Estudios Turolian (Gentry et al. 1999). Geológicos 42: 495-509 The suids referred to M. major erymanthius Alcalá, L., 1994 – Macromamíferos neógenos de la fosa by Van der Made & Moyà-Solà (1989) most de Alfambra-Teruel – Museo Nacional de Ciencias probably come from K1 and according to Naturales, Teruel these authors indicate a middle Turolian age. Bernor, R.L., Solounias, N., Swisher III, C.C. & Van Kostopoulos et al. (2001), however, have Couvering, J.A., 1996 – The correlation of three shown that the Kerassiá M. major is smaller classical ‘Pikermian’ faunas-Maragheh, than the Pikermi M. major erymanthius, and Samos, and Pikermi-with the European MN unit probably represents a new subspecies. The system – in: Bernor, R.L., Fahlbush, V. & Mittmann, geological prospecting of the Kerassiá sites H.-W. (eds.) – The Evolution of Western Eurasian has shown that K1 is situated at a higher Neogene Mammal Faunas – pp. 137-154, Columbia level than K3 and K4, so K1 must be slightly University Press, New York younger. K3 and K4 possibly belong to the Cordella, A., 1878 – La Grèce sous le rapport géologique same stratigraphic level, but no suids have et minéralogique – Parent, Paris been found in these sites until now (Theo- Crusafont-Pairó, M. & Petter, G., 1969 – Contribution à dorou et al. this volume). l'étude des Hyaenidae. La sous-famille des A biostratigraphic distinction between K1, Ictitheriinae – Annales de Paléontologie (Vertébrés) K3 and K4 is not possible at the moment, but 55 (1): 89-127 judging from the above data, an early to mid- De Beaumont, G., 1961 – Recherches sur Felis attica dle Turolian age is plausible for the fauna of Wagner du Pontien eurasiatique avec quelques obser- Kerassiá. In the future, we plan to continue vations sur les genres Pseudaelurus Gervais et the fieldwork, while sedimentological and Proailurus Filhol – Nouvelles Archives du Muséum taphonomic study of the numerous fossilife- d'Histoire Naturelle de Lyon 6: 1-45 rous sites at Kerassiá is already in progress. De Beaumont, G., 1969 – Brèves remarques sur Plioviverrops Kretzoi (Carnivora) – Bulletin des ACKNOWLEDGEMENTS Laboratoires de Géologie, Minéralogie, Géophysique The excavations at Kerassiá have been finan- et du Musée géologique de l'Université de Lausanne ced by the University of Athens, the 180: 1-7

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De Beaumont, G., 1975 – Recherches sur les Félidés Gaudry, A., 1862-67 – Animaux Fossiles et Géologie de (Mammifères, Carnivores) du Pliocène inférieur des l’Attique – F. Savy, Paris Sables à Dinotherium des environs d'Eppelsheim Gentry, A.W., Rössner, G.E. & Heizmann, E.P.J., 1999 – (Rheinhessen) – Archives des Sciences 28 (3): Suborder Ruminantia – in: Rössner, G.E. & Heissig, 369-405 K. (eds.) – The Miocene Land Mammals of Europe – De Beaumont, G. & Mein, P., 1972 – Recherches sur le pp. 225-258, Dr Friedrich Pfeil, München genre Plioviverrops Kretzoi (Carnivora, ?Hyaenidae)– Ginsburg, L., 1999 – Order Carnivora – in: Rössner, G.E. Comptes Rendus des Séances 25 (3): 383-394 & Heissig, K. (eds.) – The Miocene Land Mammals De Bonis, L., 1994 – Les gisements des mammifères du of Europe – pp. 109-148, Dr Friedrich Pfeil, Miocène supérieur de Kemiklitepe, Turquie. 2. München Carnivora – Bulletin du Muséum national d’Histoire Ginsburg, L., Morales, J. & Soria, D., 1981 – Nuevos naturelle, C 16 (1): 19-39 datos sobre los carnivoros de Los Valles de De Bonis, L., Bouvrain, G. & Koufos, G.D., 1988 – Late Fuentidueña (Segovia) – Estudios Geológicos 37: Miocene mammal localities of the Lower Axios 383-415 Valley (Macedonia, Greece) and their stratigraphic Hensel, R.F., 1862 – Über die Reste einiger Säugetier- significance – Modern Geology 13: 141-147 arten von Pikermi in der Münchener Ammlung – De Bonis, L., Bouvrain, G., Geraads, D. & Koufos, G.D., Monatsberichte der Akademie der Wissenschaften 27: 1992 – Diversity and paleoecology of Greek late 560-569 Miocene mammalian faunas – Palaeogeography Howell, F.C. & Petter, G., 1980 – The Pachycrocuta and Palaeoclimatology Palaeoecology 91: 99-121 Hyaena lineages (Plio- and extant species De Bonis, L. & Koufos, G.D., 1981 – New Hyaenid of the Hyaenidae). Their relationships with Miocene (Carnivora, Mammalia) in the Vallesian (Late ictitheres: Palhyaena and Hyaenictitherium – Miocene) of Northern Greece – Scientific Annals of Géobios 13 (4): 579-623 the Faculty of Physics and Mathematics, University Howell, F.C. & Petter, G., 1985 – Comparative observa- of Thessaloniki 21: 79-94 tions on some Middle and Upper Miocene hyaenids. De Bonis, L. & Koufos, G.D. 1991 – The late Miocene Genera: Percrocuta Kretzoi, Allohyaena Kretzoi, small carnivores of the lower Axios valley Adcrocuta Kretzoi (Mammalia, Carnivora, Hyaeni- (Macedonia - Greece) – Géobios 24 (2): 361-379 dae) – Géobios 18 (4): 419-476 De Bonis, L. & Koufos, G.D., 1994 – Some Hyaenidae Jacobi, B., 1982 – Zur Stratigraphie und Sedimentpetro- from the Late Miocene of Macedonia (Greece) and a graphie der Neogenen Süßwasserablagerungen im contribution to the phylogeny of the hunting hyaenas Bereich von Agia Anna im Nordosten der Insel Euböa – Münchner Geowissenschaftliche Abhandlungen 26: (Ägäis) – Diplomarbeit, Christian-Albrechts- 81-96 Universität, Kiel De Mecquenem, R., 1924-25 – Contribution à l’étude des Kaup, J.J., 1832-1839 – Description d’ossements fossiles fossiles de Maragha – Annales de Paléontologie 13: de Mammifères inconnus jusqu’à présent qui se trou 135-160; 14: 1-34 vent au Muséum grand-ducal de Darmstadt, Darm- Déprat, J., 1904 – Étude géologique et pétrographique de stadt l’île d’Eubée – Besançon, Paris Köhler, R.W., 1983 – Zur Stratigraphie, Sedimentologie Fraile, S., Pérez, B., De Miguel, I. & Morales, J., 1997 – und Petrographie Neogener Ablagerungen im Gebiet Revisión de los carnívoros presentes en los yacimien Kerassiá – Papades - Ag. Anna im Nordosten der tos del Neógeno español – in: Calvo, J.P. & Morales, Insel Euböa (Ägäis) – Diplomarbeit, Christian- J. (eds.) – Avances en el conocimiento del Terciario Albrechts-Universität, Kiel Ibérico – pp. 77-80 Kostopoulos, D.S., Spassov, N. & Kovachev, D., 2001 – Gaudry, A. & Lartet, E.A.I.H., 1856 – Sur les résultats Contribution to the study of Microstonyx: evidence des recherches paléontologiques entreprises dans from Bulgaria and the SE European populations – l’Attique sous les auspices de l’Académie – Comptes Geodiversitas 23 (3): 411-437 Rendus des Séances de l’Académie des Sciences 43: Koufos, G.D., 1987 – Chasmaporthetes bonisi, a new 271-274 hyaenid (Carnivora, Mammalia) from the late

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ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Miocene of Macedonia (Greece) – Bulletin de la of Europe in the Department of Geology – British Société Géologique de France 3 (5): 913-920 Museum (Natural History) Geology, London Koufos, G.D., 2000 – Revision of the late Miocene car Qiu, Z., 1985 – Restudy of "Ictitherium hyaenoides" in nivores from the Axios valley, Macedonia, Greece – the Lagrelius Collection – Bulletin of the Geological Münchner Geowissenschaftliche Abhandlungen 39: Institutions of the University of Uppsala 11: 91-112 51-92 Qiu, Z., Huang, W. & Guo, Z., 1979 – Hyaenidae of the Koufos, G.D., Koutsouveli, A., Galanakis, D., Qingyang (K'Ingyang) hipparion fauna – Vertebrata Sylvestrou, I., Vlachou, T., 1999 – A new Late Mio- Palasiatica 17 (3): 200-221 cene mammalian locality from Velestinon, Thessaly, Rook, L., Ficcarelli, G. & Torre, D., 1991 – Messinian Greece. Contribution to the biochronology of the carnivores from Italy – Bollettino della Societá Neogene deposits – Comptes Rendus de l’Académie Paleontologica Italiana 30 (1): 7-22 des Sciences 328 (7): 479-483 Roth, J. & Wagner, A., 1854 – Die fossilen Knochen- Kretzoi, M., 1952 – Die Raubtiere der Hipparionfauna Ueberreste von Pikermi in Griechenland – Abhand- von Polgardi – Annales Instituti Geologici Publici lungen der Bayerischen Akademie der Wissen- Hungarici 40 (3): 1-41 schaften 7: 371-464 Major, C.I.F., 1888 – Sur un gisement d’ossements fossi- Roussiakis, S.J., 1996 – Contribution to the study of the les dans l’île de Samos, contemporains de l’âge de mammals of the classical locality of Pikermi – Pikermi – Comptes Rendus des Séances de Unpublished PhD Thesis, University of Athens l’Académie des Sciences 107: 1178-1181 Schmidt-Kittler, N., 1976 – Raubtiere aus dem Major, C.I.F., 1894 – Le gisement ossifère de Mytilini et Jungtertiär Kleinasiens – Palaeontographica, A 155: cataloque d'ossements fossiles recueillis à Mitylini, 1-131 île de Samos, et déposés au Collège Galliard, à Semenov, Y.A., 1985 – Ictitherium pannonicum Lausanne – pp. 1-51, Lausanne (Carnivora, Viverridae) from Maeotic deposits of the Melentis, J.K., 1968 – Studien über fossile Vertebraten Northern Black Sea area – Vestnik Zoologii 6: 23-27 Griechenlands. 19. Die Pikermifauna von Halmyro- Semenov, Y.A., 1989 – Ictitheres and morphologically potamos (Euböa - Griechenland). 1. Teil: Odontologie related hyaenas from the Neogene of the USSR – und Kraniologie – Annales Géologiques des Pays Naukova Dumka, Kiev Helléniques 19: 285-411 Solounias, N., 1981 – The Turolian fauna from the island Melentis, J.K., 1969 – Studien über fossile Vertebraten of Samos, Greece, with special emphasis on the Griechenlands. 28. Die Pikermifauna von Halmyro- hyaenids and bovids – Contributions to Vertebrate potamos (Euböa - Griechenland). 2. Teil: Osteologie Evolution 6: 1-232 – Annales Géologiques des Pays Helléniques 21: 217- Sotnikova, M.V., 1992 – A new species of Machairodus 306 from the late Miocene Kalmakpai locality in eastern Mitzopoulos, M.K., 1947 – Die Verbreitung der Kazakhstan (USSR) – Annales Zoologi Fennici 28: Pikermistufe auf der Insel Euböa – Annales 361-369 Géologiques des Pays Helléniques 1: 209-216 Steininger, F.F., 1999 – Chronostratigraphy, Geochrono- Morales, J. & Soria, D., 1979 – Nuevos datos sobre los logy and Biochronology of the Miocene "European carnivoros del área de Teruel. Sintesis y biostrati Land Mammal Mega-Zones" (ELMMZ) and the grafía – Estudios Geológicos 35: 497-504 Miocene "Mammal-Zones (MN-Zones)" – in: Ozansoy, F., 1965 – Étude des gisements continentaux et Rössner, G.E. & Heissig, K. (eds.) – The Miocene des mammifères du Cénozoique de Turquie – Land Mammals of Europe – pp. 9-24, Dr Friedrich Mémoires de la Société Géologique de France 102: Pfeil, München 1-92 Swisher III, C.C., (1996): New 40Ar/39Ar dates and Petter, G. & Howell, F.C., 1987 – Machairodus africanus their contribution toward a revised chronology for the Arambourg, 1970 (Carnivora, Mammalia) du late Miocene of Europe and West Asia – in: Bernor, Villafranchien d’Ain Brimba, Tunisie – Bulletin du R.L., Fahlbush, V. & Mittmann, H.-W. (eds.) – The Muséum national d’Histoire naturelle 16 (1): 97-119 Evolution of Western Eurasian Neogene Mammal – Pilgrim, G.E., 1931 – Catalogue of the Pontian Carnivora pp. 64-77, Columbia University Press, New York

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Symeonidis, N.K., 1978 – Ein Schädel von Metailurus Italiana 28 (2-3): 329-339 parvulus (Hensel) aus Pikermi (Attica, Griechenland) Wagner, A., 1848 – Urweltliche Säugthier-Ueberreste aus – Annales Géologiques des Pays Helléniques 29 (2): Griechenland -– Abhandlungen der Bayerischen 698-703 Akademie der Wissenschaften 5: 335-378 Thenius, E., 1951 – Zur odontologischen Charakteristik Wagner, A., 1857 – Neue Beiträge zur Kenntniss der von "Felis" leiodon aus Pikermi (Griechenland) – fossilen Säugthier-Ueberreste von Pikermi Abhand- Neuen Jahhrbuch für Geologie und Paläontologie 3: lungen der Bayerischen Akademie der Wissen- 88-96 schaften 8: 11-158 Theodorou, G., Athanassiou, A., Roussiakis, S. & Weithofer, A., 1888 – Beiträge zur Kenntniss der Fauna Iliopoulos, G., 1998 – Preliminary results on the von Pikermi bei Athen – Beiträge zur Paläontologie recent excavations of the Kerassiá locality (Euboea) – Oesterreich-Ungarns 6 (3): 225-292 in: abstracts Interim-Colloquium/RCMNS Werdelin, L., 1988a – Studies of fossil hyaenids: the ‘Mediterranean Neogene Cyclostratigraphy in mari- genera Thalassictis Gervais ex Nordmann, Palhyaena ne-continental palaeoenvironments’, Patras-Greece, Gervais, Hyaenictitherium Kretzoi, Lycyaena Hensel 27-29 May 1998 and Palinhyaena Qiu, Huang & Guo – Zoological Theodorou, G., Athanassiou, A., Roussiakis, S. & Journal of the Linnean Society 92: 211-265 Iliopoulos, G., this volume – Remarks on the Late Werdelin, L., 1988b – Studies of fossil hyaenids: the Miocene Vertebrates of Kerassiá (Northern Euboea, genera Ictitherium Roth & Wagner and Sinictitherium Greece) Kretzoi and a new species of Ictitherium – Zoological Theodorou, G.E., Roussiakis, S.J. & Athanassiou, A., Journal of the Linnean Society 93: 93-105 1995 – Contribution to the study of the terrestrial Werdelin, L. & Solounias, N., 1991 – The Hyaenidae: Neogene of Greece: Artiodactyla and Rhinocerotidae , systematics and evolution – Fossils and from the Kerassiá and Chalkoutsi localities – in: Strata 30: 1-105 abstracts 10th RCMNS Congress, 4 - 9 September Woodward, A.S., 1901 – On the bone beds of Pikermi, 1995, Bucharest - Romanian Journal of Stratigraphy Attica and on similar deposits in Northern Euboea – Torre, D., 1989 – Plioviverrops faventinus n. sp., a new Geological Magazine 8 (11): 481-486 carnivore of late Messinian age – Bollettino della Zdansky, O., 1924 – Jungtertiäre Carnivoren Chinas – Societá Paleontologica Italiana 28 (2-3): 323-327 Palaeontologia Sinica 2 (1): 1-149 Van der Made, J. & Moyà-Solà, S., 1989 – European (Artiodactyla) from the Late Miocene Received 19 May 2001 onwards – Bollettino della Societá Paleontologica Accepted 16 September 2002

488 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

PLATE 1 Plioviverrops sp., Kerassia 3. Fig. 1: Left humerus (K3/B1/4), anterior view. Fig. 2: Head of a left humerus (K3.206), juvenile, proximal view. Fig. 3: Distal epiphysis of a right femur (K3/B1/15), juvenile, distal view. Fig. 4: Left Mt IV (K3.191), lateral view. Fig. 5: Right calcaneum (K3.192) medial view. x 1. cf. Ictitherium pannonicum, Kerassia 3. Fig. 6: Right mandibular ramus (K3/204), labial view. Fig. 7: idem, occlusal view. Fig. 8: idem, lingual view. x 3/4.

489 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

PLATE 2 Adcrocuta eximia, Kerassia 4. Fig. 1: Skull fragment (K4/∆388/1), occlusal view. Fig. 2: idem, labial view. x 3/4.

490 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

PLATE 3 Adcrocuta eximia, Kerassia 4. Fig. 1: Left mandibular ramus (K4/∆388/2), labial view. Fig. 2: idem, occlusal view. Fig. 3: idem, lingual view. x 3/4.

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PLATE 4 Machairodus giganteus, Kerassia 4. Fig. 1: Mandible (K4/∆69/1), labial view. Fig. 2: idem, occlusal view. x 1/2.

492 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

PLATE 5 Machairodus giganteus, Kerassia 4. Fig. 1: Right upper canine (K4.14), labial view (x 1/2). Fig. 3: Left radius (K4/∆110/16) anterior view (x 1/3). Machairodus giganteus, Pikermi. Fig. 2: Left upper canine (No 1967/7), lingual view (x 1/2). Fig. 4: Left mandi- bular ramus (P.G.01/100), labial view. Fig. 5: idem, occlusal view. x 1/2.

493 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

PLATE 6 Metailurus cf. parvulus, Kerassia 1. Fig. 1: Left mandibular ramus (K1/∆18), labial view. Fig. 2: idem, occlusal view. Fig. 3: idem, lingual view. x 1. Metailurus parvulus, Pikermi. Fig. 4: Left mandibular ramus (P.G.01/103), labial view. Fig. 5: idem, occlusal view. x 1.

494 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Table 1 Measurements of the humerus. Lmax: maximum length; Lf: maximum functional length; DTpr: maximum transverse dia- meter of the proximal epiphysis; DAPpr: maximum anteroposterior diameter of the proximal epiphysis; DTdist: maximum trans- verse diameter of the distal epiphysis; DAPdist: maximum anteroposterior diameter of the distal epiphysis.

Table 2 Measurements of the calcaneum. Hmax: maximum height; DTpr: maximum transverse diameter of the head; DTcol: transverse diameter at the middle of the neck; DTmax: maximum transverse diameter: DAPpr: maximum anteroposterior diame- ter of the head; DAPcol: anteroposterior diameter at the middle of the neck; DAPmax: maximum anteroposterior diameter.

Table 3 Measurements of the fourth metatarsal. Lmax: maximum length; DAPpr: maximum anteroposterior diameter of the proximal end; DAPdistart: anteroposterior diameter of the distal articular surface; DTpr: maximum transverse diameter of the proximal end; DTdistart: transverse diameter of the distal articular surface; DTdistmax: maximum transverse diameter of the distal end.

Table 4 Measurements of cf. I. pannonicum from Kerassia 3 and I. pannonicum from Polgardi and the Black Sea.

495 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Table 5 Measurements of A. eximia upper teeth from Kerassia and Pikermi.The data concerning the A. eximia from Pikermi are from Roussiakis (1996).These are based on the specimens mentioned by Howell & Petter (1985, tab. 6a) as well as on AMPG P.A.490/91 described by Roussiakis (1996) and original measurements of MNHNP PIK. 3000 and AMPG P.G.95/1507.The last specimen is mentioned as unnumbered by Howell & Petter (1985, tab. 6a).The statistical data have been recalculated, since there are some statistical inaccuracies in Howell & Petter (1985, tab. 7).

Table 6 Measurements of A. eximia lower teeth from Kerassia and Pikermi.The data concerning the A. eximia from Pikermi are from Roussiakis (1996).These are based on the specimens mentioned by Howell & Petter (1985, tab. 6a) as well as on AMPG P.A. 57/91, P.A.445/91 and P.A.1296/91 described by Roussiakis (1996) and original measurements of MNHNP PIK. 3001, AMPG P.G.95/1505 and P.G.95/1506.The last two specimens are mentioned as No 113 and unnumbered respectively, by Howell & Petter (1985, tab. 6a).The statistical data have been recalculated since there are some statistical inaccuracies in Howell & Petter (1985, tab. 7).

496 ROUSSIAKIS & THEODOROU: Late Miocene carnivora from Kerassiá

Table 7 Measurements of M. giganteus from Kerassia 4 and Pikermi

Table 8 Measurements of the radius. Lmax: maximum length; DTpr: maximum transverse diameter of the proximal epiphysis; DTcol: transverse diameter of the neck; DAPpr: maximum anteroposterior diameter of the proximal epiphysis; DTdia: transverse diameter at the middle of the diaphysis; DAPdia: anteroposterior diameter at the middle of the diaphysis; DTdist: maximum trans- verse diameter of the distal epiphysis; DAPdist: maximum anteroposterior diameter of the distal epiphysis.

Table 9 Measurements of M. cf. parvulus from Kerassia 1 and M. parvulus from Pikermi, Chomateri and China.

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