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Zeitschrift/Journal: Beiträge zur Paläontologie

Jahr/Year: 2009

Band/Volume: 31

Autor(en)/Author(s): Koufos George D.

Artikel/Article: The Late Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection - 5. 57-105 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Beitr. Paläont., 31:57-105, Wien 2009

The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 5. Carnivora

by

George D. Koufos*)

Koufos, G.D., 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 5. Carnivora. — Beitr. Palaont., 31:57-105, Wien.

Abstract Keywords: Late Miocene, Samos, Greece, Mammalia, Carnivora, Systematics. The new collection from the late Miocene localities of the Mytilinii Basin, Samos, Greece include several carnivores which are studied in the present article. The study and Zusammenfassung comparison of the material allow the determination of seven different taxa which are distributed in the various Die neuen Aufsammlungen aus den obermiozänen fossiliferous sites. Fundstellen des Mytilinii Beckes (Samos, Griechenland) MLN: Protictitherium crassum, Hyaenictitherium cf. beinhalten einige Carnivoren, die in der vorliegenden wongii. Arbeit untersucht werden. Die Untersuchungen erlauben M TLA: Parataxidea maraghana, Hyaenictitherium wongii, eine Zuordnung der Funde, die aus unterschiedlichen Adcrocuta eximia, Metailurus parvulus. Lokalitäten stammen, zu sieben Arten. MTLB: Plioviverrops orbignyi, Hyaenictitherium wongii. M LN : Protictitherium crassum, Hyaenictitherium cf. MTLC: Hyaenictitherium wongii. wongii. PMMS (?MTLA): Adcrocuta eximia, gi- M TLA: Parataxidea maraghana, Hyaenictitherium wongii, ganteus. Adcrocuta eximia, Metailurus parvulus. All the determined taxa, exceptProtictitherium crassum, are MTLB: Plioviverrops orbignyi, Hyaenictitherium wongii. known from the old collections of Samos but in the new M TLC: Hyaenictitherium wongii. collection there are some specimens, adding new data to PMMS (PMTLA): Adcrocuta eximia, Machairodus gi- them. The speciesProtictitherium crassum was found for the ganteus. first time in the late Miocene of Samos; it is known from Mit Ausnahme von Protictitherium crassum, sind alle an­ the neighboring area, as well as being reported from Con­ deren Taxa von früheren Aufsammlungen bekannt, aber tinental Greece and Asia Minor. A ll the studied material unter den neuen Funden ergaben sich neue Erkenntnisse was described and compared with the known material from zu den Arten. Protictitherium crassum ist zum ersten Mal Greece and Eurasia. Bearing the stratigraphic distribution aus dem Obermiozän von Samos beschrieben. Diese A rt of each taxon and their presence in the Samos fauna in ist aus angrenzenden Gebieten bekannt, zum Bespiel vom mind, a middle Turolian age is possible for the localities griechischen Festland und aus Kleinasien und wurde auch Mytilinii-1 A, B (M TLA, MTLB). The carnivoran fauna mit dem bisher bekannten Material aus Griechenland und of the other localities is poor and does not allow a precise Eurasien verglichen. Berücksichtigt man die stratigraphi­ age determination, except that of late Miocene. However, sche Verbreitung dieses Taxons und seine Anwesenheit in the various localities have been dated using the whole fauna der Samos Fauna, so ist Mittel-Turolium als Alter für die and magnetostratigraphy (see chapter 16 of this volume). Lokalitäten Mytilinii-1 A, B (MTLA, MTLB) möglich. Die Carnivorenreste aus den anderen Lokalitäten sind gering und lassen eine genauere Alterseinstufung, die über das Obermiozän hinausgeht, nicht zu. Die Datierung der verschiedenen Fundstellen erfolgte über die gesamte Fauna und mit Hilfe der Magnetostratigraphie (siehe 0 Prof. George D. Koufos, Aristotle University ofThessaloniki. Kapitel 16, dieser Band). Department of Geology. Laboratory of Geology and Palaeon- tology. GR-54124 Thessaloniki, Greece. e-mail:koufos@geo. Schlüsselworte: Obermiozän, Samos, Griechenland, auth.gr Mammalia, Carnivora, Systematik. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

58 Beitr. Paläont., 31, Wien, 2009

Forsyth M ajor Pilgrim (1931) Solounias (1981) Bernor et al. N O W , 2 0 0 7 Present Article (1894) (1996)

Mus tela palaeattica Undarctos atticus Ursavus cf. depereti Ursavus cf. depereti Ursavus cf. depereti Parataxidea maraghana Promephitis larteti Promeles palaeatticaIndarctos atticus atticus Indarctos atticus Plioviverrops orbignyi Meles maraghana Promephitis larteti Promeles palaeatticaPromeles palaeatticaPromeles palaeatticaProtictitherium crassum

Parataxidea (?) Ictitherium orbignyi Promephitis larteti Promephitis larteti Promephitis larteti Hyaenictitherium wongii maraghana Parataxidea marag­Parataxidea polaki Parataxidea / marag­ Ictitherium robustum Ictitherium orbignyi Adcrocuta eximia hana maraghana hana Ictitherium hippari- Ictitherium hippari- Plioviverrops Plioviverrops Plioviverrops Metailurus parvulus onum onum orbignyi orbignyi orbignyi Ictitherium viver- Ictitherium viver- Ictitherium viver- Lycyaena chaeretis Lycyaena chaeretis Machairodus giganteus rinum rinum rinum Hyaenotherium Hyaenotherium Hyaena eximia Crocuta eximia lhalassictis wongii wongii wongii Thalassictis hyae- Machairodus Felis neas Lycyaena chaeretis Lycyaena chaeretis sp. noides

Thalassictis (Lycyae- Felis neas Belbus beaumonti Belbus beaumonti no) n. sp.

Thalassictis (Lycyae­ Adcrocuta eximia Adcrocuta eximia na) chaeretis Hyaenictis eximia Felis attica Felis attica Felis attica Metailurus major Metailurus major Metailurus major Metailurus parvulus Metailurus parvulus Machairodus gigan- Metailurus parvulus teus giganteus Machairodus gigan- teus

Table 1: Various faunal lists of the carnivoran taxa found on Samos.

1. Introduction ity of the carnivores belong toHyaenictitherium wongii, a well-known and widespread late Miocene hyaenid. In the The late Miocene mammal fauna of Samos has been present article, the new collection of carnivores is described known since the last decades of the 19th century, when and compared with the old Samos material, as well as with C.J. Forsyth Major found and described several fossils. the late Miocene Eurasian material. The material has been However, the presence of the mammal fossils on Samos originally compared with the material of Axios Valley, was known earlier but without description (Koufos, (Macedonia, Greece), housed at LGPUT (the best dated this volume). Since 1994, a team from the Laboratory of Late Miocene collection of the Eastern Mediterranean Geology and Palaeontology of the Aristotle University of under certain stratigraphic control) for the first time, as Thessaloniki, led by the author, excavated on Samos and well as with the old collections of Samos and Piker mi collected a large amount of fossils. As all the old material stored at A M PG , MNHN, NHML and NHMW. The of Samos is mixed and without or of limited stratigraphic carnivores of Prof. J. Melentis collection from Samos were indication, there are difficulties in the determination of also included in this article. The collection was described the material and the age of the fauna. Although the new earlier (Koufos & M elentis, 1982) but it is included in collection is not as rich as the old ones, its composition, the present paper with more details and comparisons. A ll as well as the certain and precise stratigraphic control can the studied material is housed at NHM A, which is located give good taxonomic and biochronologic results. The old in the village of Mytilinii, Samos. collections of Samos include several carnivores, which The studied material has been unearthed from the follow­ have been described by various authors. Few carnivores are ing localities of the Mytilinii Basin: mentioned in the original list ofF orsyth M ajor (1894) Locality Mytilinii-4 (MLN). The locality “Mytilinii-4”, who was the first systematic excavator and collector of MLN is situated in the southern outcrops of Potamies Samos mammal fossils. The late Miocene carnivoran fauna ravine and seems to be identical to B. Brown’s Q4(K oufos of Samos slowly increased at first (Pilgrim, 1931), while et al., (1997); Kostopoulos et al., 2003). This locality later, it increased remarkably(S olounias, 1981; Bernor yielded few carnivores. It is dated to middle Turolian (MN et al., 1996; NOW, 2007), (Tab. 1). 12), but it is older than M TL, having an age of ~7.5 Ma The new collection of Samos includes several carnivores, (Kostopoulos et al., 2003; Koufos et ah, this volume). mainly hyaenids, and less mustelids and felids. The major­ Locality Mytilinii-1 (MTL). It is located in the Adrianos ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 59

ravine and can be correlated to B. Brown’s Q l or QA. Four SAM = Samos old collections different fossiliferous sites have been recognized: M TLA, VAT = Vathylakkos-3, Axios Valley, Macedonia, Greece MTLB, M TLC, and MTLD. VTK = Vathylakkos-2, Axios Valley, Macedonia, Greece M T LA corresponds to the site where Prof. Melentis excavated; MTLB and M T LC are new sites. M TLD The specimens of the various museums or institutes, except corresponds to the site where B. Brown excavated, as NH M A, are given in the text by the abbreviation of the we found several remains of his excavations (Koufos et museum or institute, the locality and the serial number ah, 1997; Kostopoulos et ah, this volume). The studied of the specimen. material was found in M TLA, MTLB, and M TLC. A ll sites are dated to middle Turolian (MN 12), more precisely from 7.1-7.0 Ma, according to the palaeomagnetic record 2. Palaeontology (Kostopoulos et ah, 2003; K oufos et ah, this volume). PM M S Collection. The collection made by Prof. J. M e­ Order Carnivora Bowdich, 1821 lentis, who excavated on Samos in 1963, originated from Family Mustelidae Swainson, 1835 Adrianos ravine. Prof. J. Melentis showed me the exact site Subfamily Melinae Burmeister, 1850 from where he collected the fossils which corresponds to MTLA. Although this information is certain, the mate­ Parataxidea Z dansky, 1924 rial is described separately. Over all this time it is quite possible, that fossils found by the villagers in various sites Parataxidea maraghana (K it t l , 1887) on Samos were mixed with Prof. J. Melentis’ collection, (PI. 1, figs. 1-5) falsifying the origin of the material. More details about the localities and their stratigraphic position are given in Locality: Mytilinii-lA (MTLA), Adrianos ravine, My­ Kostopoulos et al. (this volume). tilinii Basin, Samos, Greece. Age: Middle Turolian, MN 12 (late Miocene); 7.1-7.0 Ma. Abbreviations: Material: Skull, MTLA-283; partial skull, MTLA-465; AMNH = American Museum of Natural History, New York m l sin, MTLA-7. AM PG = Athens Museum of Geology and Palaeontology, Measurements: The cranial measurements are given in University of Athens Table 2. BSPM = Bayerische Staatssammlung für Paläontologie und Historische Geologie, München. Description: HAL = Halmyropotamos, Evia Island, Greece Skull. The skull MTLA-283 (PI. 1, fig. 1) lacks the muz­ KER = Kerassia, Evia Island, Greece zle and the occipital region with the condyles; the right LGPUT = Laboratory of Geology and Palaeontology, University zygomatic arch and the bone in the posterior part of the of Thessaloniki braincase are broken. The specimen MTLA-465 is a MGL = Musée de Géologie de Lausanne MLN = Mytilinii-4, Samos, Greece M TLA-283 M T L A -465 MNHN = Museum Nationale d’Histoire Naturelle, Paris dex sin dex sin MRG = Maragheh, Iran MTL= Mytilinii-1, Samos, Greece LC — — 5.0 4.8 MTLA = Mytilinii-1 A, Samos, Greece BC — — 4.0 4.2 MTLB = Mytilinii-1B, Samos, Greece LP1 — — — — MTLC = Mytilinii-lC, Samos, Greece — MTLD = Mytilinii-1D, Samos, Greece BP1 — — — NHMA = Natural History Museum of Aegean, Mytilinii, Samos LP2 4.5 — 5.1 5.0 NHML = Natural History Museum of London BP2 3.6 — 3.5 3.7 NHMW = Naturhistorisches Museum, Wien LP3 5.7 5.6 5.7 5.6 NOW = Neogene Old World database PER = Perivolaki, Thessaly, Greece BP3 4.4 4.3 4.8 4.5 PIK = Pikermi, Attica, Greece LP4 8.1 7.8 8.5 7.8 PMMS = Palaeontological Museum of Mytilinii, Samos BP4 ant. 8.0 8.0 7.7 7.5 PXM = Prochoma, Axios Valley, Macedonia, Greece LM 1 buccal 7.8 7.7 7.6 7.4 QA = Quarry-A, Samos, Greece Ql = Quarry-1, Samos, Greece LM l lingual 11.0 11.6 11.3 11.2 Q2 = Quarry-2, Samos, Greece BM 1 11.4 11.2 11.2 11.2 Q4 = Quarry-4, Samos, Greece Q5 = Quarry-5, Samos, Greece MTLA-466 ml = 11.3 x 7; L tal = 5.4 RP1 = Ravin de la Pluie, Axios Valley, Macedonia, Greece RZl = Ravin des Zouaves-1, Axios Valley, Macedonia, Greece Table 2: Cranial dimensions of Parataxidea maraghana and RZO = Ravin des Zouaves-5, Axios Valley, Macedonia, Greece Promeles palaeattica. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

60 Beitr. Paläont., 31, Wien, 2009

MTLA-283 NHMW-MRG-TYPE

MTLA-465 -Ex.2 (ZDANSKY, 1924)

.1 vi— iu-A.

AMNH-FS-23011 CHINA-Ex.1 (ZDANSKY, 1924

LGPUT-PER-1280

Figure 1: Comparison of the upper dentition ofParataxidea and P rom eles. a. Tooth row of the skull, MTLA-283, b. Tooth row of the skull MTLA-465, c. P. m araghana, tooth row of the skull, AMNH-FS 23011, Samos; d. P. maraghana, tooth row of the TYPE, Maragheh; e. P. sinensis, tooth row of the skull Ex. 2, China (taken from Zdansky, 1924); f. P. crassa, tooth row of the skull Ex. 1, China (taken from Zdansky, 1924); g. Promelespalaeattica, tooth row of the skull LGPUT-PER-1280. All tooth rows shown as if from right-hand side. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 61

Pararaxidea maraghana, Upper Teeth

-Prpalaeattica, LGPUT-PER-1280 X MTLA-283 - = X — MTLA-465 P. maraghana, Samos, no270 (PILGRIM, 1931) P. maraghana, Maragheh, n=1 (KITTL, 1887) ■A- P. sinensis, China, n=2 (ZDANSKY, 1924) P. crassa, China, n=1 (ZDANSKY, 1924) Pr. palaeattica, Pikermi, n=3-4 (orig, meas.) Pr. macedonicus, Maramena, n=1-3 (SCHMIDT-KITTLER, 1995) Pr. smyrnensis, Turkey, n=1 (KAYA et al., 2005)

Figure 2: Logarithmic ratio diagram comparing the upper teeth ofParataxidea and P rom eles from various localities. partial skull lacking the posterior part and the zygomatic P3: It is similar to P2 but larger. The lingual cingulum is arches. The muzzle is short and relatively wide (PI. 1, fig. stronger and has a strong projection in its distal part. 3c). The nasal cavity is oval and its lateral borders are al­ P4: It is short and wide, having a triangular shape and most vertical to the palate. Its posterior border is situated strong protocone, while the paracone and metacone are above the canine. The frontals are narrow. The infraorbital relatively short. There is no parastyle, but there is a small foramen is large. The orbit is rounded and its anterior border cingular projection in the mesial base of the paracone, is above the middle of the P4. The zygomatic arches are giving the idea of a rudimentary parastyle (PI. 1, fig. 4). relatively strong and project buccally; they begin above the The protocone is robust, well separated from the paracone contact between the P4 and M l. The postorbital projection by a deep mesial valley and situated in front of the mesial is strong. A projection from the zygomatic arch to the pos­ border of the tooth. The metacone is blade-like, very short torbital projection almost closes the orbit; only a small part and faces buccally. There is a strong lingual cingulum of the orbit’s periphery remains open (PI.1 , fig. lb). There connecting the protocone and the metacone. There is an is no sagittal crest. The bullae are elliptical and their buccal elevation in the middle of the lingual cingulum, forming surface inclines roughly buccally, while the lingual surface a small cuspule (PI. 1, fig. 4) which gives a more squarish is more vertical. The anterior border of the bullae is level shape to the tooth. In the worn teeth the lingual cingulum with the posterior surface of the post-glenoid processes. The forms a crest connecting the protocone and metacone (PI. choanae are relatively wide and their anterior border is well 1, fig. 2). behind the M l, situated in the middle of the zygomatic M l: The shape of the tooth is trapezoid with the mesial arch, at about 8 - 9.5 mm behind the posterior border of the border perpendicular to the mesiodistal axis of the tooth. M l. The palate is elongated and narrow with almost parallel The posterior border has a deep groove with the lingual borders giving it a more or less rectangular shape. part projecting strongly distally. The paracone is larger and Upper Dentition. The skull M TLA-283 preserves the more buccally projected than the metacone. The protocone tooth rows P3-M 1 dex and P2-M 1 sin (PI. 1, figs 1, 2). is strong and situated in the center of the occlusal surface. The teeth are well preserved but worn, while the right M l The hypocone is strong and is extended to the distolingual is slightly broken at its buccal border. The skull M TLA- part of the tooth. There is a strong and elevated mesial, 465 preserves the tooth rows P4-M 1 dex and C -M l sin; lingual and distal cingulum. the teeth are little worn (PI. 1, figs 3, 4). Lower Dentition. The lower dentition is represented by C: It is elongated and slender, slightly curved distally only one isolated m l, M TLA-7 (PI. 1, fig. 5). and with a round transverse section. It seems to have a m l: It is short and wide with an elongated talonid. The mesial crest. paraconid is a large cuspid, situated mesially. The proto- P2: It is monocuspid, single-rooted with rounded crown. conid is equal in size to the paraconid. The metaconid is There is a slight cingulum all around the tooth. also large and situated level with the protoconid; both are ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

62 Beitr. Paläont., 31, Wien, 2009

XMTLA-7 ■ P. maraghana, Maragheh (KITTL, 1887)

• P. maraghana, Samos (PILGRIM, 1931) A P . sinensis, China (ZDANSKY, 1924)

♦ P. crassa, China (ZDANSKY, 1924) □ Pr. palaeattica, Pikermi (orig, meas.; ROUSSIAKIS, 2002)

O Pr. pallaeattica, Perivolaki (KOUFOS, 2006) A Pr. macedonicus, Maramena (SCHMIDT-KITTLER, 1995)

Figure 3: Scatter diagram length/breadth for the ml ofParataxidea and P rom eles from various localities.

separated by a small valley. The paraconid is distinguished directly compared with the material ofPromeles palaeat­ from the metaconid and protoconid complex by a deep tica from Pikermi, stored at NHML, and from Perivolaki valley, which is open lingually. The talonid is large and (Thessaly, Greece), stored at LGPUT and described by wide, being about half of the tooth’s length. The entoco- K oufos (2006a). The comparison allows recognizing the nid is large, situated behind the metaconid, from which differences between the generaPromeles and Parataxidea. it is separated by a very narrow valley. The hypoconid is The general shape and size of the skull is similar in the stronger than the entoconid and is separated from the two genera, but the comparison of the dentition gives the protoconid by a wide valley. The hypoconulid is large and following differences: crest-like, covering the whole distal border of the tooth. • the P2,3 ofParataxidea are more robust and have a strong There is another small cusplet behind the entoconid, con­ cingulum which is absent in Promeles (Fig. 1). necting it with the hypoconulid. Thus, the talonid is closed • the P4 of Parataxidea is robust (short and wide), all around and has a basin-like form (PI. 1, fig. 5c). There triangular-shaped and with a strong protocone, while in is a strong buccal cingulum. P. palaeattica it is slender (elongated and narrow), with a Discussion: The taxon was originally described as Meles weaker and more mesiolingually projected protocone. In maraghanus from Maragheh, Iran. The type specimen Promeles, the P4 lacks the small mesial cusp which cor­ is a right maxillary fragment with C-M l; the canine is responds to a rudimentary parastyle, as well as the small broken (Fig. Id), (Kittl, 1887; Taf. 15, fig. 4). Later, lingual cuspule developed in the middle of the lingual when Zdansky (1924) described some Chinese material cingulum (Fig. lg). and erected the new genusParataxidea, the species was • the M l is also completely different. InParataxidea it is transferred to it and reported as Parataxidea maraghana. larger, with an elongated lingual mesiodistal diameter. On A complete skull associated with the mandible is reported the contrary, the buccal mesiodistal diameter is shorter and as Meles maraghana in the list of Samos material given parallel to the alveolar line, while it is oblique inPromeles by Forsyth M ajor (1894) and is stored at M GL. This (Fig. 3g). The lingual cingulum is stronger in Parataxidea. skull was described by Pilgrim (1931). The taxon is also On the mesial border of the M l, there is a strong groove reported from Q l and QA of Samos as Parataxidea ma­ in Promeles which is absent in Parataxidea (Fig. 1). raghana (Solounias, 1981), while later it is referred to as • the m l of Parataxidea is larger, more robust with shorter P polakHmaraghana (Bernor et al., 1996). Some material and narrower talonid, which is deeper than that ofProme­ of P. maraghana from Samos, coming from Q l is stored at les. The paraconid of the Parataxidea m l is shorter than that AM N H (http://paleo.amnh.org/fossil/list.html). of Promeles. There is strong buccal cingulum in Parataxidea, The genus Parataxidea resembles Promeles and can be which is absent in Promeles. easily confused with it, but a more careful comparison • The dental size ofPromeles is generally smaller than that clearly distinguishes the two genera. The Samos skulls are of Parataxidea, the Promeles teeth are especially narrower ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 63

than those of Parataxidea (Figs 2, 3). These morphologi­ Another partial skull with a mandible from the Lok. 30 of cal and metrical differences clearly distinguish the genera China are described as Parataxidea crassa (Zdansky, 1924; Parataxidea and P ro m eles. The morphology and dimen­ taf. XI, figs. 1-4). The dentition of the skull is quite worn sions of the studied material from Samos fit quite well (Fig. If), but its morphology is similar to that of MTLA- with those of Parataxidea and can be certainly ascribed 283, 465. Especially the teeth of MTLA-283, which to this genus. are worn, closely resemble those ofP. cra ssa .The dental The type specimen ofParataxidea maraghana has worn dimensions ofP. cra ssa are slightly larger than those of the teeth (Fig. Id). The species is characterized by a large studied skulls. The most significant difference is the size of infraorbital foramen situated above the P4. The premolars the P4, whose length is smaller than that ofP. cra ssa; the are monocuspid, single-rooted with strong cingulum and other teeth have similar dimensions (Figs. 2, 3). Bearing a lingual cingular projection in the P3. The P4 is short in mind that the available material from both localities and wide with a large protocone well-separated from (M TLA and Lok. 30) is rare, the dental size variability is the paracone; there is no parastyle, but a mesial cingular not well-known and with their morphological similarity, P. projection while the lingual cingulum is strong. The M l cra ssa and P. maraghana could be synonyms. Unfortunately, is trapezoid with a large paracone and a smaller metacone; I have not seen the skull ofP. cra ssa from China and thus its lingual anteroposterior diameter is remarkably longer it is difficult to ascertain such a conclusion. than the buccal one (Fig. Id). This morphology fits in quite The genus Parataxidea is mainly an Asian taxon known well with the studied skulls from M TLA, which can be from Middle Maragheh and Samos. The species is also ascribed to P. maraghana with certainity. Moreover, the known from the locality Kara5ahasan (Turkey), dated to size of the upper teeth of MTLA-283, 465 is very close early Turolian, MN 11 (NOW, 2007). Bearing in mind, to P. maraghana (Fig. 2). The studied lower carnassial is that Middle Maragheh and M T L A are dated to middle also similar to that of P. maraghana but slightly smaller Turolian (MN 12) then the stratigraphic distribution of (Fig. 3). The upper canine of MT LA-465 is smaller than P. maraghana covers early - middle Turolian. that of P. maraghana (Fig. 2); this difference could be due to sexual dimorphism. The canine of the type skull Family Hyaenidae G ray, 1821 is even larger than that ofP. cra ssa, which has the largest teeth (Fig. 2). Genus Plioviverrops K retzoi, 1938 The Samos skull and mandible (M GL-SAM -no 270), reported byF orsyth M ajor (1894) and described by Pil­ Plioviverrops orbignyi (Gaudry & Lartet, 1854) grim (1931), is an aged individual with almost completely (PI. 2, figs. 1, 2) worn P4, M l, p4 and m l, 2. The characterisitcs of this skull, as well as its dental morphology given byP ilgrim Locality: Mytilinii-lB (MTLB), Adrianos ravine, My- (1931), fit in quite well with those of the studied skulls. The tilinii Basin, Samos, Greece. upper teeth and m l dimensions are also close to those of Age: Middle Turolian, MN 12 (late Miocene); 7.1-7.0 Ma. the studied material, although they are slightly larger (Figs Material: Skull, MTLB-170. 2, 3). The upper canine of the MGL skull is remarkably Measurements: The measurements of the skull are given larger than that of MTLA-465, but this is probably due to in Table 3. sexual dimorphism. The studied material is also compared with some photos of the material stored at AM NH. The Description: AMNH material includes a skull from Samos connected Skull. The skull is almost complete, lacking the incisor’s with the mandible (AMNH-FS-20700), a maxillary frag­ border and the right zygomatic arch. The narial opening is ment with C-M ldex (AMNH-FS-23011), (Fig. lc), and a slightly distorted on the distal border; it is oval, relatively mandibular fragment with il-p3 sin (AMNH-FS-23006). wide, with its borders running parallel to the canine’s root. The skull AM N H -FS-20700 lacks the braincase, but its The nasal bones are absent (PI.2 , fig. la). The frontal bones frontal part is very similar to that of the studied skulls. The are extended to the front, like spines, at about the distal upper teeth of the specimen AM N H -FS-23011 (Fig. lc) border of the narial opening. A groove is running across are well-preserved and their morphology is quite clear and the contact of the frontals. The orbit is elliptical and large, fits in well with that of MTLA-283, 465 (Figs. la-c). with its anterior border above the middle of the P3. The Two skulls associated with the mandible from Lok. 305 of China are described as Parataxidea sinensis (Zdansky, dex sin dex sin 1924; Taf. X, figs 5-10). The cranial and dental morpho­ 5.0 x 4.0 6.6 x 4.3 Ml — x8.4 6.2 x 8.3 logical characteristics of P. sinensis (Fig. le) are similar c P3 7.7 x 4.7 76x4.6 M2 4.6 x 6.3 4.4 x 6.4 to those of M TLA-283, 465. The dental similarity of P. P4 — x7.1 10.8x7.0 C-M2 46.5 46.8 sin en sis to the type ofP. maraghana is reported byZ dansky (1924) too. Although they are morphologically similar, FP4bbde — x 3.8 C-P4 38.0 38.1 P- sinensis has smaller dimensions (Figs. 2, 3). The upper C-Ml 43.0 43.4 teeth of the studied skulls M TLA-283, 465 and the type of P. maraghana are significantly larger than those ofP. Table 3: Cranial dimensions of Plioviverrops orbignyi from vari­ sin en sis (Fig. 2), and clearly distinguished from them. ous Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

64 Beitr. Paläont., 31, Wien, 2009

Figure 4: Comparison of the upper dentition ofPlioviverrops orbignyi from various Greek localities. a. Tooth row of the skull MTLB-170;b. Tooth row of the skull NHMW-A.4750, Samos; c. Tooth row of the skull MNHN-PIK-3016, Pikermi; d. Tooth row of the skull MNHN-PIK-3022, Pikermi, TYPE; e. Tooth row of the maxilla LGPUT-PER-1, Perivolaki, Thessaly; f. Tooth row of the skull LGPUT-PER-1295, Perivolaki, Thessaly; g. Tooth row of the skull LGPUT-VTK-35, Axios Valley. A ll specimens shown as if from left-hand side. infraorbital foramen is large situated in front of the orbit inclined distal part, a arge buccal fossa and a slight groove and above the P3. The postorbital constriction is strong in their mesiolingual surface. The zygomatic arches are while the postorbital projections are partially broken, but strong, projecting buccally. The palate is relatively short seems to be large. A quite strong sagittal line starts from and wide. It is narrow between the canines and the first each postorbital projection and faces posteriorly. The two premolars, but it widens from the mesial border of the P3, sagittal lines are connected at about the beginning of the reaching its maximal breadth at the distal border of the parietals, forming a sagittal crest which ends at the external carnassial (PI. 2, fig. 2). The anterior border of the choanae occipital protuberance (PI. 2, fig. la). The occipital bone is behind the posterior border of the M2. is triangular-shaped with a strong crista nuchae which Upper Dentition. The skull preserves the tooth rows C, projects distally, exceeding the level of the occipital con­ P3-M2 on both sides. The right P4 and M l are partially dyles (PI. 1, fig. le). The occipital condyles are strong and broken, while the left P3 is somewhat inside the bone and the foramen magnum is elliptical. The glenoid fossa is well- thus not clearly shown (PI. 2, fig. 2). developed with strong post-glenoid processes (PI. 2, figs, C: The tip of both canines is broken. They are relatively lb, c, e). The auditory bullae are ovoid with a backwardly strong, flattened buccolingually and having an elliptical ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 65

Plioviverops orbignyi, Upper Teeth

■P. orbignyi, Pikermi, Greece, MNHN-PIK-3022, HOLOTYPE > P. orbignyi, Mytilinii-IB, Samos, MTLB-170

■P. orbignyi, Pikermi, Greece, MNHN-PIK-3016 P. orbignyi, Samos, Greece, NHMW-A.4750

■P. orbignyi, Vathylakkos-3, Greece, n=2 (KOUFOS, 2000) P. orbignyi, Perivolaki, Greece, n=3-4 (KOUFOS, 2006)

■P. faventinus, Brishigella, Italy, n=2-6 (TORRE, 1988)

Figure 5: Logarithmic ratio diagram comparing the upper teeth ofPlioviverrops from various localities.

cross-section. They are curved distally and they are point­ Discussion: The species P. orbignyi is well-known from ing buccally. Greece, found in several late Miocene localities. Origi­ P3: It is short and wide, lacking the anterior accessory nally it was described from Pikermi as Viverra orbignyi cusp. A strong crest begins from the top of the main by G audry & L artet (1856). Later, it was transferred to cusp, running across its mesial border and ending at the Thalassictis by G audry (1861), to Ictitherium by G audry mesiolingual corner of the tooth. There is a small posterior (1862-67) and finally to Plioviverropsby K retzoi (1938). accessory cusp on a distal projection of the distal cingulum. Then afterwards, it was transferred to this taxon (Beau­ The cingulum is strong all around the tooth. In the middle mont, 1969; Beaumont & M ein, 1972; Koufos, 1979, of the lingual cingulum there is a strong projection giving 1980,2000,2006a; Solounias, 1981; Bonis & Koufos, the idea of a rudimentary protocone; this feature indicates 1991; W erdelin & S olounias, 1991; Roussiakis, 1996). a relatively molarized premolar. The type material is housed at MNHN and includes two P4: The protocone is relatively large, well separated from skulls (MNHN-PIK-3022, 3016) associated with the the paracone and projects lingually. Its mesial border is mandible. I have seen this material and it is character­ projected in front of the parastyle. The parastyle is smaller ized by a relatively robust carnassial with a strong lingual than the protocone and paracone and it has a small cin- cingulum; the P3 has a strong projection of the lingual gular projection at the mesiobuccal corner. The metacone cingulum, being quite molarized. In these features the is blade-like and strongly curved buccally. The lingual studied skull MTLB-170 resembles the type material from cingulum is well-developed and continues with a crest Pikermi (Fig. 4). There are several skulls ascribed toP. to the top of the protocone. One weak crest starts from o r b ig n y i from various localities (Tab. 3), but their metrical the tip of the paracone, and running through the valley comparison is questionable as most of them are deformed between the paracone and protocone, ends at the tip of the and distorted, making the accurate measurement of their protocone. The buccal cingulum is weak. dimensions difficult. Nevertheless, the size ofM TLB-170 M l: It is triangular with three cuspids. The paracone is is similar to that of the two skulls from Pikermi; its veri­ stronger than the metacone, from which it is distinguished fied dimensions coincide with those of the type material by a deep valley. The paracone and protocone are similar in (Tab. 3). The comparison of the teeth shows better results height, and they are connected by a crest running across the and indicates a general metrical similarity between them. mesial border of the tooth. Another weak crest begins at However, the teeth ofM TLB-170 are slightly larger than the tip of the protocone connecting it with the base of the those of the type material from Pikermi (Fig. 5). metacone. There is a well-developed buccal cingulum. The species is reported by a skull, some isolated teeth and M 2: More rounded than M l with the paracone stronger postcranials from Adrianos ravine in the list ofF orsyth than the metacone. The protocone is similar to that of Ml. M ajor (1894) as Ictitherium orbignyi, but without descrip­ The central basin is large and open distally near the base of tion. The skull was described by Beaumont (1969) and the distobuccal cingulum. There is a strong buccal cingulum. later by Solounias (1981). Another skull from Samos ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

66 Beitr. Paláont., 31, Wien, 2009

MAMMAL LOCALITIES MAMMAL LOCALITIES MAMMAL LOCALITIES WITH P ORBIGNYI WITH P. GUERINI WITH P. FAVENTINUS

LA GLORIA-4

LA GLORIA

BRISHIGELLA LA ALBERCA

ALEZAR B

CERRO DE LA GARITA CONCUD PIKERMI LOS MANSUETOS MYTILINII-1B LOS ALJEZARES — PERIVOLAKI >VATHYLAKKOS-2,3 PROCHOMA

PUENTE MINERO PIERRA GREVILLENTE-2

RAVIN DES ZOUAVES-5

The arrow indicates the Greek LOS VALLES DE localities with GPTS age FUENTINUEVA

Figure 6: Stratigraphic distribution of the genus Plioviverrops during late Miocene of . The data was taken from the biblio­ graphy and NOW (2007).

(NHMW-SAM-A.4750) is housed in Vienna; its direct curved metastyle-blade are characters fitting in well with comparison to the studied one indicates that it is mor­ those of M TLB-170 (Fig. 4). The dental dimensions of phologically and metrically similar to M TLB-170 (Tab. NHMW-SAM-A.4750 seem to be slightly smaller than 3). The absence of an anterior accessory cusp, the small the type material from Pikermi and smaller than those of posterior cusp and the strong lingual cingular projection MTLB-170 (Fig. 5). of the P3, as well as the robust P4 with well-separated Several specimens of Plioviverrops orbignyi are known from protocone, strong lingual cingulum and buccally strongly the localities of Axios Valley (Macedonia, Greece). It was ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 67

found in the Turolian localities of Ravine des Zouaves-5 Sc Petter, 1969). A mandibular fragment from the local­ (RZO), Vathylakkos-3 (VAT) and Prochoma-1 (PXM) ity VAT of Axios Valley (Macedonia, Greece) is referred with cranial and mandibular remains (Koufos, 1979, to as P. cf. guerini because it has a rudimentary posterior 1980,2000; Bonis & K oufos, 1991). The RZO material accessory cuspid in p2,3 and a slightly larger size (Bonis contains only mandibular remains and the comparison Sc Koufos, 1991). However, the size of the teeth and the with the studied skull is impossible. A partially preserved development of the cuspids varies in the material ofP. skull with maxilla is known from Vathylakkos-3 (LGPUT- orbignyi, and in the re-examination of the taxonomic value VAT-135) described byB onis & Koufos (1991). A direct of P. guerini, it is necessary to all the new material comparison with M TLB-170 suggests that both skulls and data in mind (Koufos, 2006a). The Late Turolian P. have similar cranial and dental morphological features and faventinus is known from Italy (Torre, 1988) and differs dimensions (Tab. 3, Fig. 4). Although the dimensions of from P. orbignyi in having highly molarized premolars, a the skulls are similar, the teeth of LGPUT-VAT-135 are marked posterior accessory cusp in P3 and a remarkably smaller than those of the studied skull (Tab. 3, Fig. 5). The larger size (Fig. 5). skull from Prochoma (LGPUT-PXM-251) preserves the The oldest appearance of the genusPlioviverrops is known opisthocranium without the maxilla (Bonis &c Koufos, from the locality of Laugnac (France) with the speciesP. 1991) and its available dimensions are close to those of collectus, dated to late Agenian, MN 2 (NOW, 2007). P. MTLB-170 (Tab. 3). gervaisi is known from the locality of Vieux Collonges Recently P. orbignyi was found in the middle Turolian (France), dated to late Orleanian, MN 4-5, and the spe­ locality of Perivolaki, PER (Thessaly, Central Greece), a cies P. gaudryi is known from the locality of La Grive/St. complete skull associated with the mandible (LGPUT- Alban, France, dated to late Astaracian, MN7/8 (NOW, PER-1295), and one frontal part of the skull associated 2007). Plioviverrops sp. is mentioned from the early with the mandible, (LGPUT-PER-1), (Koufos, 2006a). Vallesian (MN 9) locality of Los Valles de Fuentinueva, The direct comparison of this material to MTLB-170 Spain (NOW, 2007). However, the maximum spread of suggests that the cranial and dental morphology of both the genus is observed in the Late Miocene, where it is very samples fit together quite well (Fig. 4). The metrical com­ common, especially in the Turolian (Fig. 6). Two species parison also indicates that the cranial size is similar (Tab. are mentioned: P. orbignyi, known from the Eastern Medi­ 3), while the teeth of the Perivolaki material are slightly terranean (mainly Greece, but it is also known from the smaller than those of MTLB-170 (Fig. 5). Spanish locality of Los Aljezares), and P. guerini, known The small size differences in the cranial dimensions ofP. from Spain. A form similar to P. guerini is known from orbignyi from the various localities can be explained by the Middle Turolian (MN 12) locality of Vathylakkos-3 deformation or distortion of the material. The differences (Koufos, 2000). The large-sized P. faventinus appeared in the teeth could either be due to sexual dimorphism or in Italy at the end of Miocene, but is also recognized in to the different geological age of the material(K oufos, the Pliocene of Spain (Fig. 6). 2006a). As is mentioned above, the dental dimensions of MTLB-170 are smaller than those of LGPUT-VAT-135 Genus Protictitherium K retzoi, 19 24 and LGPUT-PER-1, 1295. The age of these localities is very similar as all are middle Turolian. The magne- Protictitherium crassum (D eperet, 1892) tostratigraphic record indicates an age of 7.1-7.0 Ma for (PI. 3, figs. 1, 2) M ytilinii-lA, ~7.5 Ma for Vathylakkos and 7.3-7.1 Ma for Perivolaki (Sen et al., 2000; Kostopoulos et al., 2003; Locality: M ytilinii-4 (MLN), Mytilinii Basin, Samos, Koufos et al., 2006). This age similarity strengthens the Greece. first hypothesis of sexual dimorphism. The prevalence of Age: Middle Turolian, MN 12 (late Miocene); ~7.5 Ma. this hypothesis is supported by the presence of some man­ Material: Mandible, MLN-42; isolated lower canine, dibles from RZO and PER, which have larger teeth than MLN-26. the type material from Pikermi; however, these mandibles Measurements: The measurements of the mandible are are also associated with larger canines than those of the given in Tab. 4. Pikermi material and probably indicate male individuals (Koufos, 2006a). MLN-42 dex sin dex sin Besides P. orbignyi, the genus includes other species known Lc 7.0 7.2 Lp3 11.5 11.0 mainly from Spain, France and Italy. The Early-Middle Be 5.5 5.4 Bp3 4.6 4.5 Miocene P. gervaisi, known from France, differs from P. Lpl — 2.7 Lp4 12.0 — orbignyi by being smaller, having avery narrow P3 and a Bpl — 2.0 Bp4 5.3 5.0 protocone higher than the parastyle in P4 (Beaumont Lp2 9.5 9.4 Lml 15.5 15.6 &M ein, 1972; Petter, 1976). The Middle Miocene P. Bp2 4.0 3.9 Bml 6.9 6.7 gaudryi has a significantly smaller size than P. orbignyi Ltrig 10.4 10.6 (Beaumont & M ein, 1972). Some mandibular remains of MLN-26 c = 7.2 x 5.3 the Turolian P. guerini were found in Spain. It differs from the studied one in having a larger m l with a longer trigonid Table 4: Mandibular dimensions of various Protictitherium and a posterior accessory cuspid in the p2,3 (Crusafont species, in mm. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

68 Beitr. Palaont., 31, Wien, 2009

Figure 7: Comparison of MLN-42 with various ictitheres. a. Right and left tooth row of the mandible MLN-42; b. Protitctitherium aegaeum, tooth row of the mandible IKE-46, Turkey (taken from K aya et ah, 2005); c. Protictitherium crassum, tooth row of the mandible LGPUT-DKO-20, Axios Valley, Macedonia, Greece; d. Protictitherium cf. crassum, tooth row of the mandible LGPUT-PNT-68, Pentalophos-1, Axios Valley, Macedonia, Greece; e. Protictitherium crassum, tooth row of the mandible LGPUT-XIR-2, Xirochori-1, Axios Valley, Macedonia, Greece; f. Protictitherim crassum, Sofpa, Turkey; g. Protictitherium llopisi, tooth row of the Can Bayona mandible, Spain (cast); h. Protictitherium arambourgi, tooth row of the mandible MNHN-TRQT015, Sinap, Turkey; j. Hyaenictitherium intub erculatum, tooth row of the mandible MNHN- TRQG013, Sinap, Turkey. All specimens shown as if from left-hand side.

Description: inferior border of the mandibular ramus is slightly curved. Mandible. The studied mandible consists of the two man­ i3: It is small, not canine-like, with a strong buccal projec­ dibular rami of the same individual; the right mandibular tion, giving it a triangular shape. ramus preserves the tooth row c, p2-ml and the left one c: It is relatively strong, high, strongly curved backwards, the tooth row i3-m l (PI. 3, figs. 1, 2). Both mandibular with a pointed tip. Its lingual surface is flattened and has fragments lack the ascending ramus. The mandibular cor­ two crests running across its mesiolingual and distolingual pus is relatively deep and thin, with a large mental foramen border. below the anterior root of the p2. The symphysis is elon­ pi: It is very small, monocuspid and single-rooted. gated and narrow, and inclines gradually backwards. The p2: It is elongated and narrow. It lacks an anterior accessory ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 69

Protictitherium crassum, Lower Teeth

■P. orbignyi, MNHN-PIK-3022 TYPE —X — MLN-42

-P. aegaeum, Esendere IKE-46 (KAYA et al., 2005) M P. crassum, Xirochori, Axios valley, XIR-2 -P. crassum, Dytiko-3, Axios valley, LGPUT-DKO-20 —♦ — P. cf. crassum, Pentalophos, Axios valley, LGPUT-PNT-1, 68 -P. crassum, Can Bayona (CRUSAFONT & PETTER, 1969) —A— P. crassum, Sofga, BSPM-1968/758 (SCHMIDT-KITTLER, 1976) -P. arambourgi, Sinap (orig. meas.)

Figure 8: Logarithmic ratio diagram comparing the lower teeth of the various species ofProtictitherium.

cuspid but there is a mesial cingular projection which is France (D eperet, 1892). The holotype is characterized by connected with the top of the main cuspid by a crest. The a highly molarized p4 and a carnassial with a high meta­ posterior accessory cuspid is well distinguished, situated conid. Both characterisitcs exist in the studied mandible slightly buccally on a small distal projection of the distal M LN -42. In Greece P. crassum is known from several late cingulum. Miocene localities of Axios Valley (Macedonia, Greece). p3: Similar morphology to p2, but the distal part of the A right hemimandible with c-p3-m2 (DKO-20) is known tooth is larger and more talonid-like. from the late Turolian locality of Dytiko-3 (Koufos, 1980, p4: The anterior part of the left p4 is slightly broken (PL 3, 2000; Bonis & Koufos, 1991). The dental comparison of fig. lc). It is relatively short and narrow with a small ante­ this specimen with MLN-42 indicates similar morphology rior accessory cuspid and a large distal one, like a talonid (Fig. 7) but the premolars of DKO-20 are relatively shorter (PI. 3, fig. 2c). The posterior accessory cuspid is situated and wider than those of MLN-42 (Fig. 8 ); the robusticity buccally, while a smaller one is developed lingually and a index differs remarkably from those of DKO-20 (Tab. 4). cingular elevation closes the distal border of the talonid. The carnassial has a similar morphology in both specimens This morphology closely resembles that of the carnassial’s (Fig. 7). Although the carnassial of DKO-20 is worn, talonid, indicating high molarization. the metaconid and entoconid are high and this feature m l: It is relatively short and wide, with a strong and wide resembles M LN -42, but those of M LN -42 seem to be tricuspid talonid (PI. 3, figs. 1, 2). The cuspids are high. higher and more slender. The talonid in both specimens is The paraconid and protoconid are blade-like; the pro- large, wide and well separated from the trigonid by a deep toconid is higher than the paraconid. The metaconid is valley. However, the ml of MLN-42 is larger than that well-developed and high, reaching the two-thirds of the of DKO-20 (Fig. 8 ). A mandible with c-p3-ml-m3 dex protoconid. The talonid is distinguished from the trigonid and c-p2-m2 sin (XIR-2) of P. crassum has been described by a wide valley. The entoconid is the highest cuspid of the from the late Vallesian locality Xirochori-1 (XIR) of Axios talonid. The hypoconulid is smaller and connected to the Valley, Macedonia, Greece (Fig. 7e),(K oufos, 2000). The base of the hypoconid by a crest, closing the posterior end dental morphology of XIR-2 resembles that of MLN-42 of the talonid. There is a strong buccal cingulum. but there are some differences. The premolars of XIR-2 Discussion: The studied mandible is large and has rela­ are relatively shorter and wider; their robusticity index is tively long and narrow premolars, as well as high cuspids in smaller than that of MLN-42. The p2, 3 of XIR-2 lacks the carnassial, characteristics which suggest a large-sized an anterior accessory cuspid like M LN-42, but they have Protictitherium. Such a large-sized Protictitherium is P. a smaller posterior accessory cuspid than that of MLN -42 crassum and the recently describedP. aegaeum. (Figs. 7, 8 ). The p4 of XIR-2 has a smaller anterior and P crassum is known from various Eurasian localities. It posterior accessory cuspid and wider distal cingular projec­ was originally described asHerpestes crassus from La Grive, tion than those of M LN-42. The carnassial of XIR-2 is ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

70 Beitr. Paläont., 31, Wien, 2009

Protictitherium crassum, Lower Teeth

------P. orbignyi, MNHN-PIK-3022 TYPE ---- X ---- MLN-42 — A — P. aegaeum, Esendere, IKE-46 (KAYA et al., 2005) -P. arambourgi, Sinap (orig. meas.) — □ — I. viverrinum, Pikermi, mean (WERDELIN, 1988) -I. viverrinum, China, mean (WERDELIN, 1988) □ I. viverrinum, Pikermi, min (WERDELIN, 1988) •I. viverrinum, Pikermi, max (WERDELIN, 1988) 0 I. viverrinum, China, min (WERDELIN, 1988) •I. viverrinum, China, max (WERDELIN, 1988) — • — H. intuberculatum, Sinap (orig, meas.)

Figure 9: Logarithmic ratio diagram comparing the lower teeth of some Protictitherium species with Ictitherium viverrinum and Hyaenictitherium intuberculatum.

smaller and with relatively lower cuspids than M LN-42 llopisi (W erdelin 8 cS olounias, 1991). The premolars of (Figs 7, 8 ). A mandibular fragment with p3-m l sin (PNT- M LN -42 are longer and narrower, having more slender 68 ) of P. cf. crassum is known from the Vallesian locality features than those ofP. crassum from Can Bayona. Their of Pentalophos-1 (PNT) of Axios Valley, Macedonia, morphology is more or less similar, but the cingulum Greece (Bonis 8 c K oufos, 1991; Koufos, 2000). The is more developed in the premolars and there is a small premolars of both specimens, PNT-68 and MLN-42, have anterior accessory cuspid in the p3 of the Can Bayona a similar morphology with high molarization, especially teeth. The carnassial has a similar morphology, but the in the p4 (Fig. 7). However, they are more robust and the metaconid of M LN -42 is slightly higher. The talonid of anterior accessory cuspid in the p4 of PNT-68 is stronger the Can Bayona m l is broken buccally, but it has a high than that of MLN-42. The carnassial morphology is also entoconid. The size of MLN -42 is visibly larger than that similar in both specimens, with a high metaconid and of Can Bayona. entoconid. Although they are morphologically similar, A mandible (IKE-46) of a large-sized Protictitherium was the size of PNT-68 is significantly smaller than that of recently described from the locality of Esender, Turkey, M LN -42, as well as smaller than all the other specimens as a new species, P. aegaeum (Kaya et al., 2005). The of P. crassum (Fig. 8 ). studied mandible M LN -42 is compared with a cast of this Some material ofP. crassum is also known from Asia Minor species. The premolars of IKE-46 are generally shorter (Schmidt-K ittler, 1976). The M LN -42 is compared and wider than those of MLN -42, having a more robust with a cast of P. crassum from the locality of Sof^a, which is aspect (Tab. 5, Fig. 8 ). In this characteristic they are closer a mandibular fragment with p2-ml sin (BSPM-1968/758). to those of P. crassum. The p2, 3 of MLN-42 differ from The teeth of BSPM-1968/758 are smaller than those of P. aegaeum in lacking the anterior accessory cuspid. The MLN-42, but the morphology of the premolars is simi­ posterior accessory cuspid of p2 is stronger in M LN-42 lar (Figs. 7, 8 ). The premolars of the Sof£a specimen are and not circled by an elevated cingulum, as inP. aegaeum. relatively elongated and narrow, like those of M LN-42. The posterior accessory cuspid of the p3 has a similar size The carnassial is different in having a smaller and lower in both specimens, but there is a more elevated distal metaconid, a relatively smaller talonid with a smaller and cingular projection in P. aegaeum. The p4 of P. aegaeum is less high entoconid and hypoconid. shorter and broader than that of M LN -42 (Fig. 8 ) with P. crassum is also known from Spain, and M LN -42 is a stronger anterior accessory cuspid and a lingually less compared with a cast from the Vallesian locality of Can elevated distal cingular projection (Fig. 7). In this feature, Bayona housed at LGPUT; the material was originally M LN-42 is closer to P. crassum. The lower carnassial of described as P. crassum llopisi (Crusafont 8 c P etter, both specimens has similar a size, which is very close to 1969) and was later considered as a separate species, P. the minimum values for Ictitherium viverrinum (Fig. 9). ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 71

Species Reference Lp2/Bp2 Lp3/Bp3 Lp4/Bp4 Lml/Bml

P crassum MLN-42 239 247 233 229 P crassum LGPUT-XIR-2 189 197 200 220 P. crassum LGPUT-DKO-20 — 215 200 216 P. crassum Can Bayona 209 213 219 — P crassum BSPM-1968/758 221 233 205 198 P. cf. crassum LGPUT-PNT-1, 68 — 205 216 216 P. aegaeum IKE-46 198 228 197 216 P. aram bourgi MNHN-TRQ4L015 226 214 181 215 I. v iv errin u m mean, Pikermi (W erdelin , 1988) 188 203 199 227 I. viverrinum mean, China (W erd elin , 1988) 189 204 196 229 H. intuberculatum MNHN-TRQT013,1014 218 224 195 209

Table 5: Robusticity Indices of the lower teeth of variousProtictitherium species.

In spite of their metrical similarities, the carnassial ofP. a right hemimandible with p2-ml, stored at MNHN. The aegeum differs morphologically in the talonid. The talonid direct comparison of P. arambourgi with MLN-42 sug­ of M LN-42 is stronger, with a similarly high, but more gests that it differs in having a smaller m l with a relatively robust entoconid. Similar differences are also observed in shorter talonid, more robust premolars, a stronger anterior the size of the metaconid. The hypoconid and hypoconulid accessory cuspid in the lower premolars and a wider distal of MLN-42 are stronger than those ofP. aegaeum. But the cingular projection. Thus, the studied material is clearly most significant difference between the two specimens is different from P. arambourgi. the strong entoconid of MLN-42. Moreover, the buccal Another ictithere from Sinap (Turkey) was described by cingulum of the talonid of P. aegaeum is very strong; in O zansoy (1965) as Ictitherium intuberculatum and later MLN-42, the cingulum at this point is faint. Although transferred to Hyaenictitherium (Bonis, 2004). A maxillary there are similarities between M LN-42 andP. aegaeum, fragment with P2-M 1 and a mandible are known from this there are also some differences which distinguish them. species which are stored at MNHN. It is quite larger than The problem with these small hyaenids is how these dif­ M LN-42, with a relatively lower metaconid and entoconid ferences can be explained: are they intraspecific differences in the m l. The premolars have a similar morphology to or are they related to palaeoecology or geography? those of M LN -42, an absent or weak anterior accessory Considering the presence ofP. crassum in Eurasia, the cuspid in p2, 3 but the p2 of MLN-42 is relatively more rarity of the material and its startigraphic and geographic elongated, having a stronger mesial cingular projection. dispersion are given as the main reasons for providing However, the greater size ofH. intuberculatum clearly difficulties in the comparisons and the explanation of distinguishes the studied material from it. their relationships (Koufos, 2000). This is also clear from Bearing in mind all the above-mentioned comparisons, the comparison of MLN -42 and P. aegaeum. Both are of M LN -42 is morphologically closer to P. crassum, to which similar age (Turolian), geographically close, but have some it has some differences which could be considered as in­ differences. The question is if these differences are quite traspecific. Thus, the specimen M LN-42 is assigned toP. strong enough for a specific or subspecific distinction. As crassum at the moment, waiting for more material which was mentioned above, the size of the m l of M LN-42 is will allow: a) a better comparison with the known material, near the minimum values forIctitherium viverrinum from and b) a better knowledge of its morphology. Although the Pikermi and China (Fig. 9). However, the comparison known material from the late Miocene deposits of Samos of MLN-42 with I. viverrinum from Pikermi and Axios is very rich, this taxon was unknown and this is the first indicates that the premolars of MLN -42 are remarkably evidence of its presence in the Samos fauna. smaller and more slender than those ofI. viverrinum, and the cuspids of the m l in I. viverrinum are relatively lower. Genus Hyaenictitherium K retzoi, 1938 These features and especially the size clearly distinguish MLN-42 from I. viverrinum . Hyaenictitherium wongii (Zdansky, 1924) Ozansoy (1965) described a hemimandible of a small (PI. 3, fig. 3; Pis. 4-5; PI. 6, figs. 1, 2) ictithere from Sinap (Turkey) asIctitherium arambourgi', he states that it is larger than Plioviverrops orbignyi and smaller Localities: Mytilinii-lA, B, C (MTLA, MTLB, MTLC), than Ictitherium intuberculatum and Hyaenictitherium “hip- Adrianos ravine, Mytilinii Basin, Samos, Greece; Mytili- parionum” (= wongii). Later, Schmidt-K ittler (1976) nii-4 (MLN), Potamies ravine, Mytilinii Basin, Samos, synonymized this taxon with Protictitherium crassum, Greece. while even later on, W erdelin &c S olounias (1991) Age: Middle Turolian, MN 12 (late Miocene). transferred it to Protictitherium and kept the specific name M ytilinii-lA (M TLA, B, C): 7.1-7.0 Ma. of Ozansoy (1965). The latter opinion was also accepted Mytilinii-4 (MLN): ~7.5 Ma. by Bonis, (2004). The type specimen ofP. arambourgi is Material: ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

72 Beitr. Paláont., 31, Wien, 2009

Tableó: Cranial dimensions of Hyaenictitherium wongii from various Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 73

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 [36] 11.0 34 .0 25.0 29.0 29.0 MNHN- S L Q .-930

1 1 1 1 1 1 1 1 1 [29] 67.0 13.0 31.0 29.5 33.0 32.0 36.0 SH 30 .0 143.0 o 154.0

MNHN- S L Q .-929 o

i 1 1 1 1 1 1 1 1 61.3 13.3 27.0 35.2 23.7 2 2 .7 2 8 .4 26 .0 2 4 .0 O 136.5 V A T -100 LGPUT- u* \ x i o s v alley

O & 1 1 1 1 i 1 1 1 1 1 1 1 1 1 1 1 23.0 29.8 26 .5 28 .5 R Z O - 10 L G P U T -

1 1 1 1 1 1 1 1 1 1 1 1 1 37.0 13.6 25.5 23 .0 2 3 .2 28 .8 2 4 .0 2 2 .6 R Z 1 -2 LGPUT- 1

1 1 1 1 1 1 1 1 1 TT 1 1 1 12.5 49.8 24.5 2 4 .0 1 25.0 26 .5 62.0 42 .2 n n -2 00 SAM- c NHMW- <ü a bO V S 3 1 1 1 1 1 1 1 1 1 1 1 11.0 27.0 32.5 25.0 25.0 45.3 2 3 .0 63 .0 86 .2 50.3 nn-1 SAM-

NHMW- rt

S am o s E o 1 1 1 1 o 1 1 1 1 1 1 1 1 18.5 49.5 61.8 25.0 49.5 rt 2 2 .3 (N 2 0 .5 44 .0 SAM-

A .4743 C MHNW-

o 00 o o 1 1 1 1 1 7.8 1 1 1 1 19.0 Ö 32 .0 20.0 61.2 42.7 49.0 24.3 24.5 2 0 .6 00 SAM- A .4745 NHMW- Z 9 0

18 1 1 1 1 1 1 1 17.0 67.0 31.7 95.7 21.5 26.9 47.5 53.8 3 0 .6 [73.5] n i i- lC [101.5] M y tili- MTLC-

VO 1 1 1 1 1 00

171 1 13.1 47.0 58.8 82.6 40.7 64.7 88.2 20 .6 2 2 .4 24 .0 (N 25.3 25.5 MTLB-

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 100 20.5 22.1 MTLB- M ytilinii-1B

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 > 2 2 .0 2 2 .0 MTLB-

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 468 17.6 13.6 2 0 .5 MTLA-

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 26 6 19.6 MTLA- n i i- lA

o 00 1 1 1 1 1 1 1 1 6.4 IQ. 1 1 1 1 1 1 21.0 22 .0 22 .5 [80.5] M T L A - • • 237

2 1 1 1 1 1 1 1 1 1 1 1 1 1 87.5 11.5 31.5 24.7 64 .0 24 .5 26 .8 44 .3 MTLA-

E £

33 2 2 2 Q

Table 7: Mandibular dimensions ofHyaenictitherium wongii from various Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

74 Beitr. Paläont., 31, Wien, 2009

¿0 S’l sr

o ^H^ I ;s o (U rH oo oo 9.5 5.7 4.4 3.2 6.1 8.0

1 1 8.0 1 3.0 s- 3.0 1 00 H3 17.9 10.7 13.0 *4-» H 21.8 '

PP 1 I > so o so o I CS U-) Os

PP hP 1 I 1 1 I I I CS CS ro CO

1—1 CO oo Os o so ls m CO CO hP I I o I I 1 1 i—I \o

R U-) ls U-) 1-1

oo S; S ; so CO IS 00 s so s; 00 io s- í ñ 1 F 7 1-0 S; so CO s- CO SO up d so Up CO SO [1—11 Os up oo Os so 1—1 CS CO T—1 <1 i-H t hP H IS U"} vn s CO so i CO SO o o c s o o S’ o p S;

_g YL ‘■0 00 o S’ IS cs s^ uT1 LL o 7.5 is CO CO Os so so iT"H| 1 4.0

o 15.5 [9.5] CS 12.6] [23.3] hP u-} CS CO oo cs U-J o S; o 1--1 CS CO H XI s^ CO ro CO CO

u <-> in $ c O

Table 8 : Dental measurements ofHyaenictitherium wongii from Samos. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 75

MTLA-1 Figure 10: Upper and lower dent­ ition of H. wongii, Ictitherium viverrinum, Thalassictis robusta and Lycyaena chaeretis.

a. H. w ongii, upper tooth row of the maxilla MTLA-1, Samos, Greece; b. Ictitherium. viverrinum, upper tooth row of the skull NHML-PIK-M. 8983, Pikermi, Greece; c. L. chaeretis, upper tooth row of the skull NHMW- PIK-A.4744, Samos, Greece; d. L. chaeretis, upper tooth row of the skull NHML-PIK-M. 8978, Pikermi, Greece; e. T. ro- busta, upper carnassial, Kishinev, Moldavia (taken from K urtén, m & 1982); f. H. wongii, lower tooth e row of the mandible MTLC-18, Samos, Greece; g. I. viverrinum , lower tooth row of the mandible MTLC-18 NHML-PIK-M.8989 NHML-PIK-M.8989, Pikermi, Greece; h. L. chaeretis, lower tooth Iii% —* row of the mandible NHML- PIK-M.8979, Pikermi, Greece; f 9 j. Lycyaena chaeretis, lower tooth row of the mandible MNHN- PIK-3384, Pikermi, Greece; NHML-PIK-M. 8979 MNHN-PIK-3384 k. T. robusta, lower carnassial, Kishinev, Moldavia (taken from K urtén, 1982).

All specimens shown as if from right-hand side.

M T LA: Skull, MTLA-200; maxillary fragment with broken, while the nasal cavity is broken in its anterior C-M2 dex and sin, MTLA-1; left mandibular fragment part (PI. 4, figs 1, 2). The specimen M TLA-1 is the fron­ with c-p2-p4, MTLA-237; left mandibular fragment tal part of the skull with the maxilla (PI. 3, fig. 3). The with p3-ml, MTLA-266; left mandibular fragment with nasal cavity is elliptical, elongated and almost vertical to c-ml, MTLA-2; right mandibular fragment with il-p3, the palate with its posterior border above the middle of M TLA-468. the diastema P2-C, or rather above P i. In M TLA-1 the MTLB: Left hemimandible with i3-m2, MTLB-171; left posterior border of the nasal cavity is above the middle mandibular fragment with p4 and ml, MTLB-1; right of the canine. The nasals are narrow, pointed like spines mandibular fragment with p4 and ml, MTLB-100. and end at the anterior border of the orbits. The posterior MTLC: Right mandibular fragment with i2-ml, MTLC- border of the maxillary bones is situated at the middle of 18. the orbits. The frontals are triangular-shaped and separated MLN: Isolated P2 sin, MLN-27. by a shallow groove running across them. The infraorbital Measurements: The cranial mandibular and dental meas­ foramen is large situated above the contact between the urements are given in Tabs 6-8. P3 and P4. The orbits are oval-shaped and their anterior Description: border is above the middle of the P4. There is a strong Skull. The skull MTLA-200 is complete, but slightly postorbital constriction. The zygomatic arches are strong distorted in its left posterior part; the prosthion is slightly and strongly projected buccally. The braincase is relatively ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

76 Beitr. Palaont., 31, Wien, 2009

Hyaenictitherium wongii, Upper and Lower Teeth

■H. wongi, China (WERDELIN, 1988), n=36-108 H. wongii, MTLA mean, n=2-5 ■H. wongii, MTLB mean, n=1-3 H. wongii, MTLC mean, n=1 ■H. hyaenoides, China (WERDELIN, 1988), n=7-22 T. robusta, Kishinev, (KURTEN, 1982) n=1-15 ■I. viverrinum, Pikermi, WERDELIN 1988 n=2-13 I. viverrinum, China WERDELIN 1988 n=8-17

Figure 11: Logarithmic ratio diagram comparing the teeth ofHyaenictitherium wongii from various localities.

small and rounded. A weak sagittal line starts from the tip P i: It is single-rooted with a rounded crown and a strong of the postorbital projection, and is facing backwards. The cingulum, especially lingually. The PI of MTLA-1 is two sagittal lines are connected near the bregma, form­ stronger relative to that of MTLA-200. ing a sagittal crest, which is connected with the occipital P2: It is short and wide without an anterior accessory cusp. protuberance at the acrocranion. The occipital is strong, In its mesiolingual corner there is a small cingular projec­ with a strong crista nuchae and external occipital crista. tion, from which a crest starts to the tip of the main cusp. The occipital linea nuchae is strongly projected posteriorly, The posterior accessory cusp is well developed and situated well behind the posterior surface of the condyles (PI. 4, fig. on a distal cingular projection. There is a well-developed le). The condyles are relatively weak and do not laterally cingulum around the tooth. exceed the occipital bone. The foramen magnum is large P3: It is relatively short and wide. The great width is due and rounded. The postglenoid processes are large. The to a strong cingular projection situated in the distal lin­ bullae are relatively small, elliptical-shaped, and their gual half of the tooth. There is a small anterior accessory anterior border is slightly behind the anterior surface of cusp, situated in the mesiolingual corner of the tooth. The the postglenoid process (PI. 4, fig. Id). The palate is short posterior accessory cusp is large. There is a well-developed and relatively narrow. The palatal borders are parallel till buccal and lingual cingulum. P2, but then they curve bucally, reaching their maximal P4: The carnassial is elongated and narrow. The protocone breadth at the posterior end of the P4. The cheek teeth is large and well separated from the parastyle. In M TLA-1, rows are straight without imbrication of the teeth, which the anterior border of the protocone is slightly behind the are strongly in touch with each other. There is a small anterior border of the parastyle, while in M TLA-200 it is diastema between the canine and the third incisor, and a on the same level. The parastyle is large and slightly smaller smaller one between the canine and Pi. than the paracone. The metacone blade is relatively long Upper Dentition. The dentition of both specimens is and curves buccally. There is a strong lingual cingulum. well-preserved, except for some minor damages in that of M l: It is large, triangular, with well-developed and labi- M T LA -200 (PI. 3, fig. 3d; PI. 4; fig. 2). ally strongly projected paracone. The metacone is smaller, II, 2: The incisors are situated in a curved line. They have an while the protocone is large and high. elliptical crown and a strong buccal cingular projection. M 2: It is very small with an elliptical crown. 13: It is canine-like with a strong cingular projection Mandible. There are several mandibular fragments from distolingually. the various fossiliferous sites of the locality M TL (PI. 4, C: The canine is strong with a flattened root and an el­ figs. 3,4; PI. 5; PI. 6; figs. 1,2). The symphysis is relatively liptical transverse section. It has a strong mesiolingual and elongated and narrow, strongly inclining backwards. Part distal crest. The canine of MTLA-1 is relatively stronger of the ascending ramus is preserved in M TLB-171 (PI. 5, and less flattened than that of M TLA-200. fig. 6) and seems to be long and vertical to the mandibular ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 77

Hyaenictitherium wongii, Upper and Lower Teeth

LP2 LP4 LM2 Lp3 Lm1 Lm2 BP3 BM1 Bp2 Bp4 Bm2

■H. wongi, China (WERDELIN, 1988), n=36-108 H. wongii, MTLA mean, n=2-5 ■H. wongii, MTLB mean, n=1-3 H. wongii, MTLC mean, n=1 ■H. hyaenoides, China (WERDELIN, 1988), n=7-22 T. robusta, Kishinev, (KURTEN, 1982) n=1-15 -I. viverrinum, Pikermi, WERDELIN 1988 n=2-13 I. viverrinum, China WERDELIN 1988 n=8-17 -L. chaeretis, Samos NHMW-A.4744 L. chaeretis, Pikermi (orig. meas.)

Figure 12: Logarithmic ratio diagram comparing the teeth ofHyaenictitherium, Ictitherium, Lycyaena and Thalassictis.

corpus. The masseteric fossa is oval-shaped, deep and its ated on a distal cingular projection. The lingual border anterior border is situated below the m2 . The mandibular of the distal cingular projection is elevated and forms a corpus is elongated and relatively shallow. The inferior crest, like an entoconid. There is a well-developed buccal border of the corpus is curved and after the m l it points cingulum. upwards. There is a single mental foramen situated below m l: It is relatively short with a small and narrow talonid. the p2. The metaconid is relatively small, about the % of the Lower Dentition. The studied mandibular remains pre­ protoconid. The trigonid is about the 2A of the tooth’s serve well the various teeth (PI. 4, figs. 3, 4; PI. 5; PI.6 , length. The protoconid and paraconid are blade-like and figs. 1, 2). they are separated by a deep valley. The hypoconid is il, 2: They are slightly compressed laterally with an el­ strong and situated somewhat to the center of the talonid liptical occlusal aspect. basin. The entoconid is smaller and is connected with the i3: Larger than the other incisors and canine-like. It has hypoconulid, forming a crest covering the distal border of a small cingular projection situated bucally (on the side the talonid. A crest from the tip of the hypoconid is facing of the canine). obliquely to the junction of the metaconid and protoconid. c: The canine is strong, situated obliquely to the tooth A well-developed cingulum runs all around the tooth. row axis, curved backwards, compressed laterally, with Discussion. The taxonomy of the small-medium sized late a strong root and a rounded transverse section. It has a Miocene hyaenids is complicated. These hyaenids known mesiolingual and distal crest running from the top to the as “ictitheres” have a long taxonomic history (Kurten, base of the canine. 1982; Semenov, 1989; W erdelin Sc Solounias, 1991; pi: It is very small, single-rooted and with a rounded Koufos, 2000; Bonis, 2005). The “ictitheres” are common crown. in the Greek late Miocene faunas and were found in several p2: It is elongated relative to the p3, without an anterior localities: Pikermi, Samos, Axios Valley, Perivolaki (Thes­ accessory cuspid. The posterior accessory cuspid is strong, saly), Kerassia (Evia Island), (Koufos, 2006d). Among the situated on a distal projection of the distal cingulum. It has hyaenids of Pikermi there is one smaller thanA d crocu ta a strong lingual and weaker buccal cingulum. ex im ia and larger than Ictitherium viverrinum , referred p3: Relatively short and wide, with a vestigial-rudimentary to as Ictitherium hipparionum for a long time. Its generic anterior accessory cuspid. The posterior accessory cuspid name changed several times betweenPalhyaena, Thalas­ is small, situated on a wide distal cingular projection, sictis, Ictitherium and Hyaenotherium. The name P a lh y a en a giving the idea of a talonid. There is a well-developed hipparionum was given to a hyaenid from Mt. Luberon, buccal cingulum. France (Gervais, 1859); the type of this taxon was stored p4: It is short and wide, with a strong anterior accessory at the museum of Avignon, but it was lost (Kurten, 1982). cuspid and a very strong posterior accessory cuspid, situ­ This hyaenid is a nomen nudum and it can only include only ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

78 Beitr. Palaont., 31, Wien, 2009

the type from Mt. Luberon until it is found (W erdelin, fact that the latter is relatively young. However, the skull 1988; W erdelin &c Solounias, 1991), an opinion with M T LA-200 is narrower and has smaller teeth (Fig. 12). which I agree. A similar hyaenid, intermediate between Morphologically, the teeth of NHMW-SAM-A.4744 Adcrocuta eximia and Ictitherium viverrinum, was also de­ differ from those ofH. w ongii from Samos in having more scribed from the late Miocene of China under the name robust (more hyaenid-like) premolars, and a relatively Ictitherium wongii (Zdansky, 1924). After that, the similar shorter upper carnassial, as well as in lacking a lingual European material was included in this species. Finally, cingular projection in the P3, and in the absence of the this hyaenid was included in the genusHyaenotherium with M 2. A skull of L. chaeretis is also known from Pikermi H. wongii as the type species of the genus(S emenov, 1989). and housed at NHML. This skull (NHML-PIK-M.8978) However, there is another medium-sized one among the differs in the same characteristics as that from Samos, but Chinese hyaenids, named Ictitherium hyaenoides Zdan­ there is a lingual cingular projection near the root like a sky, 1924 which was later transferred to Hyaenictitherium protocone in the P3 (Fig. 10). Moreso, the cranial dimen­ (Kretzoi, 1938). Although the two hyaenids are different sions of NHML-PIK-M.8978 are clearly larger than those at specific level, they must belong to the same genus and of the studied H. w ongii from Samos. A hemimandible as Hyaenictitherium has the priority, both are included (NHML-PIK-M.8979) ofL. chaeretis is also known from in this genus (Zhang et al., 2002; B onis, 2004, 2005). Pikermi and stored at NHM L. This mandible differs Today the late Miocene hyaenid betweenAdcrocuta eximia from the studied ones ofH. w ongii from Samos in hav­ and Ictitherium viverrinum is known as Hyaenictitherium ing a larger size and teeth (Fig. 12), a stronger anterior w ongii (Zdansky, 1924). accessory cuspid in the p4, a weaker talonid significantly Such a medium-sized hyaenid, known as Ictitherium hip- smaller than that of H. wongii, and in the absence of the parionum , is reported in the old collections from Samos m2. Another mandibular fragment (MNHN-PIK-3383) (Forsyth M ajor, 1894; Pilgrim, 1931). In the descrip­ of L. chaeretis from Pikermi also differs from the studied tion of the Samos hyaenids it is referred to as Ihalassictis material in the same characteristics as mentioned above, wongii (Solounias, 1981; W erdelin, 1988), while later it and its teeth are larger (Fig. 12). was transferred to Hyaenotherium (W erdelin & Solou­ The morphological characteristics and the size of the nias, 1991). In our new collection from Samos localities a studied skulls and mandibles fit in well with those ofH. medium-sized hyaenid without description is reported as wongii from Eurasia (Solounias, 1981; W erdelin, 1988; Hyaenotherium w ongii (Koufos et al., 1997, 2004). W erdelin & Solounias, 1991; Koufos, 2000; Bonis, The most common hyaenids in the late Miocene faunas are 2005). The studied material has been compared for the Adcrocuta eximia, Ictitherium viverrinum and Hyaenictith­ first time to the material ofH. w ongii from Samos, stored erium wongii. The studied material is easily separated from at NHML and NHM W , as well as to that from Axios Adcrocuta by its smaller size, the larger protocone in the P4 Valley, housed at LGPUT, and it has a similar morphol­ and the relatively larger and tricuspid talonid of the m l. ogy. The cranial and mandibular dimensions of the studied The distinction from Ictitherium is also quite clear. The type material are very close to those of the old material from species of the genus isI. mverrinum, known from Pikermi, Samos, as well as to that from Axios Valley (Tabs. 6, 7). Greece. The studied material from Samos was directly The metrical comparison of both upper and lower teeth compared with the PikermiIctitherium housed at NHML, indicates that all the studied specimens are very close to NHMW, and some casts of the BSPM collection housed the mean values forH. w ongii from Axios Valley, and they at LGPUT, as well as with the material ofI. viverrinum are always within the ranges of variation for the Samos and from Axios Valley (Koufos, 2000). The studied material China samples of H. w ongii (Fig. 1 0 ,11). Thus, the stud­ differs from Ictitherium viverrinum in being a larger size, ied material can be ascribed to H. w ongii with certainity. less wide canines, having an elongated blade in the P4, a Some minor differences in the dental dimensions between relatively larger protocone in the P4, reduced molars and the various specimens could be considered as sexual or a relatively shorter talonid in the m l (Figs. 10-12). ecological differences. There is a clear sexual dimorphism Hyaenictitherium wongii was considered as belonging in carnivores, which could provide some minor metrical to Ihalassictis for a long time, but the distinction of the differences. H. w ongii is a widespread taxon, known from studied material from the genus Ihalassictis is quite clear. China to Spain, and some minor differences could also be Ihalassictis is known by the single species T. robusta found due to the different palaeoecological niches. in Kishinev, Moldavia, with fragmentary type material As was mentioned above, there is a medium-sized late M i­ (Kurten, 1982). Ihalassictis differs from H. w ongii in ocene hyaenid namedHyaenictitherium hyaenoides among having a shorter blade in the P4, a higher protoconid in the Chinese material. The latter differs fromH. w ongii by the m l, a stronger talonid with higher cuspids of the m l its reduced molars (especially M2 and m2), the smaller P2 and a smaller size (Fig. 12). and p2, the smaller main cuspid of the p4, the relatively Another medium-sized hyaenid from Samos is Lycy- smaller talonid of the m l and the generally larger size (Fig. aena chaeretis. There is a complete skull (NHMW- 12). Bearing in mind all the above mentioned comparisons, SAM -A.4744) from Samos housed at NHMW, which the studied material can be ascribed toHyaenictitherium is compared to the studied material ofH. w ongii from wongii. The sole P2 M LN -27 has a similar morphology Samos. The skull M T LA -200 has a similar length to and dimensions to H. wongii, but the limited material can­ that of N H M W -SAM -A.4744, but this is due to the not allow a certain determination and thus it is referred to ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 79

as Hyaenictitherium cf. H. wongii. Besides its far-reaching P2: The tooth is more or less squarish, with a vestigial expansion, the taxon also has a wide stratigraphic range anterior accessory cusp situated mesiolingually. A strong from Vallesian to the end of middle Turolian. crest connects it with the tip of the main cusp. The pos­ terior accessory cusp is larger and low, compressed on the Genus A dcrocuta K retzoi, 1938 distal part of the tooth. P3: It is short and robust; the mesial breadth is longer Adcrocuta eximia (Roth & W agner, 1854) than the distal one. Morphologically it is similar to P2 (PL 6, figs. 3-6; PI. 7, figs. 1, 2) but larger. P4: It is elongated and relatively slender; the protocone is Locality: Mytilinii-lA (MTLA), Adrianos ravine, My- partially broken but it seems to be very weak restricted to tilinii Basin, Samos, Greece. the base of the tooth, typical forAdcrocuta. The parastyle is The material labelled as PM M S belongs to the collection strong and the paracone is high. The metacone is broken. of Prof. J. Melentis and is stored at NHMA. According to M l: It is very small, short and wide, with three cuspids him it was collected from Adrianos ravine, more precisely and an almost elliptical crown. It is slightly worn and the from M TLA. preserved cusps are low. Age: Middle Turolian, MN 12 (late Miocene); 7.1-7.0 Mandible. The mandibular fragment (PMMS-70a) lacks Ma. the ascending ramus and preserves the alveoles of the inci­ Material: sors and canine as well as the tooth row pl-p4 (PI. 6, fig. M TLA. Isolated p2 sin, M T L A -6; right dp4, MTLA, 4). The symphysis is strong and elliptical, pointing upwards 290; left dp4, MTLA-466. after the p2 . The mandibular corpus is high and its inferior PMMS. Frontal part of the skull with P2-M 1 dex and P2- border is straight till m l; then it curves roughly upwards. P3 sin, PMMS-70; left mandibular fragment with pl-p4, There is a large mental foramen below the middle of the 2p PMMS-70a; left upper canine, PM M S-71. and a smaller one below the posterior border of the p2. The Measurements: The cranial and mandibular measurements masseteric fossa seems to be deep and its anterior border are given in Tabs. 9 , 10. is behind the posterior border of the ml. The tooth row is almost straight and there is no imbrication of the teeth. Description: There is no diastema between them but all are in contact Skull. The specimen PMMS-70 is the frontal part of with each other. The canine is situated somewhat labially the skull with the maxilla (PI. 6, fig. 3). The frontals are and the pi lingually to the axis of the toothrow. broken and only a part of the right orbit is preserved. The c: The canine seems to be large; the dimensions of the nasal cavity is oval-shaped and relatively wide. The lateral alveole are: 21.2 x 16 mm. borders of the nasal cavity are almost vertical to the palate pi: It is small, single-rooted, monocuspid with a rounded and its posterior border is above the anterior edge of the crown. canine. There is a large infraorbital foramen situated above p2: It is short and robust, without an anterior accessory the P3. The orbit seems to be rounded and its anterior cuspid, but with a basal mesiolingual cingular projection. border is above the posterior border of the P3. The palate The posterior accessory cuspid is larger, situated on a distal is relatively short and wide. The maxilla preserves the tooth projection of the distal cingulum. There is a faint lingual rows P2-M 1 dex and P2-P3 sin, as well as the roots of and buccal cingulum. the P4 and M l. The metacone of the right P4 is broken. p3: It is similar to the p2, but larger and with a large an­ There are all the alveoles of the incisors, canines and Pi. terior accessory cuspid situated mesiolingually. The alveole of the 13 suggests a large tooth like a canine. P4: It is relatively robust with a very strong anterior acces­ The canine’s alveole also indicates a large tooth. sory cuspid. The posterior accessory cuspid is smaller than the anterior one and situated on a large distal projection of PM M S-70 dex | sin PM M S-70a the distal cingulum, which is elevated all around the distal LP2 17.2 17.1 Lpl 4.8 part of the tooth. The distal cingulum is strongly elevated BP2 11.7 11.7 Bpl 4.7 lingually, giving the idea of an entoconid. LP3 21.7 22.0 Lp2 16.2 M ilk teeth: Two isolated dp4 are included in the new BP3 16.0 16.0 Bp2 10.8 collection. In M TLA-290 (PI. 7, fig. 1) the talonid is par­ LP4 [37] — Lp3 18.2 tially broken, very small and seems to be monocuspid. The BP4 ant. 16.3 — Bp3 12.8 protoconid has twice the height of the paraconid, which is BP4 post. 11.8 — Lp4 21.5 blade-like. The two cuspids are separated by a very deep valley open lingually. There is a faint buccal cingulum. The Lblade — — Bp4 13.2 M T LA -466 (PI. 7, fig. 2) is small and narrow. The tips L M l 5.8 — of the paraconid and protoconid are broken, but they are BM l 12.6 — blade-like. The talonid is small, monocuspid and situated MTLA-6 p2 = 15 .7 x 11.5 lingually. There is a mesial, mesiolingual and mesiobuccal M TLA-290 dp4 = 19.2 x 7.1 cingulum. M TLA-466 dp4 = 17.3 x 6.5 Discussion: Adcrocuta eximia is reported in all old col­ PM M S-71 C = 19.5 x 14.6 lections from Samos (Forsyth M ajor, 1894; Pilgrim, ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

80 Beitr. Palaont., 31, Wien, 2009

Table 9: Cranial dimensions ofAdcrocuta eximia from various Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 81

Table 10: Mandibular dimensions otAdcrocuta eximia from various Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

82 Beitr. Paläont., 31, Wien, 2009

Adcrocuta eximia, Upper Teeth

0,180

-0,140 ■ -0,160 ------LP2 LP3 LP4 LM1 BP2 BP3 BP4 BM1

------Hyaena hyaena, n=11-24 (HOWEL & PETTER, 1985) - - O -A . eximia, Pikermi, min, n=3-10 (HOWELL & PETTER, 1985) 1 B —A. eximia, Pikermi, max, n=3-10 (HOWELL & PETTER, 1985) - -O - -A . eximia, Samos, min, n=7-12 (SOLOUNIAS, 1981) — • - -A . eximia, Samos, max, n=7-12 (SOLOUNIAS, 1985) — X— PMMS-70 ♦ A. eximia Vallesian Axios —- * — A. eximia Turolian Axios

Figure 13: Logarithmic ratio diagram comparing the upper teeth ofAdcrocuta eximia from various localities.

Adcrocuta eximia, Lower Teeth

------Hyaena hyaena, n=17-21 (HOWELL & PETTER, 1985) - -Q - -A . eximia, Pikermi, min, n=5-10 (HOWELL 8. PETTER, 1985) B -A . eximia, Pikermi, min, n=5-10 (HOWELL &■ PETTER, 1985) - - O -A . eximia, Samos, min, n=11-14 (SOLOUNIAS, 1981) -A . eximia, Samos, min, n=11-14 (SOLOUNIAS, 1981) — X— MTLA-6 — X ^ “ PMMS-70a ♦ A. eximia Vallesian Axios — A ■ A. eximia Turolian Axios

Figure 14: Logarithmic ratio diagram comparing the lower teeth ofAdcrocuta eximia from various localities.

1931; Solounias, 1981; K oufos & M elentis, 1982). eximia from Pikermi (Roth & W agner, 1854). Later, its The old material, or part of it, was described earlier under name changed several times until being classified asAd­ the name Hyaenictis eximia (Solounias, 1981). However, crocuta (Schmidt-K ittler, 1976). The studied specimens Hyaenictis is a different taxon (Howel & P etter, 1985) PM M S-70, 70a, 71 have been described earlier(K oufos and thus the Samos material was soon re-classified as & M elentis, 1982) but I decided to include them in the Adcrocuta. The species was originally described asHyaena present article together with the new collection of Samos, ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 83

Figure 15: Lower deciduous dentition of Adcrocuta eximia from various localities. a. dp4, MTLA-290, M ytilinii- lA, Samos; b. dp4, MTLA- 466, Mytilinii-lA, Samos; c. Lower deciduous dentition of the mandible LGPUT-PER-1240, Perivolaki, Greece; d. Lower deciduous dentition of the man­ dible NHML-PIK-M.8973, Pikermi, Greece; e. Lower de­ ciduous dentition of the man­ dible NHML-PIK-M.8980, Pikermi, Greece.

All specimens are rotated to be right.

giving more data and comparisons for them. Although urements for the Pikermi and Samos skulls are original, A. eximia is very common in the old Samos material, it is coming from the material stored at N H M L, M NHN and almost absent in our new collection, as it is represented by NHMW). The above indices and comparisons suggest an isolated p2 and two m ilk molars only. that the studied material belongs to the typical subspecies The morphological features of the studied material, such A. e. eximia. as the robust premolars, their squarish occlusal outline Three mandibular fragments with milk molars of A. eximia with a more or less straight mesial and distal outline, the are known from Pikermi (Pilgrim, 1931). The specimen relatively slender upper carnassial with a weak protocone, NHML-PIK-M.8973 also contains the m l, whose mor­ the very small M l, and the presence of a strong anterior phology and dimensions are very similar to the typicals. accessory cuspid in the p4 are characteristics ofA. eximia. eximia. The length of the dp4 varies between 17-18.3 mm The metrical comparison of the studied material suggests and the breadth between 6.8-7.2 mm being very close to close similarities to A. eximia, too (Tabs. 9,10; figs. 13-14). those of MTLA-290, 466. A mandibular fragment with Both upper and lower dental dimensions are within the milk dentition is known from Perivolaki, Greece (Koufos, ranges ofvariation for A. eximia from Pikermi and Samos, 2006a). Its morphology and dimensions (18.3 x 7.0 mm) as well as close to the Vallesian and Turolian form of A. are very close to the studied dp4 (Fig. 15). eximia from Axios Valley, Macedonia, Greece (Figs. 13, Adcrocuta eximia is a large-sized form similar to the re­ 14). Bearing in mind all these comparisons and similari­ cent spotted hyaena Crocuta crocuta and for a long time ties, the studied material can be ascribed to A. eximiawith. it was referred to the same genus. The strong jaws, the certainity. Two subspecies are known, the typical Turolian crushing premolars, the elongated upper carnassial and A. e. eximia and the Vallesian A. e. leptoryncha, known from the strong anterior accessory cuspid in the p4 suggest Axios Valley. The second subspecies is characterized by shearing and breaking action. This hyaena was mainly an elongated and narrow palate, a higher skull, a longer a carrion eater and it lived in groups like the recent diastema C-P2 and narrower lower premolars (Bonis ones. Its first appearance in the Eastern Mediterranean & Koufos, 1981; K oufos, 2000). The index “distance and Europe dates to late Vallesian (MN 10) at ~9.6 Ma prosthion-middle of the line connecting the posterior end and it disappeared at the end of Miocene at ~5.3 Ma of the P4 / distance between the posterior ends of the P4” (Koufos, 2003). The greatest abundance of the taxon is is 154 for A .e. leptoryncha versus 130 for PM M S-70. The known from the Turolian, where it is very common in all corresponding index varies from 139-143 for the SamosA. Eurasian localities. The taxon is widespread, known from eximia and from 130-141 for the Pikermi A. eximia. The China to Spain. Considering its palaeoenvironment, it diastema C-P2 is 13.1 mm in A. e. leptoryncha versus 10.2 mainly lived under relatively open and dry conditions. mm in PMMS-70. In the Samos A. eximia, the diastema In the Eastern Mediterranean, the palaeoenvironment is 9-10 mm and in the Pikermi one 9.7-10 mm (the meas­ changed at the beginning of Vallesian from a relatively ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

84 Beitr. Palaont., 31, Wien, 2009

Table 11: Cranial dimensions of M etailurusparvulus and M . m ajor from various Greek localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 85

M etailurus p a r v u l us

m a n d ib u l a r measurements Mytilinii-lA Pikermi Pikermi Chomateres Kerassia Maragheh

AMPG- NHMW- AMPG- NHML- AMPG- MTLA-235 PIK PIK- KER- MRG-M. CHO-nn -PG 01/103 A.4712 Kl/210 3825 Original measurements 1. Length anterior c-coronoid process — — — — 110 — 2. Length anterior c-condyle — — — 101 10.5 — 3. Height inferior border at angle-coronoid — — — — 61 — 4. Height inferior border at angle-condyle — — — 21.5 22.5 — 5. Height condyle-coronoid — — — — 16.5 — 6. Symphysis length — — 26 24.5 22.8 20.5 7. Height in front of p2 20.3 — — — — — 8. Idem in the middle of p3 (lingual) 13.4 18.7 — 17.5 17.5 20.6 9. Idem in p4 (lingual) 16.3 18.5 20.2 18 17.5 20.3 10. Idem in ml(lingual) 16.4 18 — 18 — 19.9 11. Idem behind m2 — — — — 8.5 — 12.Diastema p2-c — 8.2 — 8.3 — — 12.Diastema p2-i3 — 16.3 — — — — 13.Diastema i3-c — 0 59.3 0 — — 14. Lengthp2-ml — — — — — — 15. Length c-ml 60 57 — 60 57 57 16. Length p2-p4 — — 43.8 — — — 17. Length p3-ml — 40.5 — 39.5 40.5 39.1 18. Length p2-m2 — — — — — — 19. Length c-m2 — — — — — —

Table 12: Mandibular dimensions of Metailurusparvulus from various Greek localities.

close and humid one to being more open and dry(K ou­ Measurements: The cranial and mandibular measurements f o s , 2006b). During that time A. eximia appeared in are given in Tabs. 9, 10. the Eastern Mediterranean. The environment gradu­ ally changed to more and more open and dry during dex sin Turolian, providing more favourable conditions for A. M T L A -234 P3 = 14.0 x — — eximia, which increased remarkably in numbers. During P4 = 22.2 x — 21.6x10.0 the end of Miocene another faunal change led to the MTLA-235 P3 = 10.2x4.8 more closed and wet palaeoenvironmental conditions of early Pliocene and to the disappearance ofA. eximia Description: (Koufos, 2003, 2006c). Skull. The skull lacks the muzzle, the left zygomatic arch, (the right one is partially preserved), and both orbits are Family G ray, 1821 badly preserved (PI. 7, fig. 3). The skull seems to be short with a short muzzle. The infraorbital foramen is large, Genus M etailurus Z dansky, 1924 situated above the parastyle of the P4. The orbit seems to be rounded, with its anterior border above the P3. The Metailurus parvulus Z dansky, 1924 braincase is voluminous, but its upper surface is flattened. (PI. 7, figs. 3-6) There are no sagittal lines or crests. The zygomatic arches seem to be strong and strongly project buccally. The oc­ Locality: Mytilinii-lA (MTLA), Adrianos ravine, My- cipital is small, with a well-developed external occipital tilinii Basin, Samos, Greece. crest, starting from the posterior border of the postglenoid Age: Middle Turolian, MN 12 (late Miocene); 7.1-7.0 Ma. process. The nuchal crest is very small and restricted to Material: Skull, MTLA-234; right mandibular fragment the breadth of the external occipital crest. The occipital with P3, M TLA-235. condyles are relatively small, while the foramen magnum is ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

86 Beitr. Palàont., 31, Wien, 2009

Figure 16: Upper and lower dent­ MTLA-234 MTLA-235 ition of Metailurusparvulus from various localities. a. Upper tooth row of the skull MTLA-234, Mytilinii-lA, Sa­ mos; b. Upper tooth row of the skull AMPG-CHO-nn, AMPG-CHO-nn Chomateres, Greece; c. Upper NHMW-PIK-A.4712 tooth row of the skull AMPG- HAL-1967/5, Halmyropotamos, Greece; d. Upper tooth row of the skull Ex.4, Maragheh, Iran (taken from Z dansky, 1924); e. Lower tooth row of the mandi­ AMPG-HAL-1967/5 AMPG-K1/210 ble MTLA-235, M ytilinii-lA , Samos; f. Lower tooth row of the mandible NHMW-PIK-A.4712, Pikermi, Greece; g. Lower tooth row of the mandible AMPG- KER-Kl/210, Kerassia, Greece; h. Lower tooth row of the man­ NHML-MAR-M.3825 dible NHML-MAR-M- 3825, Maragheh, Iran; j. m l, MNHN- MAR-nn, Maragheh, Iran. Ex. 4 (ZDANSKY, 1924)

All specimens shown as if from right-hand side.

large and elliptical-shaped. The bullae are ovoid and rela­ by a deep valley. There is a small cingular projection in tively large. They are well-separated from the condyles by a its mesiobuccal corner. The metacone is blade-like, low wide and deep channel. Their anterior border is on line with and curved buccally. It is distinguished from the paracone the posterior surface of the postglenoid process. The bullae by a deep valley, very well visible on the buccal surface are separated in two parts by a shallow groove across their of the tooth. long axis. The choanae are wide and their anterior border is M l: It is situated vertically to the carnassial, and, from almost level with the posterior border of the P4. The palate the alveole, seems to be small. is short and wide having a more triangular shape. Mandible. The studied mandibular fragment lacks the Upper Dentition. The dentition of the skull MTLA-234 ascending ramus and preserves the p3 and the alveoles includes the right P3-P4 and the left P4; there also are the of the p4 and m l (PI. 7, fig. 6). The mandibular corpus alveoles of the canines, as well as the alveoles of both M l is shallow and its inferior border is straight from below (PI. 7, figs. 3c, 4, 5). The tooth row is straight. the p3 to the anterior border of the masseteric fossa. The C: The alveoles of the canines indicate a relatively large symphysis is short and almost vertical to the mandibular canine. They are elliptical and their dimensions are 11.5 corpus, a characteristic feature in felids. The lower part x 8 mm. of the symphysis projects and forms an angle, giving a P3: It is mesially and labially broken. It is elongated and nar­ strong construction to the anterior part of the jaw. There row with a clearly distinguished posterior accessory cusp. are two mental foramens, one large, below the middle P4: The left carnassial is very well-preserved, elongated, of the diastema c-p3, and a smaller, below the contact narrow and felid-like (PI. 7, figs. 4, 5). The protocone is between the p3 and p4. The masseteric fossa is deep and very small, situated well posterior to the parastyle. It is a its anterior border is well behind the m l. small cusp situated at the base of the paracone, well dis­ p3: It is short and narrow with its distobuccal corner tinguished (PI. 7, figs 4, 5), but it disappears by attrition. slightly broken. There is no anterior accessory cuspid, but The parastyle is small compared to the paracone and lower. a small cingular projection, situated in the lingual part It is well-distinguished from it and from the protocone of the tooth. The posterior accessory cuspid is larger and ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 87

Figure 17: Scatter diagram length/breadth comparing the upper carnassial of Metailurus from various localities.

Length

X M . parvulus, MytiliniMA ■ M. parvulus, Plkermi (orig. meas.; ROUSSIAKIS, 2006) A M . parvulus, Chomateres (orig. meas.) A M . parvulus, Halmyropotamos (orig. meas.) Q M . parvulus, China (ZD AN SKY, 1924) □ M. major, Pikerml (ROUSSIAKIS, 2001) O M. major, Halmyropotamos (orig. meas.) O M. major, China (ZDANSKY, 1924)

Figure 18: S catter diagram length/breadth comparing the p3 of Metailurus from various localities.

X M . parvulus, Mytillnii-1 A ■ M. parvulus, Pikermi (orig. Meas.; ROUSSIAKIS, 2006) AM . parvulus, Chomateres (orig. meas.) • M. parvulus, Kerassia (ROUSSIAKIS, 2006) S M . parvulus, Maragheh (orig. meas.) BM . parvulus, China (ZDANSKY, 1924)

situated on a cingular projection. In the labial corner of Halmyropotamos and Chomateres, stored at A M P G , this projection, there is a very small cuspid level with the NHMW and PIUW, as well as to the material from posterior accessory cuspid, like a hypoconid. There is a Maragheh housed at NHML and MNHN. weak labial cingulum. A skull of Metailurus parvulus from Pikermi, stored in Discussion: PIUW, is described by M elentis (1967); a mandibular The species Metailurus parvulus was set up by H ensel fragment with P3-P4 (NHMW-PIK-4712), also from (1862), based on material found in Pikermi and descri­ Pikermi, is stored at NHMW. A skull and a mandible of bed under the name Machairodus parvulus. Later, it was Metailurusparvulus without any number from Chomateres recognized in several late Miocene Eurasian localities. A (a locality situated about 1 km from Pikermi) are stored mandible from Maragheh described asFells cf. brevirostris at A M P G (Symeonidis, 1978). The morphological cha­ by K ittl (1887) was later transferred to Metailurus parvu- racteristics of the studied skull and mandible resemble lus (Beaumont, 1961). Some machairodont material from those of the Pikermi and Chomateres material (Fig. 16), China was described as Metailurus minor by Zdansky while their dimensions are very similar (Tabs.11 , 12). The (1924), but later it was ascribed to Metailurus parvulus dimensions of the teeth are also very close to those of the (Thenius, 1951; Beaumont, 1961). In Samos, the species Pikermi and Chomateres material (Fig. 17). A skull with is known from QA and Q5 (Solounias, 1981; tab. 7), but P3-M1 (AMPG-HAL-1967/5) of Metailurus parvulus ■without mentioning the material it is based on, and with was described from the locality of Halmyropotamos, no description. The new material from Samos described Evia Island (M elentis, 1967). The morphology and size in this article is compared to the material from Pikermi, of the upper carnassial of MTLA-234 fit in well with ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Beitr. Paläont., 31, Wien, 2009

Machairodus giganteus M. aphanistus

Halmyro­ UPPER CANINE Mytilinii-1 Pikermi Thessaloniki Maragheh Kemiklitepe-D potamos

AMPG- AMPG- MNHN- MNHN- MNHN- PMMS-69 HAL-1967/6 PIK-1967/7 SLCL938 MRG-3366 KTD-66

M elentis (1967) Original measurements Dimensions at the base: 35.8x14.5 35.2x 14.3 34.2x13.8 36.8x14.9 29.0x11.9 30.8x12.7 Idem 3 cm from the base: 32.5x11.5 32.2x11.8 27.9x11.0 31.0x11.0 — Idem 6 cm from the base: 19.9x8.3 24.6x9.7 18.2x8.9 19.1x8.2 —

Table 13: Dimensions of the upper canine ofMachairodus from various localities.

those of Halmyropotamos (Figs 16, 17). The species is Montredon. All the M etailurusparvulus-bz& nng mammal also known from another locality of Evia Island, named localities are dated from middle to late Turolian (MN 12- Kerassia, from where an almost complete skeleton was 13); the locality of Montredon is dated to late Vallesian described (Roussiakis et ah, 2006). Although the skull (NOW, 2007). Thus, the species seems to appear earlier, is absent, the preserved mandibular ramus has similar during Vallesian, in Europe. This idea is supported by morphological characteristics as those of MTLA-235, its recent discovery in the late Vallesian locality Ravine like the elongated and shallow mandibular corpus with a de la Pluie of Axios Valley (Macedonia, Greece), (pers. straight inferior border, the deep masseteric fossa ending observ.). behind the m l and the two mental foramens (one below the postcanine diastema, and another below the posterior Genus Machairodus K aup, 1833 border of the p3). The Kerassia p3 lacks an anterior ac­ cessory cuspid and has a small posterior one, like that of Machairodus giganteus (W agner, 1848) M TLA-235 (Fig. 16). (PI. 7, fig. 7) The comparison of the Samos M etailurusparvulus with the Chinese sample described byZ dansky (1924) indicates Locality: ?Mytilinii-lA (MTLA), Adrianos ravine, My- close similarities. The general morphological features of tilinii Basin, Samos, Greece. the skull and mandible resemble each other, while their Material: Fragment of the upper canine, PM M S-69. The size seems to be similar too. Unfortunately, Z dansky material labelled as PM M S belongs to the collection of (1924) does not give cranial measurements for a direct Prof. J. Melentis and is stored at NHMA. According to comparison, but some measurements taken from his il­ him, it was collected from Adrianos ravine, more precisely lustrations fit in with those of the Samos material. Their from M TLA. dental measurements, however, are slightly larger than Measurements: The measurements are given in Tab. 13. those of the studied material (Fig. 17). The genus M e t a ilu r u s is known from Greece by an­ Description: The material was described byK oufos Sc other species, namedM . m a jo r and found in Pikermi and M elentis (1982). The preserved canine has a height of Halmyropotamos (M elentis, 1967; Roussiakis, 2001). about 77 mm from the base. It is flattened and curved This species is also mentioned in the lists of the old collec­ backwards. Both mesial and distal borders are indented; tion of Samos (Solounias, 1981; tab. 7). M eta ilu ru s m a jo r the mesial indented border is double, from the base to the clearly differs from M etailurus parvulus by its remarkably middle of the preserved tooth; then it becomes single. A larger size (Fig. 17). Among the known late Miocene strong crest goes over the middle of the buccal and lingual machairodonts, the species Paramachaerodus orientalis, surfaces. M achairodus aphanistus and M achairodus giganteus are Discussion: The morphological characteristics of the known from Greece. A ll of them are larger thanM e ­ studied canine fragment are similar to those ascribed to tailurus parvulus. Moreover, their upper carnassial has a M. giganteus from Pikermi, Halmyropotamos and Axios clear ectoparastyle and reduced protocone, while the lower Valley (W agner, 1848; M elentis, 1967; Koufos, 2000). carnassial lacks or has a rudimentary talonid only. Moreover, its dimensions are also similar toM. giganteus Although it is rare, M etailurus parvulus spread widely and (Tab. 13). inhabited the whole of Eurasia, from China to Spain. Besides the Greek localities, it is also known from the Spanish localities of Los Mansuetos and El Arquillo, the 3. Conclusions Italian locality of Gravitelli and the Moldavian locality of Cimislia. In Asia, except from China and Iran, it is known Although the late Miocene Samos carnivores are numerous from the Turkish locality of Sandikli Kinik. A form similar in the old collections, they are relatively fewer in the new to M etailurus parvulus is traced in the French locality of material. In the lists given byS olounias (1981), Bernor ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 89

et al., (1996) and NOW (2007), 15 different carnivoran for granting me access to the fossils at their disposal. Many taxa are mentioned (Tab. 1). Six of them are present in thanks to Dr. C. Sagne (MNHN), Dr. C. Argot (MNHN) the new collection, while Protictitherium was recognized and Dr. C. Soligo (NHML) for their help in searching the for the first time on Samos (Tab. 1). Bearing in mind that collections, and their great hospitality. I also thank Dr. D. the old material was collected a long time ago by various Geraads for providing me with casts ofProtictitherium aegeum. people excavating for many years on Samos, the new col­ Finally, I wish to thank my collaborators and several students lection is comparatively rich in carnivores. The majority who excavated with me and helped me with the collection of the found carnivores are typical for the Turolian. The and preparation of the fossils. I also thank Dr. M. Morlo for mustelid Parataxidea maraghana, apart from Samos, is his useful comments on the manuscripts. known from Middle Maragheh, which is dated to middle Turolian, M N 12 (NOW, 2007). The species is also known from the early Turolian locality of Kara9 ahasan (Turkey), 5. References dated to early Turolian, MN 11 (NOW, 2007). The strong similarity of the M TLA material ofP. maraghana to that Beaumont, G. de., 1961. Recherches sur Felis attica W ag­ of Middle Maragheh (type locality), indicates a similar ner (Mammalia, Carnivora) du Turolien de Grèce. middle Turolian age (MN 12). The hyaenidPlioviverrops — Archives des Sciences, 39:377-386, Lausanne. orbignyi is well known from Greece and mainly reported Beaumont, G. de., 1969. Brèves remarques sur le genre from the middle Turolian localities of Pikermi (type), Plioviverrops K retzoi (Carnivora). — Bulletin Labo­ Vathylakkos, Prochoma and Perivolaki; however, it is ratoire Géologie Minéralogie Géophysique, Museum also found in the early Turolian locality of Ravine des Géologique, Univ. Lausanne, 180:1-7, Lausanne. Zouaves-5. Its presence in MTLB and its similarity to the Beaumont, G. de. 8c M ein, R, 1972. Recherches sur known Greek material suggests an early - middle Turolian le genre Plioviverrops K retzoi (Carnivora, ?Hyae- age, MN 11-12; bearing in mind that its main appearance nidae). — Comptes Rendus des Séances, SPHN, is observed in middle Turolian (MN 12), a similar age is 25:383-394. quite possible for MTLB. The hyaenid Protictitherium Bernor, R.L., Solounias, N., Swisher III, C. C. 8c crassum has a wide stratigraphic distribution from MN 4 van C ouvering, J.A., 1996. The correlation of three to MN 13 (NOW, 2008). However, the similarity of the classical “Pikermian” mammal faunas - Maragheh, MLN mandible to that from the Turolian of Axios Valley Samos and Pikermi - with the European MN unit (Macedonia, Greece) indicates a possible Turolian age for system. — [in:] Bernor R-L., Fahlbusch, V. 8c the locality. The other two hyaenid taxa,Hyaenictitherium M ittmann, H-W. (eds). The evolution of western wongii vxAAdcrocuta eximia, are known from the Vallesian Eurasian Neogene mammal faunas. — 1:137—156, and Turolian, too. Thus, they cannot be very relevant to New York (Columbia University Press). biochronology, suggesting only a late Miocene age. Like­ Bonis L. de., 2004. Carnivores hyaenidés du Miocène wise, Metailurusparvulus is reported from late Vallesian supérieur de Turquie. — Zona Arquaeologica Mis­ to late Turolian (MN 10-13), and Machairodus giganteus celánea en homenaje a Emiliano Aguirre, 2. Pale- from late Vallesian to early Pliocene (MN 10-15), (NOW, ontologia. Museo Arqueológico Regional, Madrid, 2007); their wide stratigraphic distribution cannot help pp. 108-117. with biochronology. Bonis L. de., 2005., Carnivora (Mammalia) from the late Tie available data from the study of the carnivores sug­ Miocene of Akkaçdagi, Turkey. — Geodiversitas, gests that the age of the localities must be Turolian; more 27(4):567-589, Paris. precisely, the M T LA and MTLB faunas could be dated to Bonis, L. de&KouFOS, G.D., 1981. A new hyaenid (Car­ middle Turolian. These results fit in quite well with those nivora, Mammalia) in the Vallesian (late Miocene) of from the study of the whole fauna and magnetostratigraphy Northern Greece. — Scientific Annals, Faculty of (Koufos et al., this volume). Physics and Mathematics, University of Thessaloniki, 21:79-94, Thessaloniki. Bonis, L. de 8c Koufos, G.D., 1991. The late Miocene 4. Acknowledgements small carnivores of the lower Axios Valley (Macedo­ nia, Greece). — Geobios, 24(2):361-379, Lyon. Tie excavations on Samos have been supported by the Pre­ C rusafont-Pairó, M. ôcPetter, G., 1969. Contribution fecture of Samos and the “Konstantinos and Maria Zimalis” à l’étude des Hyaenidae. La sous-famille des Ictith- Foundation. The Municipality of Mytilinii provided generous eriinae. — Annales de Paléontologie, 55(1):89—127, help too. The Natural History Museum of the Aegean of­ Paris. fered us the premises for the preparation and storage of the D epéret, C., 1892. La faune des mammifères miocènes de fossils. The Synthesys Program supported my visits to Vienna la Grive Saint-Alban. — Archives Museum Histoire (AT-TAF-702), London (GB-TAF-1842) and Paris (FR- naturelle Lyon, 5:31-96, Lyon. TAF-3102). I thank Prof. M. Dermitzakis (AMPG), Prof. Forsyth M ajor, C.J., 1894. Le gisement ossifère de G. Theodorou (AMPG), Dr. S. Roussiakis (AMPG), Prof. Mitylini et catalogue d’ossements fossiles recueillis à P Tassy (MNHN), Dr. S. Sen (MNHN), Prof. G. Rabeder Mitylinii, île de Samos, et déposés au Collège Gal- (PIUW), Prof. D. Nagel (PIUW), Dr. A. Currant (NHML) liard, à Lausanne. Georges Bridel 8c Cie Editeurs, ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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pp. 1-51, Lausanne. raphy in the Eastern Mediterranean. — Deinsea, G audry, A., 1861. Résultats des fouilles entreprises 10:343-371, Rotterdam. en Grèce sous les auspices de lAcadémie des Sci­ Koufos, G.D., 2006a. The late Miocene Vertebrate local­ ences. — Comptes Rendus Academie Sciences Paris, ity of Perivolaki, Thessaly, Greece. 4. Carnivora. — 52:722-724, Paris. Palaeontographica, Abt. A, 276:39-74, Stuttgart. G audry, A. &L artet, E.A.I.H., 1856. Sur les résultats Koufos, G.D., 2006b. Palaeoecology and chronology of des recherches paléontologiques entreprises dans the Vallesian (late Miocene) in the Eastern Mediterra­ lAttique sous l’auspice de lAcademie. — Comptes nean region. — Palaeogeography, Palaeoclimatology, Rendus Académie Sciences Paris, 43:271-274, Palaeoecology, 234:127-145, Amsterdam. Paris. Koufos, G.D., 2006c. The large from the Mi- G audry, A., 1862-67. Animaux fossiles et géologie de ocene/Pliocene locality of Silata, Macedonia, Greece lAttique. Editions Savy, Paris, pp.215-218. with implications about the Latest Miocene palaeo­ G ervais, R, 1859. Zoologie et Paléontologie française, ecology. — Beiträge zur Paläontologie, 30:293-313, 2nd édition, Paris. Wien. H ensel, R.F., 1862. Uber die Reste einiger Säugetier­ Koufos, G.D., 2006d. The Neogene mammal localities arten von Pikermi in der Münchener Sammlung. — of Greece: faunas, chronology and biostratigraphy. Monatsberichte der Akademie der Wissenschaften, — Hellenic Journal of Geosciences, 41(1):183-214, 27:560-569, München. Athens. Howell, F.C. ôcP etter, G., 1985. Comparative obser­ Koufos, G.D., this volume. The Late Miocene Mammal vations on some middle and upper Miocene hyaenids Faunas of the Mytilinii Basin, Samos Island, Greece: genera: Percrocuta K retzoi, Allohyaena K retzoi, New Collection. 1. History of the Samos Fossil Mam­ Adcrocuta K retzoi (Mammalia, Carnivora, Hyae- mals. — Beiträge zur Paläontologie, 31:1-12, Wien. nidae). — Geobios, 18:419-476, Lyon. Koufos, G.D. & M elentis, J., 1982. The Late M i­ K aya, T., G eraads, D. &c Tuna, V. 2005. A new late ocene (Turolian) mammalian fauna of Samos Island Miocene mammalian fauna in the Karaburun Pe­ (Greece). Study of the collection of Palaeontological ninsula (W Turkey). — Neues Jahrbuch für Geologie Museum of Mytilinii, Samos. 1. Carnivora. — Sci­ und Paläontologie, Abhandlungen, 236(3):321-349, entific Annals, Faculty of Physics and Mathematics, Stuttgart. University of Thessaloniki, 22:175-193, Thessalo­ K ittl, E., 1887. Beiträge zur Kenntniss der fossilen niki. Säugethiere von Maragha in Persia. I. Carnivora. Koufos, G.D., Kostopoulos, D.S., & Vlachou, T.D., — Annalen des Naturhistorischen Hofmuseums, this volume. The Late Miocene Mammal Faunas 2:317-338, Wien. of the Mytilinii Basin, Samos Island, Greece: New K ostopoulos, D., Sen, S. & K oufos, G.D., 2003. Collection. 16. Chronology. — Beiträge zur Paläon­ Magnetostratigraphy and revised chronology of the tologie, 31:397-408: Wien. late Miocene mammal localities of Samos, Greece. — Koufos, G.D., Kostopoulos, D.S., V lachou, T.D. International Journal of Earth Sciences, 92:779-794, &c Sylvestrou, I.A., 2004. Reconsideration of the Wien. Mytilinii fossiliferous basin, Samos, Greece. — 5th K ostopoulos, D.S., Koufos, G.D., Sylvestrou, I.A., International Symposium on Eastern Mediterra­ Syrides, G.E. & Tsompachidou, E., this volume. nean Geology, Thessaloniki, April 2004, Abstracts, The Late Miocene Mammal Faunas of the Mytilinii 1:326-329, Thessaloniki. Basin, Samos Island, Greece: New Collection. 2 . Koufos, G.D., Sen, S., K ostopoulos, D.S., Sylvestrou, Lithostratigraphy and Fossiliferous Sites. — Beiträge I.A. & Vlachou, T.D., 2006. The late Miocene zur Paläontologie, 31:13-26, Wien. Vertebrate locality of Perivolaki, Thessaly, Greece. K oufos, G.D., 1979. Preliminary report of the study of 10. Chronology. — Palaeontographica, Abt. A, mammalian fauna (Carnivora, Proboscidea, Peris- 276:185-200, Stuttgart. sodactyla) of Axios Valley (Macedonia - Greece). Koufos, G.D., Syrides, G.E., K ostopoulos, D.S., — Proc. VIIth Int. Cong. Medit. Neogene, Athens, K oliadimou, K.K., Sylvestrou, J.A., Seitanidis, 1979. — Annales géologiques de Pays Helléniques, G.C. & Vlachou, T , 1997. New excavations in the Hors series, 2:631-635, Athens. Neogene mammalian localities of Mytilinii, Samos Koufos, G.D., 1980. Palaeontological and stratigraphi- Island, Greece. — Geodiversitas, 19(4):877-885, cal study of the Neogene continental deposits of the Paris. basin of Axios river. — Scientific Annals, Faculty of K retzoi, M., 1938. Die Raubtiere von Gombaszög nebst Physics and Mathematics, University of Thessaloniki, einer Übersicht der Gesamtfauna. (Ein Beitrag zur 19(ll):l-322. Stratigraphie des Altquartärs). — Annales Musei K o u f o s , G.D., 2 0 0 0 . Revision of the late Miocene car­ Nationalis Hungarici, 31:88-157, Budapest. nivores from the lower Axios Valley. — Münchner Kurten, B., 1982. Status of the fossil hyaenids Ictith- Geowissenschaftliche Abhandlungen, (A),3 9 :5 1 - 9 2 , erium viverrinum and Thalassictis robusta (Mamma­ München. lia). — Zeitschrift für geologische Wissenschaften, Koufos, G.D., 2003. Late mammal events and biostratig­ 10:1009-1018, Berlin. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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M e l e n t is , J.K., 1967. Studien über fossile Vertebraten Sen, S., Koufos, G.D., Kondopoulou, D. & Bonis, L. Griechenlands. 19. Die Pikermifauna von Halmyro- de, 2000. Magnetostratigraphy of the late Miocene potamos (Euböa, Griechenlands). I. Teil: Odontologie continental deposits of the lower Axios Valley, Mac­ und Kraniologie. — Annales Géologiques des Pays edonia, Greece. — [in:] K oufos, G.D. & Ioakim, Helléniques, sér. 1, 18:283-411, Athens. Ch. (eds). Mediterranean Neogene cyclostratigraphy NOW, 2007-08. Neogene of the Old World, database with in marine-continental deposits. — Bulletin of the the Neogene localities and their faunal lists. — www. Geological Society of Greece, Athens, sp. publ., helsinki.fi/science/now/database.htm 9:197—206, Athens. Ozansoy, R, 1965. Etude des gisements continentaux et Solounias, N., 1981. The Turolian fauna from the island des Mammifères du Cénozoique de Turquie. — Mé­ of Samos, Greece. — Contributions Vertebrate Evolu­ moires de la Société Géologique de France, 102:1-92, tion, 6:1-232. Paris. Symeonidis, N., 1978. Ein Schädel von M etailurus Petter, G., 1976. Etude d’un nouvel ensemble de petits parvulus (Hensel) aus Pikermi (Attica, Griechen­ carnivores du Miocène d’ Espagne. — Géologie land). — Annales Géologiques des Pays Helléniques, Méditerranéenne, 3(2):135-154, Paris. 29:698-703, Athens. Pilgrim, G.E., 1931. Pontian Carnivora of Europe. Thenius, E., 1951. Zur odontologischen Charakteristik — British Museum Natural History, pp. 1-174, von “Felis leiodon” aus Pikermi (Griechenland). — London. Neues Jahrbuch für Geologie und Paläontologie, 3: Roth, J. & W agner, A., 1854. Die fossilen Knochenüber­ 88-96, Stuttgart. reste von Pikermi in Griechenland. — Abhandlungen Torre, D., 1988. Plioviverrops faventinus n.sp. a new der Bayerischen Akademie der Wissenschaften, carnivore of the late Messinian age. — Bolletino Math.-Phys. KL, 7:371-464, München. della Société Paleontología Italiana, 28:323-327, Roussiakis, S., 1996. Contribution to the study of the Modena. mammalian findings in the type locality of Pikermi. W agner, A., 1848. Urweltliche Säugetier-Ueberreste aus — PhD thesis, University of Athens, pp. 1-259. (in Griechenland. — Abhandlungen der Bayerischen Greek, unpublished). Akademie der Wissenschaften, 2. KL, 5(2):155-170, Roussiakis, S., 2001. Metailurus major Zdansky, 1924 München. (Carnivora, Mammalia) from the classical locality of W erdelin, L., 1988. Studies of fossil hyaenas: the genera Pikermi (Attica, Greece). — Annales de Paléontolo­ Thalassictis G ervais ex Nordmann, Palhyaena G er­ gie, 87(2):119-132, Paris. vais, Hyaenictitherium K retzoi, Lycyaena H ensel Roussiakis, S., Theodorou, G.E. ôcIliopoulos, G., 2006. and Palinhyaena Qiu, Huang & Guo. — Zoological An almost complete skeleton ofMetailurus parvulus Journal of Linnean Society, 92:211-265, London. (Carnivora, Felidae) from the late Miocene of Kerassia W erdelin, L. & Solounias, N., 1991. The hyaenidae: (Northern Euboea, Greece). — Geobios, 39:563-584, taxonomy, systematics and evolution. — Fossil and Lyon. Strata, 30:1-104, Oslo. Schmidt-K ittler, N., 1976. Carnivores from the Neo­ Z dansky, O., 1924. Jungtertiäre Carnivoren Chinas. — gene of Asia Minor. — Palaeontographica, Abt. A, Paleontológica Sinica, C, 2:38-45, Beijing. 155:1-131, Stuttgart. Z hang, Z.P., G entry, A.W., K aakinen, A., Liu, L., Semenov, Y.A., 1989. Iktiteri i morfologicheski skhodnye LunkkaJ .R , Q iu, Z.D., Sen, S., Scott, R.S., W er­ hieni neogenaUSSR. (Ictitheres and morphologically delin, L., Z heng, S. ScFortelius, M., 2002. Land related hyaenas from the Neogene of USSR). — Nau- mammal sequence of the late Miocene of China: new kova Dumka, Kiev, pp. 178 (in Russian, with English evidence from Lantian, Shanxi Province. — Verte- summary). brata PalAsiatica, 40(3):165-176, Beijing. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE 1

Parataxidea maraghana, M ytilinii-lA (MTLA), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Skull, M TLA-283; a. right lateral, b. left lateral, c. dorsal, and d. occlusal view.

Fig. 2. Skull, M TLA-283; left tooth row, occlusal view.

Fig. 3. Skull, M TLA-465; a. dorsal, b. left lateral, and c. occlusal view.

Fig. 4. Skull, M TLA-465; left tooth row, occlusal view.

Fig. 5. Left m l, M TLA-7; a. buccal, b. lingual, and c. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE2

Plioviverrops orbignyi, Mytilinii-lB (MTLB), Samos, Greece, middle Turolian (MN12 ).

Fig. 1. Skull, M TLB-170; a. dorsal, b. right lateral, c. left lateral, d. occlusal, and e. occipital view.

Fig. 2. Skull, M TLB-170; both tooth rows, occlusal view.

Protictitherium crassum, Mytilinii-4 (MLN), Samos, Greece, middle Turolian (MN 12).

Fig. 3. Left lower canine, MLN-26; a. buccal, and b. lingual view. K oufos , G.D., Carnivora, G.D.,

f'V U -fi'llO ©Verein zurFörderungderPaläontologieamInstitutfürPaläontologie,GeozentrumWien PLATE2 95 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE3

Protictitherium crassum, Mytilinii-4 (MLN), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Left mandibular corpus of the mandible, M LN -42; a. buccal, b. lingual; and c. lateral view.

Fig. 2. Right mandibular corpus of the mandible, M LN-42; a. buccal, b. lingual; and c. lateral view.

Hyaenictitherium wongii, M ytilinii-lA (MTLA), Samos, Greece, middle Turolian (MN 12).

Fig. 3. Frontal part of the skull, M T L A -1; a. left lateral, b. right lateral, c. frontal, and d. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE4

Hyaenictitherium wongii, M ytilinii-1 A (MTLA), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Skull, MTLA-200; a. dorsal, b. left lateral, c. right lateral, d. occlusal, and e. occipital view.

Fig. 2. Skull, M TLA-200; both tooth rows; occlusal view.

Fig. 3. Left mandibular fragment with c-ml, M TLA-2; a. buccal, b. lingual, and c. occlusal view.

Fig. 4. Right mandibular fragment with c-ml, M TLA-237; a. buccal, b. lingual, and c. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE5

Hyaenictitherium wongii, M ytilinii-lA, B, (MTLA, MTLB), Samos, Greece, middle Turolian (MN12 ).

Fig. 1. Left mandibular fragment with p3-m l, M TLA-266; a. buccal, b. lingual, and c. occlusal view.

Fig. 2. Right mandibular fragment with il-p3, M TLA-468; a. buccal, b. lingual, and c. occlusal view.

Fig. 3. Left mandibular fragment with p4-m l, M TLB-1; a. buccal, b. lingual, and c. occlusal view.

Fig. 4. Left p3 of the mandibular fragment M TLB-1; a. buccal, b. lingual, and c. occlusal view.

Fig. 5. Right mandibular fragment with p4-m l, M TLB-100; a. buccal, b. lingual, and c. occlusal view.

Fig. 6. Left hemimandible with i3-m2, M TLB-171; a. buccal, b. lingual, and c. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE6

Hyaenictitherium wongii, Mytilinii-lC, 4 (MTLC, MLN), Samos, Greece, middle Turolian (MN12 ).

Fig. 1. Right mandibular fragment with i2-m l, M TLC-18; a. buccal, b. lingual, and c. occlusal view.

Hyaenictitherium cf. w ongii, Mytilinii-lC, 4 (MTLC, MLN), Samos, Greece, middle Turolian (MN12 ).

Fig. 2. Isolated left P2, MLN-27; a. buccal, b. lingual, and c. occlusal view.

Adcrocuta eximia, possibly Mytilinii-1 (?MTL), Samos, Greece, middle Turolian (MN 12).

Fig. 3. Frontal part of the skull with P2-M 1 dex and P2-P3 sin, PM M S-70; a. right lateral, and b. occlusal view.

Fig. 4. Left mandibular fragment with pl-p4, PM M S-70a; a. buccal, b. lingual, and c. occlusal view.

Fig. 5. Left upper canine, PM M S-71; a. buccal, and b. lingual view.

Adcrocuta eximia, M ytilinii-lA (MTLA), Samos, Greece, middle Turolian (MN12 ).

Fig. 6. Isolated p2 sin, M T L A -6; a. buccal, b. lingual, and c. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnívora, 103

PLATEÓ ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

104 Beitr. Palaont., 31, Wien, 2009

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MetailurusparvuluSy M ytilinii-lA (MTLA), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Right m l, M T LA, 290; a. buccal, b. lingual, and c. occlusal view.

Fig. 2. Left m l, M T LA, 466; a. buccal, b. lingual, and c. occlusal view.

Fig. 3. Skull, M TLA-234; a. dorsal, b. right lateral, and c. occlusal view.

Fig. 4. Skull, M TLA-234; right tooth row, occlusal view.

Fig. 5. Skull, M TLA-234; left P4; occlusal view.

Fig. 6. Right mandibular fragment with p3, M TLA-2325; a. buccal, b. lingual, and c. occlusal view.

Machairodusgiganteus, possibly Mytilinii-1 (?MTL), Samos, Greece, middle Turolian (MN 12).

Fig. 7. Fragment of the upper canine, PM M S-69; a. buccal, and b. lingual view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Carnivora. 105

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