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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 Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection - 6. Tubulidentata 107-125 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Beitr. Paläont., 31:107-125, Wien 2009

Tlie Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 6. Tubulidentata

b y

George D. Koufos*)

K o u f o s , G.D., 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 6. Tubulidentata. — Beitr. Palaont., 31:107-125, Wien.

Abstract den neuen Geländearbeiten in Samos befanden sich unter den zahlreichen Fossilresten einige neue Orycteropodidae. The orycteropodids are known from Samos since the last Diese stammen aus den Fundstellen Mytilinii-lA, B decades of 19th century when the first mammalian fossils (M TLA, MTLB) der Adrians Rinne und sind ins mittlere were found. Since 1941 there were alsmost undescribed Turolium zu stellen. Die magnetostratigraphischen Er­ but at that time Colbert studied the material of Brown’s gebnisse legen ein Alter zwischen 7,1-7,0 Millionen Jahre collection in the American Museum of Natural History nahe. Das neue Material wurde mit dem bereits aus Samos and gave an extensive and detailed description of the whole sowie aus Eurasien und Afrika Bekannten verglichen. Es skeleton. During the new field campaigns in Samos we gehört zu der gut bekannten A rt Oryctoropus gaudryi, die collected a great amount of fossils and among them several man aus dem Turolium des östlichen Mittelmeeres kennt. orycteropodids. They were found in the fossiliferous sites In dieser Arbeit wird auch die stratigraphische Verbreitung Mytilinii-1 A, B (MTLA, MTLB) of the Adrianos ravine der Orycteropodidae diskutiert. and they are dated to middle Turolian; their magneto- stratigraphic record suggests an age from 7.1-7.0 Ma. The Schlüsselworte: Obermiozän, Samos, Griechenland, new material is described and compared with the old one Mammalia, Tubulidentata, Systematik. trom Samos as well as with Eurasian and African one. It belongs to the species Orycteropus gaudryi a well known taxon from the Turoiian of Eastern Mediterranean. The Introduction stratigraphic distribution of the orycteropodids in the Old World is also discussed in this article. Samos Island is situated in the eastern Aegean Sea and is well known for its late Miocene mammal localities since Keywords: Late Miocene, Samos, Greece, Mammalia, the second half of the 19th century, when Forsyth Major ubulidentata, Systematics. found and collected mammalian fossils in the island. Later, several palaeontologists and fossil dealers excavated and collected fossils from Samos. The fossils recovered from Zusammenfassung Samos localities are numerous, but they are dispersed in several museums and collections all over Europe and ^ (V r ? podidae sind aus Sam° s sdt dem Ende des U.S.A. The most important collections are those of ,rJ ,r -1Ur* erts Staunt, als die ersten fossilen Säugetiere Lausanne, London and New York. A small collection , v pri ^ ei) Wurden. Bis 1941 waren sie allerdings un- from Samos was also assembled at the A M P G by Prof. [p, ,.Vn j "n,’ 1S Ecddert das Material aus der Sammlung T. Skoufos. Following the last excavations of B. Brown •:'d d is ^meiacan Museum ofNatural History studier- in 1925, Prof. J.K. Melentis excavated in Adrianos ravine ganZe Skelett im Detail beschrieb. Während in 1963 and made a small collection. This was on display in the city hall of Mytilinii on the Island of Samos for a long time until being transferred to the NHM A. In 1985 Prof. J.K. Melentis and the author excavated in Adrianos ravine and more exactly in the fossiliferous site Mytilinii- sntnf(^0yFOS,AristotleUniversityoflhessaloniki. lA (MTLA). The collected material (mainly giraffids and (; H->4 ! Eaboratory of Geology and Palaeon- hipparions) is included in the new collection. essaloniki, Greece. e-mail:koufos@geo. In 1994, a group of palaeontologists from the Laboratory of Geology and Palaeontology of the Aristotle University ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

108 Beitr. Palàont., 31, Wien, 2009

Figure 1: Composite skeleton of Orycteropusgaudryi from Samos, housed at the AMNH. It con­ sists of the skull and mandible AM N H -SAM -20694 and the postcranial elements AMNH- SAM-22762. After C olbert (1941:fig. 19).

of Thessaloniki, leaded by the author, started a new series MNHN = Museum National d’Histoire Naturelle, Paris of excavations in Samos mammal localities. During these MTLA = Mytilinii-lA field campaigns, a great amount of fossils have been un­ MTLB = Mytilinii-1 B earthed and are now housed in the NH M A. More details MTA = Geological Survey of Turkey about the history of the Samos mammal fossils and their NHMA = Natural History Museum of the Aegean, Mytilinii, collection are given in the first chapter of this volume Samos (Koufos, this volume). The fossiliferous localities are lo­ NHML = Natural History Museum of London cated in the Mytilinii area situated in the eastern Neogene NHMW = Naturhistorisches Museum Wien basin of the Samos Island. The stratigraphy of the Mytilinii PNT = Pentalophos-1, Axios valley, Macedonia, Greece deposits has been studied by various authors; for more data SAM = Samos (old collections) about the stratigraphy and localities seeK ostopoulos et al. (this volume). The specimens from these museums and institutions, ex­ Among the collected material from Samos localities cept NHM A, are reported in the text by the abbreviation there are several specimens of aardvarks. The aardvarks of the museum or institute, the locality and their serial are referred in the first faunal lists of Samos (Forsyth number. M ajor, 1888,1894; A ndrews, 1896). The known mate­ rial is abundant and includes some nice specimens, like a mounted skeleton made on the basis of various individuals Palaeontology and housed in the AM NH (Fig. 1). In comparison to the other Turolian localities of Eastern Mediterranean, the Order Tubulidentata Huxley, 1872 fossil aardvark collection from Samos is the largest. In Family Orycteropodidae G ray, 1821 this respect, the new collected specimens add more data about this rare animal. The studied aardvark material Genus Orycteropus G eoffroy, 1796 was found in the Adrianos ravine (locality Mytilinii-1, MTL) and more precisely in the fossiliferous sites A and Orycteropus gaudryi Forsyth M ajor, 1888 B (Kostopoulos et al., this volume). The fauna of both (Plates 1, 2) sites suggests a middle Turolian age, MN 12. The mag- netostratigraphic record indicates an age of 7.1-7.0 Ma for Localities: Mytilinii-lA, B (MTLA, MTLB), Adrianos M T L A and MTLB. More details about the age are given ravine, Mytilinii Basin, Samos, Greece. in K ostopoulos et al. (2003) and Koufos et al. (2004, Age: Middle Turolian, MN 12 (late Miocene); 7.1-7.0 Ma. this volume). Material: MTLA: Partial skull and right mandibular fragment, Abbreviations: M TLA-115; partial skull, M TLA-240; left M l, M TLA- AH G = Ahmet Aga or Dragi or Prokopion, Evia Island, 239; left mandibular ramus with m3, MTLA-116; Greece right mandibular ramus with p4-m3, MTLA-280; left AMNH = American Museum of Natural History mandibular ramus with m2-m3, MTLA-306; left ml, AM PG = Athens Museum of Geology and Palaeontology, MTLA-18; left m 2, M TLA-17. University of Athens MTLB: Right mandibular ramus with p2-m3, MTLB- B = Breadth 49. BHA = Bou Hanifia, Algeria Measurements: The measurements of the material are CA = Çandir given in Tabs 1-3. KTB = Kemiklitepe-B, Turkey Description: KTD = Kemiklitepe-D, Turkey Partial skull M T LA -115. The frontal part of the skull and L = Length a large part of the muzzle are preserved (PI. 1, fig. 1). The MGL = Musée de Géologie de Lausanne palate is long and relatively narrow as in all Orycteropus. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 109

Table 1: Comparative cranial measurements ofOrycteropus from various localities. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

110 Beitr. Palaont., 31, Wien, 2009

Table 2: Comparative mandibular measurements ofOrycteropus from various localities; the measurements of 0.sen i were taken from van der Made (2003). ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. I ll

Table 3: Dental measurements of the studied material ofOrycteropus gaudryi from Mytilinii (Samos). ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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Figure 2: Comparison of the Mytilinii cranial material of Orycteropus gaudryi with other specimens from Samos. a. partial cranium MTLA-115 in dorsal, lateral (right), and ventral view; b. partial skull MTLA-240 in dorsal, lateral (left), and ventral view; c. skull NHMW-SAM-A.4759 in dorsal, lateral (right), and ventral view; d. skull NHMW-SAM-A.4760 in dorsal, lateral (right), and ventral view.

The nasals are elongated and wide and they are separated the distal lobe of the M 3. The anterior border of the post­ by a shallow groove running sagittally between them, palatine groove is situated at M3 but its anterior part is at along the suture. The frontal region is bulging (PL 1, fig. the level of the distal lobe of the M 2. The greater part of lc). The preserved dentition consists of the tooth row P l- the nasals is broken but the groove following the suture P4 sin and the isolated P3 and M 3 dex (PI. 1, fig. Id). The between them is slightly deeper than that of MTLA- tooth row is curved after the P4. There are small diastemas 115 (PI. 1, fig. 3c). The frontals are bulging more than between the premolars. The presence of the canine is not in MTLA-115. The crista facialis is also stronger than ascertained as the maxilla in this area is broken on both in MTLA-115. The skull is slightly compressed dorsov- sides and it is difficult to recognize the presence or absence entrally but this deformation cannot justify the greater of an alveolus. The premolars are small and narrow except width of the skull. The preserved dentition consists of the for the P4, which is wide. The P i and P2 have one lobe and tooth row P2-M3 sin, and the isolated P2 and M3 dex, their occlusal surface shows a single wearing facet inclined but most of the teeth are badly preserved. The tooth row distally. The P3 and P4 have also one lobe, but their oc­ is slightly curved and is characterized by the dominance of clusal surface depicts a sharp angle between their mesial the M 2. This character is not distinct however, as the M 2 and distal wearing facets (PI. 1, fig. Id). The M l and M 2 is partially broken. There is no trace of a canine and the are absent but the alveolus of the M 2 is the largest of the diastemas between the premolars are absent. The anterior tooth row. The M 3 is smaller, triangular-shaped with deep molars are bilobed, 8-shaped whereas the M3 is smaller buccal groove and almost straight lingual wall. and triangular-shaped. The morphology of the other teeth Partial skull MTLA-240. It represents more or less the is similar to that of MTLA-115, except for the slenderer same part of the skull as MTLA-155; however a larger P4 and the smaller M3. part of the distal palate and a shorter part of the muzzle Mandibular fragment MTLA-115. It is the right man­ are preserved (PI. 1, fig. 3). The palate is long and narrow dibular corpus lacking the ascending ramus, but preserving but slightly wider than that of MTLA-115 (Tab.l). The p3, ml-m3; the mesial lobe of the m l is absent. The alveoli anterior border of the choanae is situated at the level of of the other premolars are distinct, whereas it seems to ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 113

Figure 3: Comparison of the M ytilinii cranial material of Orycteropus gaudryi with other specimens from Samos and Evia. a. cast of the skull AMNH-SAM-20560 in dorsal, lateral (right), and ventral view; b. cast of the skull AMNH-SAM-20562 in dorsal, lateral (right), and ventral view; c. skull NHML-SAM-M.5690 in dorsal, and lateral (left) view; d. partial skull and endocast NHML-AHG-M.8938 in ventral view. be no alveolus for a canine (PI. 1, fig. 2). The mandibular lobe wider than the mesial one in the anterior molars. The corpus is elongated with a height increasing from the p3 to m2 is the largest tooth of the tooth row, whereas the m3 m3. The ventral border of the mandibular corpus is slightly is smaller than in the other specimens, triangular-shaped concave (PI. 1, fig. 2a). There are no large diastemas be­ with narrow distal lobe. Both the lingual and buccal tween the premolars. The molars are bilobed and 8-shaped groove of the molars is well developed but the lingual one with deep grooves. The mandibular corpus is thicker in the is slightly deeper. In the m l and m2, the occlusal surface of area of the molars, showing its greatest thickness at the the mesial lobe has two wearing facets, a narrow anterior m2-m3 level (PI. 1, fig. 2c). The m2 is the largest tooth of and a larger posterior forming an angle between them. the tooth row and the m3 has a narrow distal lobe. The molars are larger than in the other specimens except Mandibular fragment M T L A -116. The mandibular for the m3; in this feature seems to be similar with the corpus and a small part of the ascending ramus are pre­ skull M TLA-240. served (PI. 2, fig. 1). A single and large mental foramen Mandibular fragment MTLA-306 . It is a small mandibu­ is situated almost in front of the pi. The ascending ramus lar fragment, bearing badly preserved m2-m3 sin (PI. 2., forms a wide angle (-123°) with the mandibular corpus. fig. 3). The mandibular corpus is deep and the m3 is as The other features of the mandible, as well as that of the small as in MTLA-280. sole preserved m3 are similar to those of MTLA-115. Mandibular fragment MTLB-49. It is a mandibular Mandibular fragment M T L A -280. It bears the tooth fragment with well preserved p2-m3 dex (PI. 2, fig. 4). row p4-m3 (PI. 2, fig. 2). The mandibular corpus of The size and morphology of the mandibular corpus, as M i LA-280 and MNHN-KTB-94 is the deepest of the well as the dental morphology, are similar to those of comparative set (Tab. 2). The anterior border of the mas­ M TLA-115, 116. seteric fossa is situated at the level of the distal border of the Discussion: The systematics of the aardvarks is still dis­ tn j. The morphology of the mandibular corpus is like that cussed and there are several debates (Patterson, 1978; °f I TTLA-115. The molars are 8-shaped with the distal P ickford, 1975; L ehmann, 2007). Certainly the aim ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

114 Beitr. Paläont., 31, Wien, 2009

Orycteropus, Upper Teeth Figure 4: a. Logarithmic ratio di­ agram comparing the upper teeth of the Mytilinii material with Orycteropusgaudryi and O ryctero­ pus pottieri. b. Logarithmic ratio diagram comparing the upper teeth of the Mytilinii material with various orycteropodids.

O. afer, Africa (LEHMANN, 2004a) ■ O. gaudryi, MTLA-240 O. gaudryi, MTLA-115 - • — O. gaudryi, NHMW-SAM-A.4760 (orig, meas.) O. gaudryi, NHMW-SAM-A.4759 (orig. meas.) -O — O. gaudryi, BMNH-SAM-M.5690 (orig, meas.) O. gaudryi, AMNH-SAM-20694 (COLBERT, 1941) -O — O. gaudryi, AMNH-SAM20560 (COLBERT, 1941) O. gaudryi, AMNH-SAM-20561 (COLBERT, 1941) -%— O. gaudryi, BMNH-AHG-M.8938 (orig, meas.) O. gaudryi, MNHN-KTB-95, 97, Kemiklitepe (SEN, 1994) -----O. pottieri, LGPUA-PNT-127, Pentalophos (BONIS etal., 1994)

Orycteropus, Upper Teeth of this paper is not to discuss these issues but it is necessary to give some information. The order Tubulidentata consists of the sole family Oryctero- podidae, which according to Patterson (1975, 1978) is separated in two sub-families Plesiorycteropodinae and Orycteropodinae. The sta­ tus of the first sub-family is debated (Patterson, 1975, ------O. afer, Africa (LEHMANN, 2004a) ■ MTLA-240 1978; M acPhee 1994). The — A — MTLA-115 — B — O. mauritanicus, Bou-Hanifia, Algeria, (ARAMBOURG, 1959) — 0— O. depereti, Perpignan, France (HELBING, 1933) —is— O. browni, Pakistan (PICKFORD, 1978) second sub-family includes — O— M. africanus, Kenya (PICKFORD, 1975) — X — O. abundulafus, Tchad (LEHMANN, 2005) the following genera: M yo- — X — O. djourabensis, Tchad (LEHMANN, 2004b) rycteropus, Leptorycteropus and Orycteropus. The last one includes all the Eurasian Vryssoula (Solounias, 1981); but this is not sure as the aardvarks, as well as the modern O. afer. However, the author mentions that it is impossible for Forsyth Major to taxomonic status of the Eurasian material of the genus be in Samos and not to visit Adrianos ravine, the richest Orycteropus is doubted (Lehmann, 2004). The last author fossiliferous area of the Mytilinii Basin. The morphology suggests that the Eurasian aardvarks belong to another of these skulls (narrow palate, shape of the tooth row, genus, but he did not give a name. In the meantime, the dental morphology, dominance of the M2 and m2) as name Orycteropus still remains and will be used in the well as the size are similar to both studied cranial frag­ present paper. ments (Figs 2, 3, Tab. 1). Despite their similarities, the As it was mentioned in the introduction, Orycteropus specimen M T L A -240 seems to be slightly larger than gaudryi is known from Samos since the last decades of the other Samos skulls (Figs 2, 3, Tab.l). Likewise, the the 19th century (Forsyth M ajor, 1888, 1894). The proportions of the teeth are similar to the ones compared comparison of the present material with the old one from to but the M T L A -240 teeth are among the largest (Fig. Samos is quite useful. For this reason, the new material 4). The morphology of the studied skulls and mandibles was compared to the old one housed at the N M H W and fits quite well with those o f the Samos material housed NHM L, as well as to some casts of the AM N H collec­ at the AM N H and described by C olbert (1941). The tion stored in the MNHN. Two skulls of 0. gaudryi from comparison of the studied material with the casts from Samos are stored in the M N H W and one in the NHML; the AMNH indicates similar morphology and a slightly their origin is unknown as there are not locality indica­ larger size (Fig. 3; Tab. 1). Likewise, the proportions of tions. Those o f NH M W (N H M W -SA M -A.4756,4760) the teeth of the AM N H material are very similar to those have labels referring that they were purchased, while that of the studied specimens (Fig. 4). According to these ob­ of NHML (NHML-SAM-M.5690) belongs to Forsyth servations, there is noticeable size variation in O.gaudryi Major’s collection, which was bought by the Museum. It from Samos. This variation is probably intra-specific as is reported that this collection comes from a site named no sexual dimorphism is observed in the recent aardvarks ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 115

Orycteropus, Lower Teeth Figure 5: a. Logarithmic ratio di­ agram comparing the lower teeth of the Mytilinii material with Orycteropusgaudryi and Oryctero- puspottieri. b. Logarithmic ratio diagram comparing the lower teeth of the Mytilinii material with various orycteropodids.

O. afer Africa (LEHMAN, 2004a) O. gaudryi» MTLA, mean O. gaudryi, MTLB O. gaudryi, Kemiklitepe (SEN, 1994) O. pottieri, Pentalophos-1 (BONIS et ah, 1994) O. pottieri, Middle Sinap (LEHMAN, 2004a)

Orycteropus, Lower Teeth

A small mandibular frag­ ment with p4-ml is known from the Turolian locality of D ytiko-1 (DTK) of Axios Valley, Macedonia, Greece (Bonis et al., 1994). The size of its teeth is similar to that of the Mytilinii material. Recently, Orycteropus was recognized in the locality of LP/3 LP/4 LM/1 LM/2 LM/3 BP/3 BP/4 BM/1 BM/2 BM/3 Kerassia, Evia Island (Theo-

------O. afer, Africa (LEHMAN, 2004a) ■ O. gaudryi, MTLA mean dorou et al., 2003) but no A O. gaudryi, MTLB —I------O. seni, Candir (VAN DER MADE, 2003) description of the material O. maurrtanicus, Algeria (ARAMBOURG, 1959) - A — O. browni, Pakistan (PICKFORD, 1978) - X — O. abundulafus, Chad (LEHMANN et al., 2004) - X — O. djourabensis, Chad (LEHMANN et al., 2004) was given till now. - 0 — O. chemeldoi, Kenya (PICKFORD, 1975) - O ^ M . africanus, Rusinga (PICKFORD, 1975) Orycteropus gaudryi is also recorded from the Turkish (Shoshani et al. 1988; L ehmann, 2004). Orycteropus locality of Kemiklitepe (Sen, 1994). The morphology gaudryi is also reported from three other Greek locali­ and dimensions of the present material is similar to that ties (Achmet Aga, Dytiko, Kerassia). A badly preserved of the Kemikilitepe (KTA-B) material (Fig. 4a). It is skull (NHML-AHG-M.8938) is known from the late also reported, without description, from the Turkish Miocene locality of Achmet Aga (Drazi or Prokopion) localities of Kufuk Qekmece, Mahmutgazi and Kayadibi of Evia Island (W oodward, 1901). The skull is very de­ (Sen, 1994). Besides Orycteropus gaudryi, several other formed and weathered (Fig. 3d). Because Woodward’s ar­ tubulidentate species are known from Eurasia and Africa. ticle is entitled “On the bone-beds of Pikermi, Attica and Their comparison with the studied material from Samos the similar deposits in Northern Euboea”, the specimen is given below: was wrongly considered to come from Pikermi. I have Orycteropus pottieri O zansoy, 1965. This taxon was seen the specimen at the NHM L and compared it with originally described from the Vallesian levels of the Mid­ the studied material. On the specimen, a label states that dle Sinap, Turkey (Ozansoy, 1965). Although Ozansoy it comes from the locality of Dragi from the Evia Island listed a mandible, a maxilla and some isolated teeth, he and it is clearly reported in the text ofW oodward (1901; described only the mandible. Later, additional material P- 485). The confusion of its origin gave rise to the inclu­ of O. pottieri was found in the Sinap (Sen, 1994; Bonis sion of Orycteropus in the Pikermi fauna. In the major et al., 1994). This taxon is also known by some maxillary collections from Pikermi, which I have seen in European and mandibular fragments from the Vallesian locality of museums as well as in A M P G , there is no evidence for Pentalophos-1 (PNT, Axios Valley, Macedonia, Greece) the presence ofOrycteropus in Pikermi. The preservation (Bonis et al., 1994). Recently, new material of O. pot­ of the skull NHML-AHG-M.8938 does not allow a tieri has been described from the Vallesian localities of comparison with the new material, but its dental dimen­ the Sinap Formation (Turkey), dated from 10.1-9.6 Ma sions are similar to those of the M ytilinii specimens and (Fortelius et al., 2003). The Mytilinii material differs falls into the range of variation of O. gaudryi (Fig. 4a). from O. pottieri by the larger upper teeth (Fig. 4b), the ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

116 Beitr. Palaont., 31, Wien, 2009

larger and deeper mandibular ramus (Tab.2 ), the absence lobe of the M 2 (PI. I; fig. 3d). Second, the origin of the of canines, the absence or very small diastemas between zygomatic arches is situated above the mesial lobe of the the upper and lower premolars, the longer and wider P4, M 2, whereas in O. gaudryi it begins above the distal lobe. the dominance of the M 2 and m2 in the tooth row, the In the studied skull M T LA-115, the zygomatic arches relatively deep labial and the shallow lingual grooves in originate above the M 3. Finally, a fragmentary skull of the upper molars. an aardvark has been discovered in the early Turolian Orycteropus seni T e k k a y a , 1 9 9 3 . It was originally de­ Bulgarian locality of Kocherinovo-1 and was referred to scribed from the middle Miocene locality of £andir (Tur­ as O. cf. browni (S p a s s o v et al., 2006). key), while later it was recognized in the middle Miocene African Orycteropodids. The aardvarks are slightly more Turkish locality of Pajalar (F o r t e l iu s , 1990; T e k k a y a , frequent in the Neogene of Africa and are represented by 1993; v a n d e r M a d e , 2003). Recently, comparable several species. The oldest known species isO. minutus specimens were found at the Loc. 64 of the Sinap, dated from the early Miocene of Kenya(P i c k f o r d , 1975). to Vallesian (F o r t e l iu s et al., 2003). In his description of The taxon is mainly represented by postcranial elements the £andir material, va n d e r M a d e (2003) noted that “a (P ic k f o r d , 1975) and the comparison with the Mytilinii smaller average size might be a justification for the species material is thus impossible. Another early Miocene A f­ O. seni”. A comparison between the mandible from £andir rican species is Myorycteropus africanus M a c I n n e s , 1956, (v a n d e r M a d e , 2003; pi. 1) and the Mytilinii ones does which is significantly smaller than the studied material not show any morphological difference; likewise, the size from Samos (Figs 4b, 5b). O. chemeldoi is known from the of the mandible and teeth of the Samos material and0. middle Miocene of Kenya. It was originally described on a seni is similar (Fig. 5, Tab. 2). hemimandible with p2-m3 from the localities of Ngorora Orycteropus depereti H e l b i n g , 1 9 3 3 . The taxon is based and Fort Ternan (P ic k f o r d , 1975). Its teeth are longer on a single skull, found in the early Pliocene (MN 15) and narrower than those of the Mytilinii ones (Fig. 5b). locality of Perpignan, France. The skull of O. depereti is There are small diastemas between the premolars like in larger than that of 0. gaudryi\ in comparison, the My­ several specimens of O. gaudryi, but the slenderer teeth tilinii skulls are smaller than the skull of O. depereti and of O. chemeldoi help distinguishing the two taxa. Some thus different (Tab. 1). The palate of 0. depereti is broader material (skull, mandibular fragments, postcranials) re­ than 0. gaudryi (A r a m b o u r g , 1939; P a t t e r s o n , 1975) ferred to as O. mauritanicus are known from the Vallesian and in this feature differs clearly from the studied skulls locality of Bou Hanifia, Algeria (A r a m b o u r g , 1959). 0 . of Samos. Although the skull of O. depereti is larger than mauritanicus differs from the studied material in having: O. gaudryi, its upper teeth are shorter than it, while their diastemas between the premolars, canines, as well as the breadth is close to it (Fig. 4b). On the other hand there M l and m l larger than the M 2 and m2 (Figs 3b, 4b), are some similarities between the Mytilinii skulls andO. respectively. Its teeth are larger than those of the studied depereti, like the bulging frontals, the elongated and curved material (Figs 4, 5). Two new species ofOrycteropus have tooth row, the absence of diastemas between the premolars recently been described from Chad(L e h m a n n et al., and the similar morphology of the molars. As there is only 2004, 2005, 2006). O. djourabensis comes from the early one skull of O. depereti, its comparison is limited and more Pliocene and is close in size to the recentO. afer from which material is necessary in order to ascertain its similarities it differs in having longer premolars, longer lower molars or differences from O. gaudryi; thus it could be referred and shorter-slenderer hands (L e h m a n n et al., 2004). It separately from O. gaudryi, at the moment. is different from the studied material in having a larger Siwaliks Orycteropus. The tubulidentata are reported from general size and teeth (Fig. 4, 5), and diastemas between the Siwaliks area, Pakistan by two species: O. browni the molars (L e h m a n n et al., 2004; fig. 2). Although the and O. pilgrimi (C o l b e r t , 1933). The first species was teeth of O. djourabensis are not well preserved(L e h m a n n originally described from a maxillary fragment with et al., 2004, fig. 2) the lingual groove of the upper molars is M 2-3, before additional material was found (C o l b e r t , deeper than that of the studied material. The other Chad­ 1933; P i c k f o r d , 1978). According to C o l b e r t (1933) O. ian species is O. abundulafus, dated between 7.0 Ma and pilgrim i was erected on a single right m2 but later a skull the Mio-Pliocene boundary (~5.3 Ma) and characterized was described under the same name(L e w i s , 1938). Cur­ by very robust teeth. This species shows similarities with rently, the two species are synonymized under the name the European O. gaudryi (L e h m a n n et al., 2005, 2006). O. browni (L e h m a n n , 2004, 2007). The Siwaliks mate­ In fact, the size of its teeth is very close to the one of the rial is poor and fragmentary and cannot allow extensive studied material from Samos, but the length of the upper comparison with the studied one. The dental size of the teeth is slightly smaller and the breadth slightly larger (Figs Siwaliks aardvark is clearly smaller than the Mytilinii 4, 5). These features are in agreement with the observa­ material (Fig. 4b) and can thus be distinguished from O. tion of L e h m a n n et al. (2005) that O. abundulafus has gaudryi. Two other differences with O. gaudryi are noticed robust teeth. The upper molars of the Mytilinii material from. The post-palatine foramen is situated between M2 are more symmetric showing more clear 8 -shaped oc­ and M 3 in O. browni, whereas in O. gaudryi it is situated clusal outline, whereas in O. abundulafus they are more at the level of the M3. Noticeably, in the partial skull asymmetric (L e h m a n n , et al. 2005; fig. 2). The PI and M T LA-240, the post-palatine foramen is situated at the pi are absent in O. abundulafus (L e h m a n n et al., 2005; level of the M 3 but its anterior part is situated at the distal fig. 2), whereas they are present in the Mytilinii material. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 117

Figure 6: Stratigraphic distribution of the orycteropodids in the Old World; see also text.

3. Biostratigraphy and Chronology of the recently recognized in the locality of Arrisdrift, Namibia Orycteropodids dated to 17.5 M a (Pickford, 2003). However van der M ade (2003) considers that O. minutus is a synonym of The orycteropodids are quite common in the Old World Myorycteropus africanus, an opinion which seems to be ac­ and they probably originated from Africa. A very small cepted by Lehmann (2006) too. The last author refers to aardvark, named Orycteropus minutus has been described O. minutus only the type material from Songhor, doubting from the early Miocene locality of Songhor, Kenya (Pick- also its fossorial way of life and myrmecophagous diet. ford, 1975); according to the last author the Songhor Thus, O. minutus appeared at the end of early Orleanian fauna is dated at 19.0 Ma. A similar age between 18.0-20.0 ELM A (-19.0 Ma), it is present during middle Orleanian Ma is also given for the Songhor locality in NOW (2007). (dashed line in Fig. 6) and probably existed at the end of The species is also reported from other Kenyan localities Miocene with a great gap. If we accept its synonymy with as Nyanza-12, 25 dated to early Orleanian (MN 3) ac­ M. africanus, then its stratigraphic span is restricted at the cording to the European Land M ammal Ages (ELMA), sole locality of Songhor at about 19.0 M a; its position is Mfwangano and Rusinga, dated to middle Orleanian, MN given by a star in Fig. 6. Its presence at the end of Miocene 4 (Pickford, 1975; NOW, 2008). It is worth mentioned is questionable and it is better not to give it in Fig. 6. here that the European land mammal ages application in Another Early Miocene African orycteropodid is M yo­ the African Neogene has not proved but it is used here in rycteropus africanus reported from Rusinga, Kenya (Mac order to have comparable data between the two continents. Innes, 1956) Later it was recognized in the localities Recent K-Ar data from Kenya suggest for the Rusinga Mfwangano and Kathwanga dated to middle Orleanian group a mean age of 17.9 M a but the available data for the ELM A, M N 4 (Pickford, 1975; NOW, 2008). As it was Mfwangano are not enough as no biotite with sufficient K referred above the K-Ar dating indicates an age of 17.9 M a content exists (D rake et ah, 1988). According to NOW for the Rusinga group, while the dating of Mfwangano (2007) the youngest appearance of O. minutus is known is not clear (Drake et ah, 1988). The stratigraphic distri­ from the Kenyan locality Nyanza-2 (Chantwara-34) bution of M. africanus spans in middle Orleanian, MN 4 dated to early Turolian ELM A, MN 13. The species was but if we accept the synonymy with O. minutus, then it is ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

118 Beitr. Palaont., 31, Wien, 2009

extended to early Orleanian, MN 3 (dashed line in Fig. (MN 5-6), the Tethyan Seaway re-opened but its opening 6). Besides the systematic problems the oldest traces of the was interrupted by several closures, which corresponds to orycteropodids have been recognized in the early Miocene several mammalian migration waves. During one of these of Africa at -19.0 Ma. migration waves, Orycteropus probably entered Eurasia. Orycteropus chemeldoi is the only middle Miocene oryctero- During this period, signs of migration waves were found podid known from Africa. It was originally described on the Greek island of Chios such as the suoidSanitherium from Ngorora Formation, Tugen Hills, Kenya, dated to schlagintweiti, the giraffid Georgiomeryx georgalasi and the latest Astaracian ELMA, MN 7+8 (P ic k f o r d , 1975). ctenodactylid Sayimys intermedius. The biochronological According to NOW (2008) the species is referred from data for the Chios fauna suggests a late Orleanian (MN the Kenyan sites Maboku and Fort Ternan dated to early 5) age, whereas the magnetostratigraphy indicates an age Astaracian ELM A, MN 5 and to latest Astaracian, MN of -15.5 Ma (K o u f o s et al., 1995; B o n is et al., 1997a, b, 7+8 respectively. The Maboku material consists of two 1998; L o p e z -A n t o n a n z a s et al., 2005). Moreover, the metapodial fragments which cannot be certainly identified, faunas of the Turkish localities, including O. seni, have rela­ making questionable the presence ofOrycteropus in this tions to the African ones confirming the above hypothesis. site (L e h m a n n , pers. comm.); for this reason its span at The above mentioned data suggest that the orycteropodids MN 5 is given by dashed line in Fig. 6. dispersed into Eurasia some 16.0-15.0 Ma ago and then During late Miocene two African orycteropodids are spread over the whole area. Besides Qandir and Pajalar, known. Orycteropus mauritanicus was found in the Vallesian a form referred to O. cf. seni is known from the Turkish locality of Bou Hanifia, Algeria (A r a m b o u r g , 1959). Re­ locality of Sinap 64, dated to early Vallesian ELM A, MN cently a new speciesOrycteropus abundulafus was described 9 and magnetostratigraphically estimated around 10.7 Ma from the Chadian sites Kossom Bougoudi and Toros Me- (K a p p e l m a n et al., 2003). It is quite possible, thus, that nalla dated to 7.0 Ma and close to Mio-Pliocene boundary the stratigraphic distribution of O. seni extends up to the (-5.3 Ma) respectively(L e h m a n n et ah, 2005; L e h m a n n , beginning of late Miocene (Fig. 6). Another Eurasian 2006). In Pliocene the sole known African orycteropodid taxon is 0. pottieri, a species restricted to the Vallesian so is Orycteropus djourabensis described from the fossiliferous far (Fig. 6). It is known from the localities of Sinap 12, sector of Kolle, Chad dated to early Pliocene (5.0-4.0 Ma), 72 and 108 dated at -10.59 Ma, -10.08 Ma and -10.13 (L e h m a n n et al., 2004). The species is also known from Ma respectively (K a p p e l m a n et al., 2003), suggesting Ethiopia and Kenya (L e h m a n n , 2006). an early Vallesian age (MN 9). O. pottieri is also known During Pleistocene two species of orycteropodids are from the locality of Pentalophos-1 (PNT, Axios Valley, known from Africa: Orycteropus afer and Orycteropus Macedonia, Greece). The fauna of PNT is peculiar, but a crassidens. The previous one is that living up to now; its biochronological age at the end of early Vallesian ELMA, first appearance is discussed. Some orycteropodid remains MN 9 or the beginning of late Vallesian ELMA, MN 10 from the early Pliocene locality of Langenbaanweg have is quite possible (K o u f o s , 2006). been attributed to O. afer (P ic k f o r d , 2005). However The distribution of the well known O. gaudryi spans over L e h m a n n (2006) does not agree to this determination the whole Turolian (Fig. 6). The taxon is certainly known of the Langenbaanweg material; thus its presence in the from several Greek and Turkish sites(W o o d w a r d , 1901; Pliocene is given by dashed line in Fig. 6. 0. crassidens C o lb e rt , 1941; B on is et al., 1994; S en, 1994 and included was originally described from Kenya and it is dated from bibliography; T h e o d o r o u et al., 2003). Some oryctero­ 1.8-0.01 Ma (M a c I n n e s , 1956). However, its systematic podid remains reported as O. cf. gaudryi are known from value is under discussion as it is considered either as syno­ the Late Turolian (MN 13) locality of Brishigella, Italy nym of O. afer (P ic k f o r d , 1975, 2005) or as a separate (R o o k S c M a s s i n i , 1994) and from the middle Turolian species (v a n d e r M a d e , 2003; L e h m a n n et al., 2005; level ofMaragheh, Iran (M e q u e n e m , 1925). Although the L e h m a n n , 2006). species is mentioned in the faunal lists of several locali­ The oldest known appearances of orycteropodids in Eura­ ties of Moldova and Ukraine (NOW, 2007), its presence sia are traced in the middle Miocene Turkish localities is dubious as all these old collections need a revision. Its of (pandir and Pa^alar with Orycteropus seni. The former older appearance is traced in the locality Kufuk (pekmece is dated to late Orleanian ELM A, MN 5 (NOW, 2007) (Turkey) dated probably to late Vallesian (MN 10). The but v a n d e r M a d e (2003,2005) suggested a younger age taxon makes its last appearance in the Greek locality of for it (Early Astaracian ELMA, MN 6). The locality of Dytiko, dated to late Turolian (MN 13), (B o n is et al., Pajalar is dated from late Orleanian to early Astaracian 1994). The species Orycteropus depereti is known from ELM A, MN 5-6 (NOW, 2007). Thus, the orycteropodids Perpignan (France), dated to early Pliocene (MN 15), appeared in Eurasia during late Orleanian ELM A, MN (H e l b in g , 1933). However, a more detailed comparison of 5 or at the end of Burdigalian. According to Rogl (1999) this species with O. gaudryi is necessary in order to specify the Tethyan Seaway (the marine branch connecting the its systematic position. The Perpignan aardvark is the last Mediterranean Sea to the Indo-Pacific Ocean) closed for appearance of the orycteropodids in Eurasia, which are the first time at the end of early Miocene (Burdigalian) and later restricted to Africa. more precisely around 19.0-18.0 Ma. The “Gomphotherium- Besides these ascertained appearances of the oryctero­ landbridge” allows the first dispersal of mammals from podids in Europe, there are several traces of the family Africa to Eurasia and vice-versa. During the Langhian in various European localities from Turkey, Georgia and ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 119

Hungary (NOW, 2007). However, the available material Miocene of Macedonia (Greece). — Neues Jahrbuch is too poorly preserved to allow accurate identification and für Geologie und Paläontologie Monatshefte, 194: is often referred to as Orycteropus indet., e.g. in the middle 343-360 Stuttgart. Miocene Georgian locality of Belometchetskaja an unde­ C o l b e r t , E.H., 1933. The presence of tubulidentates in termined orycteropodid is referred to as Orycteropus sp. the Middle Siwalik beds of northern India. — Ame­ (G a b u n i a , 1956); however v a n d e r M a d e (2003) refers rican Museum Novitates, 604:1-10, New York. that its identification to Orycteropus is quite dubious. C o l b e r t , E.H., 1941. A study of Orycteropus gaudryi from In the Siwaliks (Pakistan), the orycteropodids are known the island of Samos. — Bulletin American Museum by a single species Orycteropus browni found in the fos- of Natural History, 78:305-351, New York. siliferous levels, dated toVallesian, MN 9-10 (C o l b e r t , D r a k e , R.E., v a n C o u v e r in g , J.A., P ic k f o r d , M.H., 1933), (Fig. 6). This taxon is also referred from the early C u r t i s , G.H. & H a r r i s , J.A., 1988. New chrono­ Turolian locality of Kocherinovo-1, Bulgaria by a skull logy for the early Miocene mammalian faunas of Ki- known from illustrations (S p a s s o v et ah, 2006). singiri, Western Kenya. — Journal of the Geological Society of London, 145:479-491, London F o r s y t h M a jo r , C.J., 1888. Sur un gisement d’ossements 4. Acknowledgements fossiles dans l’île de Samos, contemporains l’âge de Pikermi. — Compte Rendus de LAcademie des Sci­ Tire excavations in Samos have been supported by the Pre­ ences, Paris, C X V IL 608-610, Paris. fecture of Samos and the “Konstantinos and Maria Zimalis” F o r s y t h M a j o r , C.J., 1894. Le gisement ossifère de Foundation. The Municipality of Mytilinii provided great help Mitylini et catalogue d’ossements fossiles recueillis à too. The Natural History Museum of Aegean offered us the Mitylini, île de Samos, et déposés au Collège Galliard, place for the preparation and storage of the fossils. The Syn- à Lausanne. — Georges Bridel &c Cie Editeurs, pp. thesys Program supported my visits to Vienna (AT-TAF-702), 1-51, Lausanne. London (GB-TAF-1842) and Paris (FR-TAF-3102).I thank F o r t e l i u s , M., 1990. Less common ungulate species Prof. P.Tassy (MNHN), Dr. S. Sen (MNHN), Dr. A. Currant from Pasalar, Middle Miocene of Anatolia (Turkey). (NHML) and Dr. G. Daxner-Höck (NHMW) for giving me — Journal of Human Evolution, 19:479-487, New access to the fossils at their disposal. Many thanks to: Dr. C. York. Sagne (MNHN), Dr. C. Argot (MNHN) and Dr. C. Soligo F o r t e l iu s , M., 2003. Fossil aardvarks of the Miocene Si- (NHML) for their help in searching the collections, and nap Formation, Turkey. — [in:] F o r t e l iu s , M., K a p - their great hospitality. I also thank Dr. T. Lehmann for his p e l m a n , J., S en , S. S c B e r n o r , R.L. (eds). Geology useful information and suggestions. Finally, I wish to thank and Palaeontology of the Miocene Sinap Formation, my collaborators and several students who excavated with me pp. 194-201, New York (Columbia Univ. Press). and helped me in the collection and preparation of the fossils. G a b u n i a , L., 1956. An extinct Tubulidentate (Ory­ cteropus sp.) from the Middle Miocene deposits of Belomechetskaya (North Caucase). — DoK. AN 5. References Azerbaidzhanskoi SSR, 12:203-206 (in Russian). H e l b in g , H., 1933. Ein Orycteropus-Fund aus dem Un­ A n d r e w s , C.W., 1896. On a skull of Orycteropusgaudryi teren Pliocäne des Roussillon. — Eclogae geologicae F o r s y t h M a jo r , from Samos. — Proceedings Zoo­ Helvetiae, 26(2):255-267, Basel. logical Society of London, 1896:296-299, London. K a p p e l m a n , J., D u n c a n , A., F e s h e a , M., L u n k k a , A r a m b o u r g , C., 1959. Vertébrés continentaux du M io­ J-P., E k a r t , D., M c d o w e l l , R, R y a n , T., S w i s h e r cene supérieur de LAfrique du Nord. — Publications III, C.C., 2003. Chronology of the Sinap Forma­ Carte géologique d Algérie, Paléontologie mémoires, tion. — [in:] F o r t e l iu s , M., K a p p e l m a n , J., S en, 4:1-161, Alger. S., B e r n o r , R.L. (eds). Geology and Palaeontology B o n is, L de., K o u f o s , G.D. & S e n , S., 1997a. 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Basin, Samos Island, Greece: New Collection. 2 . — Neues Jahrbuch für Geologie und Paläontologie Lithostratigraphy and fossiliferous sites. — Beiträge Monatshefte, 210:399-420, Stuttgart. zur Paläontologie, 31:13-26. Wien. B o n is, L. de, B o u v r a i n , G., G e r a a d s , D., K o u f o s , K o u f o s , G.D., 2006. The Neogene mammal localities G.D. &c S e n , S., 1994. The first aardvarks from the of Greece: faunas, chronology and biostratigraphy. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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— Hellenic Journal of Geosciences, 41(1):183-214, NOW, 2007. Neogene Old World, Database with the Ne­ Athens. ogene localities and their faunal lists, www.helsinki. K o u f o s , G.D., this volume. The late Miocene Mammal fi/science/ now/database.htm Faunas of the Mytilinii Basin, Samos Island, Greece: O zansoy, F, 1965. Etude des gisements continentaux et New Collection. 1. History of the Samos Fossil Mam­ des Mammifères du Cénozoique de Turquie. — M é­ mals. — Beiträge zur Paläontologie, 31:1-12. Wien. moires Société géologique France, 102:1-92, Paris. K o u f o s , G.D., B o n is , L. de & S e n , S., 1995. Lophocyon Patterson, B., 1975. The fossil aardvarks (Mamma­ paraskevaidisi a new viverrid (Carnivora, Mammalia lia, Tubulidentata). — Bulletin of the Museum of from the middle Miocene of Chios Island (Greece). Comparative Zoology Cambridge, 147(5):185-237, — Geobios, 28(4):5ll-523, Lyon. Cambridge. K o s t o p o u l o s , D., S e n , S. &. K o u f o s , G.D., 2 0 0 3 . 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New evidence concerning the fossil K o u f o s , G.D., K o s t o p o u l o s , D.S., V l a c h o u , T.D. aardvarks (Mammalia, Tubulidentata) of Pakistan. — 8 c S y l v e s t r o u , I.A., 2004. Reconsideration of the Tertiary Research, 2(l):39-44, London. Mytilinii fossiliferous basin, Samos, Greece. — 5th PiCKFORD, M., 2003. Minute species ofOrycteropus from International Symposium on Eastern Mediterrane­ the early Middle Miocene at Arrisdrift, Namibia. an Geology, Thessaloniki, April 2004, Abstracts, — Mémoires Geological Survey of Namibia, 19: 1:326-329, Thessaloniki. 195—198, Namibia. L ehmann, T., 2004. Les Orycteropodidae (Mammalia, PiCKFORD, M., 2005. Orycteropus (Tubulidentata, Mam­ Tubulidentata) du Mio-Plio-Quaternaire dAfrique. malia) from Langebaanweg and Baard’s Quarry, — PhD Thesis, University of Poitiers, France, pp. Early Pliocene of South Africa. — Comptes Rendus 1-474 (unpublished). Palevol., 4(8):7l5-726, Paris. L ehmann, T., 2006. Biodiversity of the Tubulidentata R ó g l , R, 1999. Circum-Mediterranean Miocene Paleo- over geological time. — Afrotherian Conservation, geography. — [in] Rôssner, G. ScH eissig, K. (eds). 4:6—10, Switzerland. The Miocene Land Mammals of Europe, vol. 1:39-48, L e h m a n n , T., 2007. Amended taxonomy of the order München (Verlag Dr Friedrich Pfeil). Tubulidentata (Mammalia, Eutheria). — Annals of Rook, L. 8 c M asini, R, 1994. Orycteropus cf. gaudryi the Transvaal Museum, 44:179-196. (Mammalia, Tubulidentata) from the Late Mes- L e h m a n n , T , V ig n a u d , R, L i k i u s , A. &. B r u n e t , M., sinian of the Montitcino Quarry (Faenza, Italy). 2005. A new species of Orycteropodidae (Mamma­ — Bollettino della Societa Paleontológica Italiana, lia, Tubulidentata) in the Mio-Pliocene of northern 33(3):369-374, Modena. Chad. — Zoological Journal of the Linnean Society, Sen, S., 1994. Les gisements de mammifères du Miocène 143:109-131, London. supérieur de Kemiklitepe, Turquie: 5. Rongeurs, Tu- L e h m a n n , T., V ig n a u d , R, M a c k a y e , H.T. &c B r u n e t , bulidentés et Chalicothères. — Bulletin du Muséum M., 2004. A fossil aardvark (Mammalia, Tubuliden­ national d’Histoire naturelle, Paris, 4eme sér., 16, tata) from the lower Pliocene of Chad. — Journal of section C, 1:97-111, Paris. African Earth Sciences, 49:201-217, Amsterdam. Solounias, N., 1981. The Turolian fauna from the island L e w i s , G.E., 1938. A skull of Orycteropus pilgrimi. — of Samos, Greece. — Contributions Vertebrate Evo­ American Journal of Science, 5th ser., 36,216:401—405, lution, 6:1—232. New York. Shoshani, J., G oldman, G.A. Sc Thewissen, J.G.M., L ö p e z -A n t o n a n z a s , R., S en, S. & K o u f o s , G.D., 2005. 1988. Orycteropus afer. — Mammalian species, A ctenodactylid rodent (Mammalia: Rodentia) from 300:1-8. the Middle Miocene of Chios Island (Greece). — Spassov, N., Tzankov, T. ôcG eraads, D., 2006. Late Geobios, 38:113-126, Lyon. Neogene stratigraphy, biochronology, faunal diversity M a c P h e e , R.D.E., 1994. Morphology, adaptations, and abd environments of South-West Bulgaria (Struma relationships of Plesiorycteropus, and a diagnosis of a Valley). — Geodiversitas, 28(3):477-498, Paris. new order of eutherian Mammals. — Bulletin of the Tekkaya, L, 1993. Türkiye fósil OrycteropodidaeTeri. American Museum of Natural History, 220:1-214. T. C. Kültür Bakanligi Anidar ve Müzeler Genel M enequem, R. de, 1925. Contributions à l’étude des fos­ Müdürlügü. — 8th Arkeometri Sonuçlari Toplantisi siles de Maragha (suite). — Annales de Paléontologie, 1992, Ankara, pp. 291-297. 14:135-160, Paris. Theodorou, G., A thanassiou, A., Iliopoulos, G. 8 c ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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R o u s s i a k i s , S. 2003. Remarks on the late Miocene v a n d e r M a d e , J., 2005. Errata and reply to Guest Editor’s vertebrates of Kerassia (Northern Euboea, Greece). notes. — Courier Forschung-Institut Senckenberg, — Deinsea, 10:519-530, Rotterdam. 254:133-147, Frankfurt. v a n d e r M a d e , J., 2003. The aardvark from the M io­ W o o d w a r d , A.S., 1901. On the bone-beds of Pikermi, cene hominoid locality Qandir, Turkey. — Courier Attica and on similar deposits in Northen Eubo­ Forschung-Institut Senckenberg, 240:133-147, ea. — Geological Magazine, new series, decade IV, Frankfurt. 3(ll):481-486, London. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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Orycteropusgaudryi, Mytilinii-lA (MTLA), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Partial skull, M TLA-115; a. right lateral, b. left lateral, c. dorsal, and d. occlusal view.

Fig. 2. Right mandibular fragment with p3 and ml-m3 associated with the skull, M TLA-115; a. buccal, b. lingual, and c. occlusal view.

Fig. 3. Partial skull, M TLA-240; a. right lateral, b. left lateral, c. dorsal, and d. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 123

PLATE 1 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

124 Beitr. Palaont., 31, Wien, 2009

PLATE2

Orycteropusgaudryi, Mytilinii-lA, B (MTLA, MTLB), Samos, Greece, middle Turolian (MN 12).

Fig. 1. Left mandibular fragment with m3, M TLA-116; a. buccal, b. lingual, and c. occlusal view.

Fig. 2. Right mandibular fragment with p4-m3, M TLA-280; a. buccal, b. lingual, and c. occlusal view.

Fig. 3. Left mandibular fragment with m2-m3, M TLA-306; a. buccal, b. lingual, and c. occlusal view.

Fig. 4. Right mandibular fragment with p2-m3, M TLB-49; a. buccal, b. lingual, and c. occlusal view.

Fig. 5. Left M l, M TLA-239; a. buccal, b. lingual, and c. occlusal view.

Fig. 6. Left m l, M TLA-18; a. buccal, b. lingual, and c. occlusal view.

Fig. 7. Left m2, M TLA-17; a. buccal, b. lingual, and c. occlusal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Tubulidentata. 125

PLATE2