(Mammalia, Artiodactyla) from the late of Akkas ¸ dag ˘ ı, Turkey

Dimitris S. KOSTOPOULOS University of Thessaloniki, Museum of Paleontology, Geological Department, 54124 Thessaloniki (Greece) [email protected]

Gercek SARAÇ General Directorate of Mineral Research and Exploration, Natural History Museum, TR-06520 Balgat, Ankara (Turkey)

Kostopoulos D. S. & Saraç G. 2005. — Giraffidae (Mammalia, Artiodactyla) from the late Miocene of Akkas ¸ dag ˘ ı, Turkey, in Sen S. (ed.), Geology, and environments atAkkas ¸ dag ˘ ı, late Miocene of Central Anatolia. Geodiversitas 27 (4) : 735-745. KEY WORDS Mammalia, Giraffidae, ABSTRACT , A few dental and several postcranial giraffid remains from the late Miocene Samotherium , , locality of Akkas¸ dag˘ ı (Central Anatolia, Turkey) have been identified as late Miocene, belonging to Helladotherium sp., Palaeotragus rouenii Gaudry, 1861 and Turolian, Samotherium cf. major Bohlin, 1926. The comparison of the material with Akkas¸ dag˘ ı, Central Anatolia, several Eurasian representatives of the three genera indicates a middle-late Turkey. Turolian age.

RÉSUMÉ MOTS CLÉS Giraffidae (Mammalia, Artiodactyla) du Miocène supérieur d’Akkas¸ dag˘ ı, Mammalia, Giraffidae, Turquie. Palaeotragus, Quelques restes dentaires et postcraniens de giraffidés, provenant du Miocène Samotherium , Helladotherium , supérieur de la localité d’Akkas¸ dag˘ ı (Anatolie centrale, Turquie) sont déter- Miocène supérieur, minés comme appartenant à Helladotherium sp., Palaeotragus rouenii Gaudry, Turolien, 1861 et Samotherium cf. major Bohlin, 1926. La comparaison du matériel Akkas¸ dag˘ ı, Anatolie Centrale, turc avec les représentants eurasiatiques de ces trois genres permet de le dater Turquie. du Turolien moyen-supérieur.

GEODIVERSITAS • 2005 • 27 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 735 Kostopoulos D.S. & Saraç G.

INTRODUCTION RZO Ravin des Zouaves-5, Axios valley, Greece; In contrast to the contemporaneous asso- VAT Vathylakkos 3, Axios valley, ciations of Eastern Europe, giraffids seem to be rare Greece. at the late Miocene locality of Akkas¸ dag˘ ı (Central Measurements Anatolia, Turkey; Kazancı et al. 1999). The avail- DAP anteroposterior diameter; able material of 24 identifiable specimens comes DT transverse diameter; from 11 bone-pockets of a single stratigraphic hori- Hheight; Llength; zon. The material labelled as AK comes from bone Wwidth; pockets excavated between 1997 and 2001 or from alvalveolar; surface collection (AKK); this material is stored at art articular; the Natural History Museum in Ankara. A few diaph diaphysis; specimens labelled GOK derive from the excavation dist distal; latlateral; led by Émile Heintz in 1971 and are preserved at maxmaximal; the Muséum national d’Histoire naturelle in Paris. med medial; Although insufficiently documented, three forms occlocclusal; have been recognized: a large Sivatheriinae Zittel, proxproximal. 1893 ascribed to Helladotherium Gaudry, 1860, a small sized Palaeotraginae Pilgrim, 1911 referred to Palaeotragus rouenii Gaudry, 1861 and a larger SYSTEMATICS form of the latter subfamily, similar to Samothe- rium major Bohlin, 1926 from Samos. Family G IRAFFIDAE Gray, 1821 Subfamily S IVATHERIINAE Zittel, 1893

A BBREVIATIONS Genus Helladotherium Gaudry, 1860 Museums and localities AeMNH Aegean Museum of Natural His- T YPE SPECIES. — Helladotherium duvernoyi (Gaudry & tory-Zimalis Foundation, Samos Lartet, 1856). Type locality: Pikermi, Greece. island, Greece; AK (A, B, K, 1-14) Akkas¸ dag˘ ınew collection; BMNH Natural History Museum, Lon- Helladotherium sp. don; DIT Dytiko 3, Axios valley, Greece; GOK Akkas¸ dag˘ ı Heintz’ collection; M ATERIAL EXAMINED AND MEASUREMENTS (in mm). — KTA,B,D Kemiklitepe A,B,D, Turkey; P3 left (AK7-29): L occl = 34.0, W occl = 26.0, W alv = LGPUT Museum of the Geological 38.5; M2 right (AK2-441): L occl = 46.8, W max-anterior Department, Aristotle University lobe = 44.0, W max-posterior lobe = 41.0; radius left (AK7- of Thessaloniki; 64): L max = 580.0, DTprox = 123.0, DAPprox = 73.0, MGL Musée cantonal de Géologie, DTdiaph = 78.8, DAPdiaph = 60.0, DTdist-art = 101.3, Lausanne; DAPdist-art = 62.2; tibia right (AK7-129): L = 540.0, MNHN Muséum national d’Histoire DTprox-max = 166.5, DAPprox = 130+, DTdiaph = 74.5, naturelle, Paris; DAP = 53, DT = 106.2, DAP = 82.4); part MTA-MA Maden Tetkik ve Arama Mu- diaph dist dist of calcaneus (GOK-200): L sustentaculum tali = 167.0; talus seum (General Directorate of (GOK-197): L = 115.5, L = 99.3, DT = 76; Mineral Research and Explora- lat med dist tion, Natural History Museum), cubonavicular (AK7-101): DTmax = 100.0, DAPmax = Ankara; 99.0; phalanx I (AK7-152): L = 114.0, DTprox = 53.2, MTLA,B Mytilinii-1 A and B, Samos, DAPprox = 58.2, DTdist = 47.2, DAPdist = 35.3; (AK3- Greece; 310): L = 109.2, DTdist = 42.5, DAPdist = 33.7; NKT & NIK Nikiti-1 & 2, Greece; (GOK-201): L = 115.0, DTprox = 52.5, DAPprox = PIK Pikermi, Greece; 56.3, DAPdist = 41.2; phalanx II (AK7-35a): L = 62.5, RPlRavin de la Pluie, Axios valley, DTprox = 45.3, DAPprox = 46.2, DTdist = 40.8, DAPdist Greece; = 44.5; (AK7-27): L = 65.4, DTprox = 46.4, DAPprox =

736 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Giraffidae from Akkas¸ dag˘ ı

AB

F IG. 1. — Helladotherium sp., Akkas ¸ dag ˘ ı, Turkey, occlusal view; A , M2 right; B , P3 left. Scale bar: 1.5 cm.

46.2, DTdist = 43.5, DAPdist = 44.5; phalanx III (AK7- genera Samotherium Forsyth-Major, 1888 and 35b): DTart = 35.0, H art = 58.6. Helladotherium Gaudry, 1860 which however, Provisionally ascribed: metacarpal III+IV of immature belong to different phylogenetic lineages, the individual (AK7-65): DTprox = 87.5, DAPprox = 55.0, DT = 50.0, DAP = 42.5. Palaeotraginae and Sivatheriinae respectively diaph diaph (Bohlin 1926; Hamilton 1978; Geraads 1986). Geraads & Güleç (1999) mention that the type D ESCRIPTION AND DISCUSSION (F IGS 1-4) specimen of Helladotherium duvernoyi Gaudry, The cranial elements are limited to two isolated 1860, type species of the genus, belongs to a teeth with finely rippled enamel and barely visi- female individual of a different genus and, recall- ble cingulum (Fig. 1). The P3 is large with strong ing Matthew’s statement (Matthew 1929: 550 parastyle and well developed paracone rib. The fide Hamilton 1978: 218) they suggest a provi- internal side of the labial crescent is weakly divid- sional synonymy of Helladotherium with Brama- ed into paracone and metacone. The occlusal sur- therium Falconer, 1845. Although quite possible, face looks sub-quadrangular with an this synonymy is not yet formally founded and, antero-lingual protuberance of the lingual wall following Hamilton (1978), we shall continue to and a clear hypoconal spur on the central cavity use Helladotherium as a valid taxon. (Fig. 1). The M2 has simple morphology with According to Bohlin (1926) and Geraads (1974), strong parastyle, slim but well built mesostyle, Helladotherium differs from Samotherium in the weak metastyle and strong paracone rib. The lin- larger premolar row relatively to the molars, the gual wall of the protocone is rounded, while the unmolarized p3, the less developed styles on the hypocone is slightly narrower and more angular cheek teeth and the more massive limbs. lingually. The anterior flange of the protocone is At first sight the large P3 from Akkas¸ dag˘ ı, signifi- connected with the parastyle; its posterior flange cantly larger than usually recorded in Samo- is short, curves anteriorly and do not confine the therium and notably large comparatively to M2, posterior flange of the hypocone. A relatively indicate the presence of Helladotherium (Fig. 2). strong hypoconal spur is present (Fig. 1). In contrast to the studied specimen and The absence of cranial or more complete dental Helladotherium , the P3 of Samotherium is more material makes the identification of the largest rounded and presents stronger metastyle and giraffid from Akkas¸ dag˘ ı quite difficult. Similar more centrally placed paracone-metacone pillar. sized Turolian forms are usually referred to the The M2 structure (thin mesostyle, posterior

GEODIVERSITAS • 2005 • 27 (4) 737 Kostopoulos D.S. & Saraç G.

60 AB 55

5 0

4 5 WM2

4 0

3 5

3 0 2 0 2 5 3 0 3 5 4 0 4 5 W P 3

M GL Akkasdagı¸ ˘ China M a r agh a P I K V A T KTB B M NH T A S K ER

F IG.2. — Scatter diagram “Width of P3 against Width of M2” of Helladotherium sp., Akkas ¸ dag ˘ ı, Turkey ( ■ ) in comparison with Helladotherium duvernoyi from Kerassia (KER), Pikermi (PIK, several collections); Samotheriummajor from Vathylakkos (VAT), Kemiklitepe B (KTB), Tas ¸ kınpas ¸ a (TAS); Samotherium boissieri from Samos (MGL and BMNH collections); S. sinense (China) and S. neumayri (Maragha) (data from Bohlin 1926; S ¸ enyürek 1954; Geraads 1974, 1994; Iliopoulos 2003; and pers. data). F IG. 3. — Helladotherium sp., Akkas ¸ dag ˘ ı, Turkey, anterior view; A , left radius AK7-64; B , right tibia AK7-129. Scale bar: 5 cm. flange of the protocone, presence of hypoconal dal face. This character strongly recalls Hella- spur, etc.) also differentiates the studied specimen dotherium from Pikermi but also Samotherium from Samotherium ,supporting close relationships sinense (Bohlin 1926: fig. 103), while the imma- with Gaudry’s genus. However, the available M2 turity of the individual may influence the fossa appears relatively narrower than that of the type pattern (Geraads pers. comm. 2004). Regarding species Helladotherium duvernoyi from Pikermi the absolute dimensions, this young individual (MNHN, BMNH, LGPUT) and closer to the falls, however, within the range of large samoth- Maragha (Iran; MNHN, BMNH) and Kerassia eres, indicating probably an even larger and (Greece; Iliopoulos 2003) samples. stouter adult , hence, closer to those of Although more robust, the dimensions of the Helladotherium . postcranials of Helladotherium are usually hardly The rectangular epiphyses of the complete radius distinguished from those of the large samotheres AK7-64 are not significantly wider than the shaft (e.g., S. major Bohlin, 1926 and S. sinense (Fig. 3A), the lateral tuberosity is weak, the radial Bohlin, 1926). The limb proportions of the stud- tuberosity is placed below the medial proximal ied specimens from Akkas¸ dag˘ ı are placed between articular surface, the antero-lateral corner of the those of Samotherium and Helladotherium , being proximal part forms an almost right angle, the closer to the second genus. radial styloid process is more projected downwards Differently from Samotheriumboissieri Forsyth- than the ulnar one, the shallow groove of the exten- Major, 1888 and S.major , the proximal articula- sor capri rarialis muscle is defined by two blunt tion of the preserved metacarpal, provisionally crests of more or less equal length and it is sym- ascribed to Helladotherium , presents a large syn- metrically located above the medial ulnar ridge, the ovial fossa, which opens widely towards the cau- groove of the abductor digiti I longus muscle is shal-

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low and located rather anteriorly than medially, the anterior margins of the distal articular facets are low, the anterior part of the scaphoid facet is rather quadrangular with flat anterior border bended medially, the anterior part of the lunar facet is spindle-shaped with clear posterior margin, the lateral crest of the lunar is shorter, less prominent and more oblique than the medial one, the artic- ular surface for the cuneiform is rather narrow and the transverse crest of the posterior face is weakly developed (Fig. 4). The tibia (AK7-129, Fig. 3B) is relatively long and moderately robust (DTdiaph × 100/L =13.5). The tibial crest is relatively short, lateral- F IG. 4. — Helladotherium sp., Akkas ¸ dag ˘ ı, Turkey, distal articula- ly located, not very prominent and with wide- tion of radius AK7-64. Scale bar: 2 cm. shallow tuberosity. Consequently, in the upper part of the anterior face the restricted tibial sulcus is oval-shaped and not very deep. The tibial spine is relatively high. A small facet for the fibula is In the absence of adequate data we refer at the present at the lateral side. The muscle imprint at moment this form to Helladotherium sp. the posterior side of the bone is located in the medial part of the diaphysis. The antero-lateral tuberosity of the distal part is weakly developed. Both the medial malleolus and the lateral malleo- Subfamily P ALAEOTRAGINAE Pilgrim, 1911 lar facets of the distal epiphysis are strong. Genus Gaudry, 1861 The dorsal and plantar edges of the calcaneus are Palaeotragus rectangular. The large talus has strongly assyme- T YPE SPECIES. — Palaeotragus rouenii Gaudry, 1861. trical proximal trochlea, the scar for the external Type locality: Pikermi, Greece. tendon of the cubonavicular is missing and the limit between the articular facets for the calca- Palaeotragus rouenii Gaudry, 1861 neum and the cubonavicular is well marked. The cubonavicular is almost square with extremely M ATERIAL EXAMINED AND MEASUREMENTS (in mm). — developed caudal tuberosity, coating the proxi- Upper toothrow (AK3-298): LP2-M3 = 123.0, LP2- mal articular surface. The posterior metatarsal P4 = 53.0, LM1-M3 = 72.7, LP2 = 15.5, WP2 = 15.3, LP3 = 15.8, WP3 = 18, LP4 = 17.1, WP4 = 17.1, facet is present. The first phalanx is large and LM1 = 23.1, WM1 = 22.5, LM2 = 24.8, WM2 = 26, robust. LM3 = 26.5, WM3 = 25.8; lower toothrow (AK12- This set of postcranial morphological characters 78): Lp2-m3 ≈ 129.0, Lp2-p4 ≈ 49.0, Lm1-m3 = rules out the association with Samotherium and 77.0, Lp3 = 17.0, Wp3 = 10.1, Lp4 = 18.5, Wp4 = place the Akkas¸ dag˘ ı formcloser to Helladotherium 12.2, Lm1 = 22.5, Wm1 = 15.4, Lm2 = 24.5, Wm2 = (Bohlin 1926; Geraads 1974; Iliopoulos 2003; 16.4, Lm3 = 30.5, Wm3 = 15.3; proximal part of radius (AK5-392): DT = 88.2, DAP = 50.0; Iliopoulos pers. comm. 2004; pers. data). None- prox prox part of metacarpal III+IV (AK6-86): DTprox = 65.0, theless, the observed morphological features are DAP = 48.0, DT = 34.4, DAP = 41.0; not fully identical to those of the Pikermian form prox diaph diaph (AK6-87): DTprox = 56.6, DAPprox = 38.4; tibia (AK4- (BMNH sample), suggesting either a larger intra- 236): DTdist = 67.5, DAPdist = 51.7; talus (AK3-197): specific variability for Gaudry’s species or – less L lat = 76.2, L med = 69.0, DTdist = 50.0; (AK5-184): possibly – a distinction at a higher taxonomic level. L lat = 80.0, L med = 72.0, DTdist = 49.5.

GEODIVERSITAS • 2005 • 27 (4) 739 Kostopoulos D.S. & Saraç G.

A

B

F IG. 5. — Palaeotragus rouenii from Akkas ¸ dag ˘ ı, Turkey, occlusal view; A , upper dentition; B , lower dentition. Scale bar: 2 cm.

D ESCRIPTION (F IGS 5; 6) protocone of M1, 2 is angular and slightly con- The teeth are small, brachyodont and with finely stricted lingually (Fig. 5A). A strong hypoconal rippled enamel (Fig. 5). The upper premolar row spur is present in all upper molars and especially represents 72.9% of the molars, while the same in M3, which also bears a weak protoconal fold. ratio is estimated about 63-65 for the lower den- The cingulum is weakly developed both on the tition. lingual and labial faces. A very short basal pillar is also present. Upper toothrow (Fig. 5A) P2 is simple with a rudimentary hypoconal spur, Lower toothrow (Fig. 5B) directed strongly backwards. The parastyle is well p3 is highly molarized with strongly elongated developed. The paracone rib is strong and is metaconid, parallel to the anteroposterior axis of placed anteriorly. A weak cingulum is present the tooth and extremely shortened talonid. The anterolingually (Fig. 5A). P3 is morphologically elongated endoconid is independent from the similar to P2, but with stronger hypoconal spur, metaconid in early wear. The reduced endostylid tending to form a hypoconal islet. The parastyle is obliquely settled. Labially, a well developed and the lingual cingulum are also stronger than furrow separates the bulgy hypoconid from the in P2. P4 is more symmetrical than P2 and P3. strong protoconid. The parastylid is well defined In occlusal view there is a well formed hypoconal (Fig. 5B). p4 is also molariform (Fig. 5B). Its islet (Fig. 5A). The paracone rib is strong and sit- metaconid is long and the parastylid thinner than uated centrally on the labial wall, while both the in p3. The endoconid is well distinct and parastyle and the metastyle are less developed. A oblique. The endostylid is longer than in p3 and weak cingulum appears along the lingual surface. placed lingually. On the labial wall, the trigonid All the molars have moderate to well developed is distinguished from the bulgy talonid by a deep styles and ribs. The posterior flange of the para- furrow. The first molar shows a weak anterior cone does not confine with the mesostyle. The fold. A rudimentary basal pillar, a well developed

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metaconid and a strong metastylid (especially on 110 the m2, 3) are present in all molars. The third lobe of m3 is relatively small, elliptical and bi- 100 cuspid (Fig. 5B). 9 0 Postcranials -M3 8 0 1

The poor postcranial material ascribed to this M form does not allow major observations. The pre- 7 0 served part of the metacarpal clearly shows dolichopodial morphology. The external articular 6 0 groove of the distal trochlea of the tibia is shorter 5 0 than the internal one. The plantar face of the 4 0 5 0 6 0 7 0 8 0 9 0 P 2 - P 4 talus lacks the depression for the external tendon 1 3 0 of the cubonavicular and its external face is quite flat. 1 2 0 110 3 m

D ISCUSSION - 100 1

Palaeotragus is a well known genus from the m 9 0 so-called Greco-Iranian province and Eurasia in general, while its distribution area extends also to 8 0 Africa. Nonetheless, the phylogenetic relation- 7 0 ships among the referred late Miocene species are 6 0 4 0 4 550 55 6 0 6 5 7 0 7 5 8 0 not always clear and the species synonymy p 2 - p 4 appears sometimes to be confused (see Bohlin P. microdon P. rouenii DIT 1926; Bosscha-Erdbrink 1977; Hamilton 1978; P. cf. coe l o p h r y s P. rouenii Geraads 1986, 1994; Gentry et al. 1999). Over- Akkasdagı¸ ˘ P. b e rislavicu s looking the palpable nomenclature problems, it is P. de cip i e n s evident that most of the Turolian Palaeotragus could be grouped in two size categories: F IG. 6. — Scatter diagram “Length P2-P4 against Length M1-M3 (p2-p4/m1-m3)” of Palaeotragus rouenii, Akkas ¸ dag ˘ ı, Turkey ( ■ ) – a group of small-sized and slender-limbed in comparison with other late Miocene Palaeotragus species (data from Bohlin 1926; Geraads 1974, 1978; and pers. data). forms, represented mainly by the type species of Abbreviation: DIT, Dytiko 3, Axios valley, Greece. the genus P. rouenii Gaudry, 1861 from Pikermi (Greece) and its allies; and – a group of larger and stouter forms, principally represented by P. coelophrys (Rodler & Weithofer, between 57-62 [n = 7] in the first species and 1890), originally described from Maragha (Iran). between 63-73 [n = 5] in the second one; Fig. 6) The contemporaneous Palaeotragus microdon and its limb proportions are slightly different (Koken, 1885) from China (mainly from Loc. (more slender limb bones, stouter tali, etc.). 116 of Kansu; Bohlin 1926) is considered to be Therefore, we regard P. microdon as a distinct very similar to P. rouenii (Bohlin 1926; Bosscha- species. Erdbrink 1977; Geraads 1986). Nevertheless, Except for the ossicone and skull morphology P.microdon presents ossicones in both sexes while (which anyway are not available in the Akkas¸ dag˘ ı P. rouenii females appear to be “hornless” (Bohlin collection) P. coelophrys differs from P. rouenii in 1926; Geraads 1974). Moreover, the lower denti- its larger size and dental dimensions (Fig. 6), sim- tion of P. microdon presents a comparatively pler dental morphology, more robust and less shorter premolar row than that of P. rouenii (the dolichopodial limbs (Bohlin 1926; Geraads index [premolar/molar row length]% ranges 1974, 1978). Moreover, the p3 of P. coelophrys

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“smoothening” cannot be systematically or chronologically appreciated because of the insuf- ficient data, the Akkas¸ dag˘ ı P. rouenii seems to be closer to the later Turolian forms than to the early ones.

Genus Samotherium Forsyth-Major, 1888

T YPE SPECIES. — Samotherium boissieri Forsyth-Major, 1888. Type locality: Samos, Greece.

Samotherium cf. major Bohlin, 1926

M ATERIAL EXAMINED AND MEASUREMENTS (in mm). — Distal part of tibia (AK2-506): DTdist = 106.6, DAPdist = 76.0; part of tibia (AK4-203): DTdiaph = 70.0, DAPdiaph = 52.5; talus (AK7-28): L lat = 109.5, DTdist = 74.0; (GOK-198): L lat = 105.2, L med = 92.0, DTdist = 68.6; cubonavicular (AK11-65a): DTmax = 88.7, DAPmax = 77.0.

F IG. 7. — Samotherium cf. major , Akkas ¸ dag ˘ ı, Turkey, right talus AK7-28, posterior view. Scale bar: 4 cm. D ESCRIPTION AND DISCUSSION (F IG. 7) The presence of a second large giraffid in Akkas¸ dag˘ ı is poorly but certainly documented by has an independent metaconid, while the p3 of a few postcranial elements. Although the absolute P.rouenii presents a strong molarization on the dimensions of the available specimens are slightly lingual wall. smaller than those of Helladotherium , their pro- The small size, the molarized p3 and p4, the portions and some morphological characters bi-cuspid talonid of m3, the weakly developed clearly separate them from this genus: the lateral cingula and basal pillars on the molars and the malleolus surface of the distal tibia is reduced slender limbs of the Akkas¸ dag˘ ı form rule out the (large in Helladotherium ); the proximal trochlea association with P. coelophrys and related forms, of the talus (Fig. 8) is moderately unequal (clearly and match P. rouenii and P. microdon. Moreover, asymmetrical in Helladotherium ); the proximo- the “premolar/molar ratio” values for the lateral tuberosity of the calcanear facet is weak Akkas¸ dag˘ ı specimens are larger than those of (usually strong in Helladotherium ); the medial P.microdon and within the known range of ridge of the plantar trochlea is continuous (pres- P.rouenii. ence of notch in Helladotherium ) and presents a The original comparison with P. rouenii from large, shallow and round imprint at its lateral RPl, NKT, DIT (Greece; LGPUT), PIK base (absent in Helladotherium ) (Fig. 7); the (Greece; MNHN, BMNH), Samos (Greece; cubonavicular is longer transversally than antero- MGL, BMNH, AeMNH) and KTD (Turkey; posteriorly (squarish in Helladotherium ). This set MNHN), does not exhibit important morpho- of morphological features is indicative of Samo- logical or metrical differences (Fig. 6). Neverthe- therium (Bohlin 1926; Geraads 1974; pers. data). less, the accessory features of the dentition (labial The type species Samotherium boissieri Forsyth- and lingual cingula, basal pillars, spurs, etc.) seem Major,1888 (Geraads 1994), originally known to become less significant in the younger samples from the late Miocene deposits of Samos island of the species. Although the value of this (Greece), appears to present a great size variabili-

742 GEODIVERSITAS • 2005 • 27 (4) Late Miocene Giraffidae from Akkas¸ dag˘ ı

ty. Based on the works of Bohlin (1926) and 85 S ¸ enyürek (1954), Geraads (1994) refined the spe- 8 0 cific status of Samotherium from Samos, recog- 7 5 nizing two species, S. boissieri Forsyth-Major, 7 0 1888 and S. major Bohlin, 1926, the latter one dist considered as a successor of the former. We also DT 6 5 consider the classical Samotherium stock from 6 0 Samos as certainly bi-specific. Study of the 55 Forsyth-Major collections (MGL, BMNH), as 5 0 well as of the new material collected during the 7 5 9 5 115 1 3 5 L lat last years (Koufos et al. 1997; and pers. data) S. major MGL S. b o i ssier i B M NH, MGL allow us to recognize two forms of certainly dif- Samother ium K TD G OK-1 9 7 ferent stratigraphic origin, similar but not identi- H e lla d o ther ium S. major K T A,B cal in cranial morphology and different in size. A K 7 - 2 8 S. major T A S According to the new available magnetostrati- G OK-1 9 8 graphic data (Kostopoulos et al. 2003), the fossil- F IG. 8. — Scatter diagram “L lat against DTdist” of talus of Hel- iferous levels yielding S. boissieri (“Stefano”, Qx, ladotherium sp., Akkas ¸ dag ˘ ı, Turkey (GOK-197) and Samothe- Q4) are certainly older than those with S. major rium cf. major , Akkas ¸ dag ˘ ı, Turkey (AK7-28, GOK-198) in comparison with Helladotherium from several sites, Samothe- (“Andriano”, Q1). rium major from Samos (MGL collection), Kemiklitepe A,B In comparison to the known Samotherium species, (KTA,B), Tas ¸ kınpas ¸ a (TAS), Samotherium sp. from Kemiklitepe D (KTD) and Samotherium boissieri from Samos (BMNH and MGL the Akkas¸ dag˘ ıform appears dimensionally closer collections) (data from S ¸ enyürek 1954; and pers. data). to the large samotheres referred to S.major (Fig. 8) from the upper horizons of Samos, VAT, and KTA,B (Geraads 1978, 1994) and it could be age such as Çobanpınar, Balçıklidere (= Kemik- referred to as Samotherium cf. major . litepe), Eski Bayırköy, Gökdere and Kavakdere exhibit the presence of Helladotherium/Bramath- erium (Sickenberg 1975; Geraads & Güleç 1999; BIOCHRONOLOGY AND CONCLUSIONS NOW database 2003) but as the systematic value of the genus is still under discussion and the Regarded as a common element of the late Akkas¸ dag˘ ımaterial few for certain conclusions, Miocene faunas from the Greco-Iranian the presence of the genus cannot provide a more province, giraffids often constitute a quite accurate means of dating. monotonous assemblage of relatively low Palaeotragus rouenii is also widely distributed in biochronological value. Although the family is time and space. The species or a closely related poorly documented and rather sporadically pres- form ( P. pavlowae Godina, 1979; P. moldavicus ent in Turkey, all described forms had already Godina, 1979) probably appeared during late been mentioned from several localities of late Vallesian (E¸s me-Akçaköy, RPl, NKT, Poksheshty) Miocene age (S ¸ enyürek 1952; Ozansoy 1965; and became firmly established in the early Turo- Sickenberg 1975; Geraads 1994). lian faunas of the peri-Mediterranean region The first appearance of Helladotherium is proba- (Grebeniki, Nova Elizavetovka, RZO, KTD). Its bly dated at the end of Vallesian-beginning of maximum geographic extension took place during Turolian (NKT, NIK, RZO, Prochoma) and MN12 and it still exists in MN13 (DIT) (Geraads later on (MN12) the genus becomes more abun- 1994; Kostopoulos et al. 1996; Gentry et al. 1999; dant, occupying a wide territory from Western NOW database 2003). The dental characters of Europe to India (Bohlin 1926; Bosscha-Erdbrink the Akkas¸ dag˘ ı P. rouenii indicate closer affinities to 1977; Gentry et al . 1999; NOW database 2003). the middle-late Turolian forms of the species, sug- Several Turkish localities of middle-late Turolian gesting a similar age for the locality.

GEODIVERSITAS • 2005 • 27 (4) 743 Kostopoulos D.S. & Saraç G.

The common late Miocene large giraffid Samo- age (MN12) which is in agreement with the therium presents a vast geographic distribution radiometric dating data provided by Karadenizli from China to the Eastern Europe. In eastern et al . (2005). Mediterranean small-medium-sized forms appeared as early as in early Turolian (MN11; KTD, Samos, Maragha) and replaced later by Acknowledgements larger forms, namely S. major (MN12; VAT, Geological studies and excavations at Akkas¸ dag˘ ı Samos, KTA,B, Tas¸ kınpas¸ a, etc.). The presence have been authorized by the MTA and Ankara of Samotherium in the latest Turolian-earliest University,Faculty ofSciences.The work was done Ruscinian (MN13/14) locality of Maramena thanks to grants from the CNRS (ECLIPSE (Greece; Koehler et al. 1995) is questioned and it Program and DRI), the MNHN, MTA and seems that the genus falls into decline during late TUBITAK. They have financially supported field- Turolian. The large samothere from Akkas¸ dag˘ ı work and reciprocal visits to the concerned insti- shows clear metrical similarities with Samothe- tutions. The authors are grateful to G. Bouvrain, rium major from Samos (upper fossiliferous lev- L. de Bonis, L. Ginsburg, E. Heintz and all par- els), VAT (Greece), KTA,B and Tas¸ kınpas¸ a ticipants of 2000-2001 excavations. The authors (Turkey), indicating an MN12 age. thank Sevket Sen for having invited them to dig Although the giraffid association of Palaeotragus, and study the Akkas¸ dag˘ ı material. Thanks are also Samotherium and Helladotherium is not unexpect- due to Aymont Baud (UNIL, Lausanne) and the ed, it is not so common in the faunal record. In MGL, M. Bertling and the Geologische und Maragha (Iran), Helladotherium is associated with Paläontologische Institut, Münster, J. Hooker, Palaeotragus coelophrys, while Samotherium is rep- A.Currant, A. W. Gentry and the BMNH for per- resented by a medium-sized form. The old collec- mission to access their collections and overall assis- tion from Samos also includes both Samotherium tance and G. Iliopoulos, Leicester, for discussion. and Helladotherium in association with P. rouenii We sincerely thank D. Geraads (MNHN) and and the larger, badly known P.quadricornis J.Morales (MuseoNacionaldeCiencias Naturales, Bohlin, 1926 but the material comes from at least Madrid) for their useful comments and suggestions four stratigraphic horizons, while Samotherium is on the manuscript and for providing personal data represented by two distinct forms and conse- and key papers. This work is partly supported by quently, the combination at specific level cannot the IHP program of the BMNH. The photographs be fully controlled (Bosscha-Erdbrink 1977; are due to Denis Serrette (MNHN). Solounias 1981; Geraads 1994; Kostopoulos et al. 2003; NOW database 2003). P.rouenii and Hel- ladotherium duvernoyi are also present in the clas- REFERENCES sic fauna of Pikermi, but the occurrence of a B OHLIN B. 1926. — Die Familie Giraffidae. Palaeon- samothere in this locality has been not yet proved. tologica Sinica ser. C, 4 (1): 1-178. The new collection from Samos shows that in the B OSSCHA-E RDBRINK D. P. 1977. — On the distribu- tion in time and space of three giraffid genera with MN12 fauna coming from the upper fossiliferous Turolian representatives at Maragheh in N.W. Iran. horizons (MTLA,B; Koufos et al . 1997) P. rouenii Proceedings of the Koninklijke Nederlandse Akademie coexists with Helladotherium and Samotherium van Wetenschappen B 80 (5): 337-355. major . A very similar condition exists in the G ENTRY A. W., R ÖSSNER G. E. & H EIZMANN P. J. 1999. — Suborder Ruminantia, in R ÖSSNER G. & MN12 sites of Kerassia (Greece, Iliopoulos 2003), H EISSIG K. (eds), The Miocene Land Mammals of Kemiklitepe A,B (= Balçıklıdere, Turkey, Sen Europe. Verlag Dr F. Pfeil, München: 225-258. pers. comm. 2004; Geraads 1994) and probably G ERAADS D. 1974. — Les giraffidés du Miocène supérieur de la région de Thessalonique (Grèce). Thèse in Taraklia (NOW database 2003). de 3e cycle, Université Paris VI, France, 102 p. In conclusion, the giraffid assemblage of G ERAADS D. 1978. — Les Palaeotraginae (Giraffidae, Akkas¸ dag˘ ırather indicates a late middle Turolian Mammalia) du Miocène supérieur de la région de

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Submitted on 28 July 2003; accepted on 20 January 2005.

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