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

Beitr. Paläont., 31:397-408, Wien 2009

The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 16. Biochronology

by

George D. Koufos, Dimitris S. Kostopoulos & Theodora D. Vlachou*)

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 , Th.D. 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 16. Biochronology. — Beitr. Palaont., 31:397-408, Wien.

Abstract Zuordnung der Säugetierfunde von Samos für lange Zeit. Zwei Interpretationen sind bekannt: bei der ersten handelt The limited correlation of the old collections to the fos- es sich um die Möglichkeit zweier Faunenvergesellschaf­ siliferous sites and to the local stratigraphy prevented safe tungen und die andere wäre eine homogene, isochrone dating of the Samos mammal fauna for a long time. Two Fauna. Neu aufgesammeltes Material und die detaillierte main approaches are known: that supporting the presence Studie dazu erlauben eine Trennung in drei aufeinander of two faunal assemblages and that of a single homogene­ folgende Faunenhorizonte (MLN, MYT, MTL), die vom ous and isochronous fauna. The collected new material, obersten Frühturolium bis zum späten Mittelturolium and its detailed study and comparison allow the separation reichen. Die Aufarbeitung der alten Sammlungen mit der of three chronologically succeeding faunal assemblages lokalen Stratigraphie und der Vergleich mit den Neufun­ MLN, MYT, M TL, ranging from the uppermost early den, sowie die Miteinbeziehung der magnetostratigraphi­ Turolian to late middle Turolian. The updated correla­ schen Ergebnisse der fossilen Mytilinii Formation führte tion of the old collections with the local stratigraphy and zu einer präzisen Datierung aller Fundstellen: a. Q5-? their comparison with the new collection, as well as the Limitzis, unterstes MN13, 6.9-6.7 Ma; b. Q l, QA, S3,4, magnetostratigraphic study of the fossiliferous Mytilinii Adrianos, M TL, MN12, ~7.1 Ma; c. Q3, S2,3, Potamies, Fm allow the precise dating of all fossil sites: a. Q5-? MYT, MN 12, ~7.3 Ma; d. Q2, Stefano, MLN, unterstes Limitzis, lowermost MN 13, 6.9-6.7 My; b. Ql, QA, MN12, ~7.5 Ma; e. Qx, Vryssoula, upper part of M N ll, S3,4, Adrianos, MTL, MN 12, ~7.1 My; c. Q3, S2,3, 8.0-7.6 Ma. Die Kombination der alten und neuen Daten Potamies, MYT, MN 12, ~7.3 My; d. Q2, Stefano, MLN, der turolischen Säugetierfundstelle von Samos erlaubt eine lowermost MN 12, ~7.5 My; e. Qx, Vryssoula, upper part Aufstellung von vier Evolutionsstadien und verwirft damit of MN 11, 8.0-7.6 My. The combination of old and new den alten „single fauna“-Ansatz. data concerning the Samos Turolian mammal faunas implies the establishment of four stages of evolution and Schlüsselworte: Obermiozän, Samos, Griechenland, refutes the “single fauna” approach. Säugetiere, Chronologie.

Keywords: Late Miocene, Samos, Greece, Mammalia, Chronology. 1. Introduction

The Late Miocene fossiliferous deposits of Samos Island, Zusammenfassung Greece, have been known since the second half of the 19th century, when Forsyth-Major discovered them and Die eingeschränkte Korrelation der alten Aufsammlun­ collected the first fossils. Then afterwards, several scien­ gen mit den Fossilfundstellen einerseits und der lokalen tists, as well as fossil collectors and dealers visited Samos Stratigraphie andrerseits verhinderte eine sichere zeitliche and gathered fossils for various museums and institutions (K o u f o s , this volume-a). There is a great number of pub­ lications concerning the Samos fauna, referreing either to

Prof. George D. K o u f o s , Dr. Dimitris S. K o s t o p o u l o s 6c vertebrate paleontology or to chronology(M a j o r , 1888,

MSc. Theodora D. V l a c h o u , Aristotle University of Thes­ 1891, 1894; A n d r e w s , 1896; S c h l o s s e r , 1899, 1906; saloniki, Department of Geology, Laboratory of Geology 8c O s b o r n , 1898; S t u d e r , 1911; A n d r e e , 1926; B r o w n , Paleontology, 54124 Thessaloniki, Greece, e-mail: koufos@ 1927; C o l b e r t , 1941; W e h r l i , 1941; v a n C o u v e r i n g & geo.auth.gr, [email protected]; [email protected] M i l l e r , 1971; G e n t r y , 1971; S o n d a a r , 1971; H e i s s ig , ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

398 Beitr. Palaont., 31, Wien, 2009

1975; M e i s s n e r , 1979; B l a c k et al., 1980; S o l o u n i a s , Q l-6 = Quarryl-6, Samos, Greece 1981; K o u f o s S c M e l e n t i s , 1982,1984; W e i d m a n n et Qx = Quarry x, Samos, Greece al., 1984; S e n & V a l e t ; 1986; B e r n o r et al., 1996; and S = Stefano, Samos, Greece literature cited in these articles). In these studies, a great A = Adriano, Samos, greece number of fossils has been described, numerous species S2-4 = Solounias collection from Samos, Greece established and several opinions about the age of the Samos L = Limitzis site, Samos, Greece fauna have been proposed. Nevertheless, the absence of G = German quarries, Samos, Greece accurate locality descriptions from the old collections and the uncertainties about the local stratigraphy continued to plague biostratigraphic and age determinations and 2. Historical Overview left vague taxonomic definitions. Exceptions are Forsyth- Major’s collection in Lausanne (MGL) and B. Brown’s The Samos vertebrate fauna was considered equivalent collection at AM N H . Forsyth-Major marked the fossils to that of Pikermi for a long time as and was referred he collected as originating from ‘Stefano’, ‘Potamies’ and to as Pontian (Late Miocene/Early Pliocene), with the 'Adriano’, all referring to local place-names, whereas old-fashioned use of this term. A t first sight, the faunal Brown used a register code of seven “quarries’ (Qx, Ql-6), assemblages provided from these two Greek localities look corresponding to precise fossil sites. A first serious effort similar, but a thorough study indicates several differences. towards the relocation of the localities and their correlation The first serious efforts concerning age determination of with the various museum collections has been undertaken the Samos mammal fauna started in the 1970ies. At that by S o l o u n i a s (1981). The author combined personal field time v a n C o u v e r i n g & M i l l e r (1971), using Argon and laboratory observations with information from the Isotope Analysis, provided absolute radiometric datings available field books of B. Brown, the museum archives and for Samos Neogene volcanoclastic deposits, influencing the local people of Samos. But, as he realized, it is hard to various palaeontologists dealing with museum collec­ give a definite answer about the provenance of the entire tions to comment on the biochronological meaning of fossil collections as most of them were made by amateurs, the Samos fauna. sometimes including specimens purchased from villagers S o n d a a r (1971) studied the hipparion assemblage (mainly (K o u f o s , this volume-a). In the early 1970ies, absolute skulls and metapodials) of Samos, housed at AM NH, chronological methods were also used for the chronology and although he stated that “the phylogeny of the Samos of the Samos fauna, but again, due to the problems did Hipparion remains speculative” he realized that differ­ not result in a final solution as the correlation of the faunas ent Hipparion species from the various fossiliferous sites with the stratigraphic and sampling horizons remained might represent chronologically distinct assemblages questionable. (Table 1). The same author tried to check if there were W ith the aim to solve the Samos puzzle and its negative differences in other animal groups of the Samos fauna impact on European biochronology and mammalian sys- and he studied the aardwark Orycteropus gaudryi from the tematics, a team of palaeontologists from the Laboratory of Brown quarries (S o n d a a r , 1971). Comparing the length Geology and Palaeontology, University of Thessaloniki, led of the upper and lower molar row versus M 2 or m2, he by G.K., started a new series of excavations in 1993 (K o u ­ found that 0. gaudryi from Q5 is larger than that from f o s et al., 1997,2004). The main goal of this new campaign Q l-4 (S o n d a a r , 1971:figs. 4, 5). Thus, he supposed an was the relocation of the fossiliferous sites, their arrange­ age difference between the two faunas, accepting that the ment in a precise stratigraphic order, the collection of new size increase in this genus is an evolutionary trend, as the fossils and the dating of the faunas using biochronology recent 0. eriksoni is much bigger. G e n t r y (1971) arrived and magnetostratigraphy. The results of the first 12 years at a similar conclusion, studying “Pachy tragus” samples of this campaign are included in this volume. at AM N H from Q l-4 and Q5, whereas H e i s s ig (1975) distinguished four rhinocerotid assemblages on Samos and Abbreviations: later also indicated a time-distance between Q l-4 and Q5 AMNH = American Museum of Natural History, New York mammal faunas (Table 1). ELMZ = European Land Mammal Zones A ll this data failed, however, to reliably correlate with the MGL = University of Lausanne, Samos Collection radiometric datings of v a n C o u v e r i n g & M i l l e r (1971), PMMS = Melentis collection, Aegean Museum of Natural that gave unexpectedly high values, ranging from 20.8±1.7 History (NHMA), Samos M y to 7.4±0.6 M y (Table 1). As the authors mentioned, MLN = Mytilinii-4, Samos, Greece “.. .there is nothing in the geology of the basin to suggest M YT = Mytilinii-3, Samos, Greece that the pumice-breccia to which this date refers (i.e., the MTL = Mytilinii-1, Greece sample GM -101 dated at 20.8±1.7 M a and 18.5±1.5 M a ) M TLA = M ytilinii-lA, Samos, Greece should be as much as 10 m.y. older than the others”. Based MTLB = Mytilinii-lB, Samos, Greece on the three best agesv a n C o u v e r i n g & M i l l e r (1971) MTLC = Mytilinii-lC, Samos, Greece suggested an age of 9.3 M y for Q l-4. MTLD = Mytilinii-ID, Samos, Greece Later, B e r n o r (1980) studied the hipparions from Maragheh NOW = Neogene Old World database (Iran) and compared them with the Samos ones. Using the QA = Quarry-A, Samos, Greece chronological data from Maragheh he also suggested two ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Kostopoulos, D.S. & Vlachou, Th.D., Biochronology. 399

GENTRY (1971) 1 VAN COUVERING WEIDMANN ct al. BERNOR et al. SW ISHER III PRESENT CHRONOLOGY SONDAAR (1971) i &. MILLER (1971) (1984) (1996) (1996) HEISSIG (1975) ! OLD LOCALITIES NEW LOCALITIES ELMA 351

------SKI=6.14±0.19 ------,

SK2=6.14±0.04

R105=6.74±0.11 ------SfC 19=5.41 ±0.17------n i _ f 7.23±0.I4\ GM106{S0±08r 8 .9 ± 0 .9 Q5 }^5 \ R106=6.89±0.06 lL A *SK5=7.75±0.08

?SK5=8.580±0.010

GM102=9.2±0.7 7 .5 5 ± 0 .U R 1 0 2 { Q2.3 7,4 2 ± 0 .0 6 S2-4 SK18A=7.52±0.16

SKI8A=7.276±0.006

Q4 Q4 R102=7.660±0.010 Q2 Stefaoo

r 8.5 7 ± 0 .2 8 G S K I 6 { s.S 8 ± 0 .0 7

G Q6 G Q* SK3=8.26±0.08 SK3=8.26±0.008 •Q” SK6=7.80±0.40 SK16=8.38±0.07 Qx, Vryssoula

Qx Qx

NormaI= 4,Ar/MAr date; llalics=K-Ar date

Table 1: Chronostratigraphic position of Samos fossiliferous sites according to several authors associated by radiometric dating and magnetochronology. (1): lithostratigraphic division according to W eidmann et al. (1984) and Kostopoulos et al. (this volume).

fossiliferous levels for Samos, in agreement withS ondaar et al., 1984) in accordance with S olounias (1981) “single­ (1971) and G entry (1971). The first level was considered as fauna” hypothesis. being early and the other middle-late Turolian. The first magnetostratigraphic data for the late Miocene One year later S olounias (1981) published a thesis on deposits of Samos was published during the mid-1980ies the history of the Samos fossil sites and collections and by S en & Vale t (1986). The authors sampled a 132 m studied the carnivores and bovids (Table 1). He also pro­ thick section, covering the upper part of the fossiliferous vided revised faunal lists for the various Samos sites, but Mytilinii Fm, without, however, any reference to precise the number of registered taxa was extremely high, not only fossil sites. The authors estimated a 6.4-6.1 My age for significantly exceeding the usual number of mammals the Q5, Q l, L and A sites. The old radiometric samples ever recorded in the late Miocene European faunas but of W eidm ann et al. (1984) were lately re-dated with the also in recent ones. Even more surprisingly, the author 40Ar/40Ar-method (Sw ish e r III, 1996). The proposed age diverged from all known approaches at that time and he for the basal part of the Mytilinii Fm (Old M ill Beds of proposed that the fossiliferous horizons of Samos were W eidm ann et al., 1984) ranges between 8.38+0.07 and deposited during a short-timed interval and consequently 8.26+0.08 My, for the lower part of the main fossilifer­ the included mammal faunas represent an isochronous and ous beds (Main Bone Beds) from 7.66±0.01-7.28±0.01 homogeneous assemblage. This idea was going to radically My, for the upper part of the main fossiliferous beds from affect most of the following works. 7.09±0.01 M y and for the uppermost part of the Mytilinii W eidm ann et al. (1984) gave new absolute datings for Fm (Marker Tuffs) between 5.41±0.17 and 6.74±0.11 My. the Neogene deposits of the Mytilinii Basin (Table 1). Based on these datings, B ernor et al. (1996) tried to cor­ Using the K/Ar-method and the stratigraphy proposed relate the Maragheh, Samos and Pikermi faunas, combin­ by S olounias (1981), they dated the base of the Mytilinii ing faunal and chronological data. Concerning Samos, Formation at ~8.5 My, the main fossiliferous beds at ~7.35 the authors suggested (Table 1) that the lower part of the M y and the uppermost part of the Mytilinii Formation Mytilinii Fm can be correlated to MN 11 with an age of at -6.18 My. The authors stated that it was incorrect to -8.34 My, the main fossiliferous level to MN 12 with an separate the Samos mammal fauna into two associations age > 7.1 M y and the uppermost part of the Mytilinii Fm (Q l-4 and Q5), and concluded that all bone-bearing beds (Marker Tuffs) to MN 13. In spite of opposing evidence, had been deposited between 7.0 and 8.5 M y(W eidm ann the “single-fauna” hypothesis was not explicitly abandoned. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

400 Beitr. Palaont., 31, Wien, 2009

GPTS M y MN

B

Figure 1: Magnetostratigraphic correlation between the sampled sections of Mytilinii basin, Samos and correspondence of the com­ posite magnetostratigraphic section with GPTS and MN-zones (after K o s t o p o u l o s et al., 2003, modified). F = fossil levels. MN boundaries according to A g u s t i et al. (2001), GPTS according to B e r g g r e n et al. (1995)

3. Magnetostratigraphy give a short review of this work in order to correlate the bio- and magneto-chronological data. 178 horizons from As it was already mentioned, the magnetostratigraphy two main and five secondary sections, including the whole of the fossil-bearing Mytilinii Fm was partially studied Mytilinii Fm, as well as part of the underlying Hora Fm, earlier by S e n & V a l e t (1986), whose results allowed and the overlying Kokkarion Fm have been sampled and them to choose between the 8.7-7.4 M y and the 6.8-5.7 analysed (K o s t o p o u l o s et al., 2003). The correlation of M y intervals. Based on the available radiometric data by magnetostratigraphic data from individual sections led to a W e i d m a n n et al. (1984), and on the H a r l a n d et al. composite magnetostratigraphic column on which the new (1982) polarity time scale, both sources of evidence being mammal sites were correctly placed. W e also successfully extensively reviewed during the last decade, the authors coped with the problem of the stratigraphic location of suggested a 6.8-5.7 M y age for the upper part of the M y­ the old quarries, using all the available information given tilinii Fm (K o s t o p o u l o s et al., 2003). During the end of by the previous researchers, mainlyS o l o u n i a s (1981), as 1990ies we decided to re-study the magnetostratigraphy well as field and personal observations of old collections of the Mytilinii Fm and the results were presented by (K o s t o p o u l o s et al., 2003). The correlation of the com­ K o s t o p o u l o s et al. (2003). In the present article we will posite magnetostratigraphic section with GPTS (Fig. 1) ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien suggested the following age determinations(K ostopou- Uppermost part of Mytilinii Fm. It includes the old los et al., 2003): locality Q5 of Brown (Tab. 1). According to our mag- Basal part of Mytilinii Fm. It includes the localities Qx netostrati-graphy, this part of the formation should be and Vryssoula (probably a single site), situated at the NE correlated with chrons C3Ar-C3Bn, corresponding to an border of Mytilinii village inside an army campus(S olou- age between 7.1-6.5 Ma; an age estimation for Q5 should n i a s , 1981; pers. obs). This part of the Mytilinii Fm can be be 6.9-6.7 My. correlated with Chron C4n.2n, corresponding to 8.0-7.6 Figure 1 and Table 1 summarize the chrono-stratigraphic My, implying that Qx and Vryssoula correspond to the distribution of the old and new Samos sites, updating data upper part of early Turolian (MN 11). As the localities are by Kostopoulos et al. (2003). situated above the middle of the basal part of Mytilinii Fm it is clear that the deposition of the formation started within the Turolian. 4. Biochronology - Correlation with ELMZ Lower part of the main fossiliferous beds. It includes the new locality MLN, as well as ‘Stefano’ of Forsyth-Major Two main issues concern the biochronology of the Samos (Tab. 1). Following the topographic re-location of the old fauna: a) its relations with the European Land Mammal quarries by Solounias (1981) we have directly but errone­ Zones (ELMZ; Steininger, 1999), and b) the internal ously correlated MLN with Brown’s Q4 (Kostopoulos structure of the Samos mammal assemblages and their et ah, 2003). New data (Kostopoulos, this volume-a) possible discrimination into chronologically succeeding allows us to realize that for inexplicable reasons, the loca­ stages. The study of the new collection from the mammal tion of Q4 has been confused with that of Q2 and vice localities of Samos led to the determination of a quite versa. Correcting this mistake, we now suggest correlating rich fauna. The available material was collected from the MLN with Q2, both placed in the same fossil horizon localities MLN, M YT and MTL; the last locality includes with ‘Stefano’. This part of the section can be correlated to three fossiliferous sites, M T LA, M TLB and M T LC Chrons C3Br.2n-C4n.ln (between 7.45 and 7.65 My) with (details about the localities are given in K ostopoulos an average 7.5 M y of age for MLN, Q2 and ‘Stefano’. This et al., this volume). The faunal composition of M TL is age corresponds to the end of early Turolian, MN 11. the richest one and gives accurate biochronological data. Upper part of the main fossiliferous beds. The old locality Although the faunas of MLN and M YT are poor, they Q6 is now placed near its base (Tab. 1). It is situated at the give some biochronological evidence, which together with northern border of the Mytilinii basin, near the village of magnetochronology allow certain age determinations. The Kokkarion, and at the beginning of Tholoremma ravine. biochronology of each locality will be given separately, Although the locality has been relocated, we did not yet beginning with the oldest fauna. excavate there. Brown’s Q6 site was indirectly correlated Biochronology of Mytilinii-4 (MLN). The locality MLN with the radiometric samples SK3 (8.26±0.8 My) and SK6 is situated at the base of the main fossiliferous beds of (7.8±0.4 My) of W eidmann et al. (1984), and used to be Mytilinii Fm in Potamies ravine (Kostopoulos et al., placed together with Qx into the lowermost fossil-level this volume). The determined fauna is relatively poor, (Bernor et al. 1996). However, the collected poor fauna including the following taxa: Hyaenictitherium cf. w ongii, from this site undoubtedly includes Samotherium major; Protictitherium crassum, Hipparion aff. proboscideum, Hip- which certainly implies an age no older than 7.4 My, i.e. parion aff. prostylum, “Diceros” neumayri, Palaeotragus younger than Qx and even younger than MLN, Q2 and rouenii, Palaeotragus sp., Samotherium boissieri, Gazella ‘Stefano’, and maybe older than M YT (Kostopoulos, pilgrim i, Tragoportax sp., Miotragocerus sp., Walaeoryx sp. this volume-a). (Kostopoulos, this volume-a, b; Koufos, this volume-b; The new locality MYT is identical with Brown’s Q3 placed Vlachou Sc Koufos, this volume). in the middle of the main fossiliferous part of Mytilinii Most of the available taxa indicate Turolian age. Protic­ Fm (Tab. 1); S2-3 of Solounias (1981) are on the same titherium crassum is known from several Eurasian localities level, whereas ‘Potamies’ of Forsyth-Major would also be and has a long stratigraphic range from middle Miocene contemporaneous. This part of the section is correlated MN 6 up to late Turolian MN 13 (Koufos, this volume- to Chron C3Br.2r, suggesting an age of ~7.3 M y for the b). The MLN Protictitherium sample belongs to the large M Y T fauna, which should, therefore, be placed in the forms of the species, like that found in Dytiko (Axios middle Turolian, MN 12. According to the updated valley, Greece), suggesting a Turolian age. faunal data (Kostopoulos, this volume-a, b; Vlachou The hipparion sample from MLN is poor and indicates Sc Koufos, this volume), the Q4 site of Brown should be the presence of two species: a medium-sized species re­ slightly younger. sembling H. prostylum and a large-sized one, known only The new locality M T L (including the sites M T L A , by postcranials, that could be ascribed to H. proboscideum MTLB, MTLC), as well as the old localities ‘Adriano’ of by their size. The type locality ofH. prostylum is Mont Forsyth Major, ‘Adrianos’ of Melentis and Q l of Brown, Luberon, (France) dated to middle Turolian, MN 12 are located in Adrianos ravine, in the upper levels of the (Bernor et al., 1996; NOW 2007). The species is also main fossiliferous beds. This fossil level is correlated to known from the middle Maragheh and Pikermi(B ernor Chron C3Br.ln with an estimated age of 7.13-7.17 My, et al., 1996), dated from early to middle Turolian (Koufos, corresponding to the end of middle Turolian, MN 12. 2006; NOW, 2007). The type of the large-sizedH.probos- ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

402 Beitr. Palaont., 31, Wien, 2009

cideum comes from Samos but from an unknown locality et al. (2003) register the species in locality 26 of Middle (S o n d a a r , 1971); nonetheless, its presence in Brown’s Sinap (Turkey), dated at 8.1 My. Thus, the co-existence Qx indicates that the species probably originates from the o f “Diceros” neumayri, Dihoplus pikermiensis and Ancyloth­ lower fossil levels (V l a c h o u 6 c K o u f o s , this volume). H. erium pentelicum is rather indicative of an early middle proboscideum is certainly known from the locality Ravin Turolian age. des Zouaves-5 of Axios valley (Macedonia, Greece) dated The identified hipparions from M YT are slightly different to MN 11, more precisely at ~8.2 M y by magnetochronol­ than those from MLN. H. prostylum is probably still present ogy (K o u f o s , 1987, 2006). A large-sized hipparion from in MYT, whereas H. cf. proboscideum is known only by a few the Turkish locality of Kemiklitepe-A, B, dated to middle postcranial elements (V l a c h o u 6 c K o u f o s , this volume). Turolian, at ~7.2 M y (K o u f o s 6 c K o s t o p o u l o s , 1994; S e n H. matthewi is originally described from an unknown lo­ et ah, 1994), resembles H. proboscideum, too. The MLN cality of Samos (A b e l , 1926) but is well-known from the H. aff. proboscideum sample is distinguished from the Q l middle Turolian sites of Kemiklitepe, Turkey(K o u f o s 6 c sample of the species by its more slender postcranials that K o s t o p o u l o s , 1994 ).H. forstenae was originally described indicate a more primitive stage(V l a c h o u & K o u f o s , this from Loc. 30 of Shanxi, China, and it seems to be a time- volume). “Diceros” neumayri is a wide-spread rhinoceros spread species ranging from Vallesian to Turolian. A similar with a very long range from MN 9 to the end of MN 12 form is known from Giilpinar, Turkey (recorded asH. mat­ and with a clear tendency of size-increase through time thewi) and Titov-Veles, FYR of Macedonia (recorded asH. (H e i s s ig , 1975; G iaourtsakis , this volume). The MLN verae), both dated to middle Turolian, MN 12 (F o r s t e n 6 c rhinocerotids are, however, too poor for certain chrono­ G a r e v s k i , 1989; F o r s t e n 6 c K a y a , 1995). H. cf. forstenae logical suggestions. from M YT represents the first appearance of the species The MLN giraffids and bovids are rather point to an early in the Samos faunal succession. Turolian age. Palaeotragus rouenii has a wide chronos- The large palaeotragine Samotherium major replaces its tratigraphic range from late Vallesian (MN 10) to latest forerunner S. boissieri at about 7.35 My, appearing for the Turolian (MN 13) but its coexistence with a sturdier first time in the Q6Tevel (K o s t o p o u l o s , this volume-a). palaeotragine, Palaeotragus sp. in MLN is rather indica­ S. majorss well-documented in M YT by a form of slightly tive of an early Turolian, MN 11 age(K o s t o p o u l o s , this smaller size than that occurred in the overlying Samos volume-a). A robust Palaeotragus is usually present in lat­ fossil levels, suggesting an earlier age. est Vallesian-early Turolian faunas from Turkey, Greece Gazella pilgrimi is the predominant gazelle species in the and the Black Sea region, while it is much more uncom­ early Turolian localities of Macedonia, Greece, where it mon in middle Turolian faunas. Samotherium boissieri is fades out at the beginning of middle Turolian(K o s t o p o u ­ originally known from Samos, but is also documented l o s , 2006). The species occurs in the lower fossil levels

at Kemiklitepe D, dated at ~7.7 M y (S e n et al., 1994). of Samos probably together with G. cf. ancyrensis but it Gazella pilgrimi also implies chronological similarities is still present in MYT, MTLA/B and Q5, suggesting a with early Turolian faunas from continental Greece. On much wider time-distribution than in continental Greece the other hand, Palaeoryx is a common element in the (K o s t o p o u l o s , this volume-b). Palaeoryx sp. and Spora­ middle Turolian faunas of the Greek mainland. Hence, dotragus parvidens are typical middle Turolian bovids of the combination of early Turolian elements with some continental Greece, originally known from Pikermi, but middle Turolian ones suggests a late early Turolian age they seem to have a wider time distribution in the East, (late MN 11) for MLN, in accordance with the available going down to early Turolian (K o s t o p o u l o s , this volume- magnetostratigraphic data. b). S. parvidens is already well-known from Kemiklitepe

Biochronology of Mytilinii-3 (MYT). The locality MYT D, dated at ~7.7 M y (S e n et al., 1994). Skoufotragus ze­ is situated at the basal part of the main fossiliferous beds of malisorum is a possible forerunner ofSkoufotragus laticeps Mytilinii Fm, in Potamies ravine (Tab. 1). The determined fPachy tragus laticeps) from the main Samos fossil levels fauna is poor in identifiable specimens but includes an (K o s t o p o u l o s , this volume-b). important number of taxa: “Diceros” neumayri, Dihoplus In summary, a few early Turolian mammal taxa persist pikermiensis, Ancylotheriumpentelicum, H. cf. proboscideum, in the MYT fauna, which is basically characterized by H. cf. forstenae, H. prostylum, H. cf. matthewi, Samotherium the first appearance of several middle Turolian elements. major.; Sporadotragus parvidens, Gazella pilgrimi, Skou- Thus, an early MN 12 age, totally compatible with the fotragus zemalisorum n. sp., Palaeoryx sp., }Majoreas sp. magnetostratigraphic data, is suggested. (G iaourtsakis , this volume; G iaourtsakis & K o u f o s , Biochronology of M ytilinii-1 (MTL). The locality this volume; K o s t o p o u l o s , this volume-a, b; V l a c h o u M TL is situated in Adrianos ravine and includes several 6c K o u f o s , this volume). fossiliferous sites, from which a rich mammal fauna has “Diceros” neumayri continues its presence in MYT.Ancy­ been unearthed; the fauna includes both micro- and lotherium pentelicum is a rare taxon known from Pikermi, macro-mammals. upper Maragheh and Akka^dagi, suggesting a middle Tu­ M T LA. Pseudomerionespythagorasi, lKarminata provocator, rolian age; its record in M Y T most probably corresponds Spermophillinus cf. bredai, Adcrocuta eximia, Hyaenictith- to the appearance of the species and its first occurrence in erium wongii, Machairodus giganteus, Metailurus parvulus, the Samos faunal succession. H e i s s ig (1996) lists Dihoplus Parataxidea maraghana, Zygolophodon turicensis, Orycteropus pikermiensis from Samos and Pikermi whereas F o r t e l i u s gaudryi, “Diceros” neumayri, Dihoplus pikermiensis, Ancyloth- ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

erium pentelicum, Hipparion brachypus, Hipparion dietrichi, eximia (B onis 8 c K o u fo s, 1981). The resemblance of the Hipparion cf. proboscideum, Hipparion cf. matthewi, Hip­ studied material from M TL to that of the Turolian sub­ parion cf. forstenae, Microstonyx major, Palaeotragus rouenii, species A. e. eximia indicates a Turolian age for M TLA. Samotherium major, Helladotherium duvernoyi, Gazella pil- Machairodus giganteus is possibly known from M T L A by grim i, Gazella cf. capricornis, Gazella mytilinii, Miotragocerus a fragment of an upper canine in the PM M S collection valenciennesi, Sporadotragusparvidens, Skoufotragus laticeps, (K oufo s, this volume-b). Its resemblance to the material Palaeoryxpallasi, Urmiatherium rugosifrons. from Vathylakkos and Halmyropotamos (Greece) sug­ M TLB. Pseudomeriones pythagorasi, Spermophillinus cf. gests a middle Turolian age (K oufos, this volume-b). The bredai, Pliospalax cf. sotirisi, Plioviverrops orbignyi, Hy- MTLB Plioviverrops orbignyi is similar to that from early aenictitherium wongii, Ghoerolophodonpentelici, Orycteropus middle Turolian localities of Axios valley and Thessaly gaudryi, “Diceros” neumayri, Ancylotherium pentelicum, (Greece), as well as to the middle Turolian sample from Hipparion brachypus, Hipparion dietrichi, Hipparion cf.pro­ Pikermi (K o u fo s, this volume-b). The felid M etailurus boscideum, Hipparion cf. matthewi, Hipparion cf. forstenae, parvulus is mainly known from middle-late Turolian, Palaeotragus rouenii, Palaeotragus sp., Samotherium major.; although there is a single mention of its presence in the Gazellapilgrimi, Gazella cf. capricornis, Gazella mytilinii, late Vallesian locality of Montredon, France (K o u fo s, Miotragocerus valenciennesi, Tragoportax rugosifrons, Skou­ this volume-b). The similarity of the studied material fotragus laticeps, Palaeoryx pallasi, Palaeoryx majori. from M T L A to that from the Greek localities Pikermi, M TLC. Hyaenictitherium cf. wongii, Pliohyrax graecus, Chomateres, Halmyropotamos and Kerassia suggests late Samotherium major, Miotragocerus valenciennesi, Gazella middle Turolian (MN 12) age. The mustelid Parataxidea cf. capricornis, Palaeoryx majori. maraghana from M T L A establishes chronological rela­ (G iaourtsakis, this volume; G iaourtsakis8 c K oufos, this tions with middle Maragheh (Solo un ias, 1981; B ernor volume; K onidaris 8 c K oufos, this volume; K ostopoulos, et al., 1996). this volume-a, b; K oufos, this volume-b; Sylvestrou 8 c The rhino assemblage of MTL is identical to that of MYT, K ostopoulos, this volume; Vasileiadou 8 c Sylvestrou, including “Diceros” neumayri, Dihopluspikermiensis z.n& An­ this volume; Vlachou 8 c K oufos, this volume). cylotherium pentelicum, indicating middle Turolian, M N12 The rodent Pseudomerionespythagorasiwas found in M TLA age. Furthermore, the rich material of “D.”neumayri from and MTLB, but B lack et al. (1980) had already recorded M TL shows a more advanced morphology than Pikermi the species in S3 site, i.e. near the M YT fossil level. Recent and a lesser one than Akkajdagi, Turkey, being closer to data about the phylogeny of the genus suggests that it is a the Turkish samples from Mahmutgazi and Kinik, indi­ descendant ofPseudomeriones latidens, known from the late cating a late middle Turolian age (G iaourtsakis, this Vallesian - early Turolian of Turkey, whereasP. pythagorasi volume; G iaourtsakis 8 c K oufos, this volume). is classified as middle Turolian, M N -12 (Sylvestrou 8 c Five different hipparions have been recognized in M TL. K ostopoulos, 2007). Apart from Samos, P. pythagorasi As already mentioned Hipparion proboscideum and H. is also known from the locality Diizyayla, Turkey, dated forstenae suggest an early-middle Turolian age. H. cf. to MN 12 (Fah lbu sch , 1996). “ Karminata” provocator m atthewi from M TL is characterized by a larger size than shares several morphological characteristics with that MYT, suggesting a younger age (V lachou 8 c K o u fo s, from Pikermi (Vasileiad o u 8 c Sylvestrou, this volume), this volume). H. dietrichi is a medium- to large-sized form, implying a similar middle Turolian age. the holotype of which originates from an unknown local­ The mastodonts found in MTL are few and include ity on Samos (S o n d a a r , 1971). Although H. dietrichi is two species, Choerolophodon pentelici and Zygolophodon known from early-middle Turolian sites of continental turicensis. The late Miocene choerolophodonts are sepa­ Greece (K oufos, 1987a, b, 1988; V lachou 8 c K o u fo s, rated into three species: C. corrugatus, C. anatolicus and 2002, 2006), new data suggests that the mainland form C. pentelici. C. anatolicus is Vallesian and the other two might represent a distinct taxon (V lachou, in prep). H. Turolian (San d ers, 2003). The similarity of the MTL dietrichi probably derives from Q6 and the M Y TH. pros- choerolophodont with C. pentelici suggests Turolian age tylum (V lachou 8 c K oufos, this volume) and it represents (K o n idaris 8 c K oufos, this volume). The Zygolophodon the first occurrence of the species in the Samos faunal turicensis specimen belongs to the PM M S collection and succession. A more advanced form ofH. dietrichi is also possibly originates from M TLA. This taxon has a wide recorded in Akkajdagi, Turkey, dated to 7.0 My(K oufos time-distribution in Eurasia covering middle-late Miocene 8 c V lach ou, 2005; K arad en izli et al., 2005). The type (NOW, 2007). The sole known dp4 is similar to those of H. brachypus is known from Pikermi, Greece, while from the Greek localities of Ravin de Zouaves-5 (Axios the species is also documented in Hadjidimovo, Bulgaria, Valley) and Pikermi, dated to early and middle Turolian both evidences implying a MN 12 age (H ensel, 1862; respectively. K oufos, 1987; H risto va et al., 2002). The presence ofH. The carnivores from M TL are relatively abundant and offer brachypus in M TL represents the first certain occurrence some additional biochronological data. Hyaenictitherium of the species in the Eastern Mediterranean and its first w ongii has a wide chronostratigraphic range, covering the appearance in the Samos faunal succession. whole late Miocene. Adcrocuta eximia also shows a long The large morphotype ofMicrostonyx major from M T L A time-distribution but it is known by two subspecies, the is indicative of a late middle Turolian age, analogous to Vallesian A. eximia leptoryncha and the Turolian A. eximia that of Pikermi, Greece, and Kalimantsi, Bulgaria (Sy l - ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien vestrou & K ostopoulos, this volume). Palaeotragus The reader should already have realized from the thorough rouenii continues its presence in M TL. Samotherium major study of the new collection that the Samos mammal fauna from M TL appears to be larger than that from MYT, sug­ is in no case homogeneous and isochronous. Originally gesting a younger age(K ostopoulos, this volume-a). The based on bovid data (Kostopoulos, this volume-b) we species characterizes middle Turolian faunas, being present tried to restore this unfortunate misinterpretation and in Vathylakkos (Greece), Kemiklitepe A/B, Akka^dagi to credit the Samos fauna with its real meaning and and Taskinpa§a (Turkey), dated to middle-late Turolian importance. Thus, we combined information from old (Kostopoulos, this volume-a). The first appearance of and new collections in order to create a time-distribution Helladotherium duvernoyi in M T L A is also in favor of a table of Samos taxa (Fig. 2), illustrating the changes in late middle Turolian age. Although the species shows a the internal structure of the Samos mammal assemblage wide time-distribution from late Vallesian to latest Tu­ through time. Four chronologically succeeding mammal rolian, its signal significantly increases in the upper part assemblages can be made out, reflecting a “four stages- of middle Turolian, documented in Pikermi, Kerassia, of-evolution” scheme. Perivolaki (Greece), Hadjidimovo, Kalimantsi (Bulgaria) Primary Mammal Assemblage of Samos (PMAS), 7.8-7.4 and Akkajdagi (Turkey) (Kostopoulos & K oufos, 2006; My, late MN 11. Kostopoulos, this volume-a). PM AS (Fig. 2) is characterized by the presence ofProtic- Gazella pilgrim iand Sporadotragus parvidens, already known titherium eras sum, Promephitis lartetii, “Diceros” neumayri, from earlier Samos levels, persist in M TL.Miotragocerus Hipparionprostylum, Hipparionproboscideum, Microstonyx valenciennesi shows a wide time-distribution, covering the major, Samotherium boissieri, Palaeotragus rouenii, Palae­ entire Turolian. Tragoportax rugosifrons is considered to be otragus quadricornis, Miotragocerus vallenciennesi, Tragopor­ an early Turolian element, disappearing in the Balkans tax rugosifrons, Sporadotragus parvidens, Gazella pilgrimi, at the beginning of M N 12. Nevertheless, it seems to Gazella cf. ancyrensis, Majoreas woodwardi, Criotherium last longer in the East (Kostopoulos, this volume-b). argalioides, Tragoreas oryxoides, Prostrepsiceros fraasi Palaeoryxpallasi and Gazella capricornis are typical middle Gazella mytilinii, Protoryx capricornis and Palaeoryx sp. Turolian bovids, originally known from Pikermi. Gazella According to data from Turkish sites (Heissig, 1996) cf. capricornis from M TL marks the first appearance of Chilotherium samium known from non-stratigraphically the species in the Samos faunal succession and shows a controlled old Samos collections, should also be credited great resemblance to the predominant Akka^dagi gazelle, to this assemblage. A t the end of this period, Majoreas suggesting a late middle Turolian age (Kostopoulos this woodwardi, Tragoreas oryxoides and Protoryx capricornis volume-b). Palaeoryx majori is also traced in the Samos probably disappeared. fauna for the first time. Although its holotype and the rest Intermediary Mammal Assemblage of Samos (IMAS), of the Samos specimens ascribed to this species have an 7.4-7.2 My, early MN 12. unknown origin, P. majori is known from Akka^dagi, Tur­ IM AS (Fig. 2) is characterized by an enormous renewal key, dated at ~7.0 My. Skoufotragus laticeps, also originally of the mammal fauna. Promephitis lartetii, “Diceros” neu­ known from Samos, replaces Sk. zemalisorum from MYT, mayri, Pliohyrax graecus, Microstonyx major, Palaeotragus and becomes the predominant bovid in the M TL fauna rouenii, Miotragocerus valenciennesi, Tragoportax rugosifrons, (Kostopoulos this volume-b). The species is also known Gazella pilgrimi, Criotherium argalioides and Sporadotragus from Kemiklitepe A/B, dated at ~7.2 M y(S e n et al., 1994), parvidens continue in this interval together withHipparion suggesting a middle Turolian age (MN 12). prostylum and Hipparion proboscideum. Simultaneously, Hence, the mammal assemblage of MTL is rather indica­ Pliospalax cf. sotirisi, Pseudomerionespythagorasi, Byzantinia tive of a late middle Turolian age (late MN 12), such as hellenicus, “Karnimata provocator, Spermophilinus cf. bredai, suggested by magnetostratigraphy. It is worth mentioning Adcrocuta eximia, Hyaenictitherium wongii, Orycteropus that small compositional differences between the faunal gaudryi, Dihoplus pikermiensis, Ancylotherium pentelicum, assemblage of the two M TL fossil levels, i.e. M TLA- Hipparion cf. matthewi, Hipparion ci. forstenae, Palaeoryx M TLC from the one site and M TLB-M TLD from the pallasi and Skoufotragus zemalisorum appear. Samotherium other, do not signify a faunal discrepancy, even though major replaces Samotherium boissieri and Prostrepsiceros the short stratigraphic distance between them certainly zitteli probably takes the place ofProstrepsicerosfraasi. Chi­ reflects a restricted time-lapse. lotherium kowalevskii from Samos should also be credited in this assemblage, probably coexisting with Chilotherium samium. 5. Internal Structure of the Samos Mammal Dominant Mammal Assemblage of Samos (DMAS), Fauna 7.2-6.9 My, late MN 12. The end of the previous period is characterized by the last Despite remaining minor inconsistencies concerning the occurrences of H. prostylum and Criotherium argalioides correlation of the Samos old fossil collections with specific and the first appearance ofHipparion brachypus, Hipparion chrono-stratigraphic horizons, the fossil assemblage of dietrichi, Skoufotragus laticeps and probably Chilotherium Samos offers an unprecedented panoramic aspect of the schlosseri. evolution of middle Turolian mammal faunas, which was D M AS (Fig. 2) represents an advanced stage of IM AS neglected for a long time by the “single-fauna” hypothesis. characterized by a similar faunal composition but with ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Figure 2: Time distribution of Samos small and large mammal taxa and correlation with MN-zones and GPTS.

(1): Qx, Vryssoula fossil level; (2): MLN, Q2, Stefano fossil level; (3): MYT, Q3, fossil level; (4): MTLA/B/C, Q l, Adriano fossil level; (5): Q5 fossil level: Q6 is in between (2) and (3) and Q4 in between (3) and (4). Taxa marked with an asterisk are included in the new collection and described in the present volume. PMAS, IMAS, DMAS and FMAS represent primary, intermediary, dominant and final Samos mammal assemblage (for details see text).

a clear trend towards larger and/or more specialized Skoufotragus schlosseri substitutes the earlier Sk. laticeps ungulates and an enrichment in carnivores. Most IM AS and becomes the dominant bovid. The smallerHipparion taxa continue to be present, but “Diceros” neumayri, Hip- nikosi with avdeep narial opening replacesH. cf. matthewi. parion cf. matthewi, Samotherium major and Microstonyx Hystrixprimigenia is traced for the first time. major appear with larger morphotypes than previously, whereas Skoufotragus zemalisorum is replaced by the larger Skoufotragus laticeps and H. prostylum by H. dietrichi, both 6. Conclusion predominating among ungulates. New taxa also appear: Metailurus parvulus, Promeles palaeattica, Parataxidea The biochronological analysis of the new Samos mammal maraghana, Plioviverrops orbignyi, Ictitherium viverrinum, collection and its correlation with existing magnetostrati- Choerolophodon pentelici, Zygolophodon turicensis, Chilo- graphic data (Kostopoulos et al., 2003) allows dating therium schlosseri, Helladotherium duvernoyi, Gazella capri- the MLN assemblage to the end of early Turolian (late cornis, Urmiatherium rugosifrons and Palaeoryx majori. The MN 11), the M Y T assemblage at the beginning of early first occurrence of Oioceros wegneri, Tragoportaxpunjabicus Turolian (early MN 12) and the M TL assemblage at the and Samokeros minotaurus could also be also credited in this late middle Turolian (late MN 12). assemblage. A t the end of this periodPromephitis lartetii, Updated information concerning the chronostratigraphic Pliohyraxgraecus, H.proboscideum, H. brachy pus, Tragopor- location of old quarries and their correlation with new tax rugosifrons and Palaeoryx pallasi fade out. ones together with revised taxonomical data, leads us to Final Mammal Assemblage of Samos (FMAS), 6.9-6.7 refute the “single-fauna” hypothesis ofS olounias (1981) My, latest MN 12-earlyM N 13. and to propose four stages of evolution in the Samos FM AS (Fig. 2) is characterized by the reduction of the mammal fauna, representing a chronological succession overall number of mammal taxa and the replacement of of ~1.0 My. some of them by others.Adcrocuta eximia, Hyaenictitherium The results of our work resolve long-lasting chronological w ongii and Metailurus parvulus are still present, as are problems concerning one of the classical late Miocene Ghilotherium schlosseri, Hipparion dietrichi, Hipparion cf. European mammal localities and provide an unequivocal forstenae, Microstonyx major., Samotherium major, Palae- framework for the local and Eurasian biochronology, as otragus rouenii, Helladotherium duvernoyi, Miotragocerus well as for the study of mammalian evolution. valenciennesi, Gazella pilgrimi, Gazella cf. capricornis, Gazella mytilinii, Prostrepsiceros zitteli, Palaeoryx majori and Sporadotragusparvidens. Orycteropus gaudryi from Q5 7. Acknowledgements appears larger than that from Q 1-Q 4 (So n d a a r , 1971). Tragoportax amalthea could replace Tr. rugosifrons together The excavations on Samos have been supported by the Prefecture with the entrance ofTragoportaxpunjabicus and an overall of Samos and the “Konstantinos and Maria Zimalis” Foundation. increase of the boselaphine signal. The more specialized The Municipality of Mytilinii provided generous help too. The ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Koufos, G.D., Kostopoulos, D.S. & Vlachou, Th.D., Biochronology. 406

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