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Original article

A synopsis of the late Miocene Mammal Fauna of Samos Island,Aegean Sea, Greece§

Synopsis de la faune mammalienne du Miocène supérieur de l’île de Samos, Mer Egée, Grèce

George D. Koufos *, Dimitris S. Kostopoulos, Theodora D. Vlachou, George E. KonidarisAristotle University of Thessaloniki, Department of Geology, Laboratory of Geology and Palaeontology, 54124 Thessaloniki, Greece

Received 14 April 2010; accepted 9 August 2010

Available online 23 February 2011

Abstract

As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness andoverall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the localstratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of thefossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearerand more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (�1200identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to theend of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, theestablishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amendedmorphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a bettermorphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlationof the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for themammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammalassemblages reflecting a ‘‘four stages-of-evolution’’ scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland withthick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. TheSamos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.# 2011 Elsevier Masson SAS. All rights reserved.

Keywords: Turolian; Samos; Greece; Mammals; Stratigraphy; Chronology; Palaeoecology

Résumé

Comme l’une des localités les plus anciennes connues de vertébrés fossiles d’Eurasie, la faune du Miocène supérieur de Samos a suscité l’intérêtdes spécialistes par sa richesse et son importance. Néanmoins, des questions essentielles de taxinomie et des problèmes chronologiques ontobscurci sa valeur. L’étude détaillée de la stratigraphie locale, la récolte de nouveau matériel fossile et son étude, la révision des collectionsanciennes et de la magnéto-chronologie des dépôts fossilifères ont permit de rediscuter la plupart des problèmes dans un volume spécial édité en2009 par Koufos et Nagel et de donner une idée plus claire et précise sur la faune de Samos et son âge. Une synthèse de ce travail est donnée ici.L’étude systématique de la nouvelle collection (�1200 spécimens identifiés) permet d’identifier 42 espèces provenant de trois horizons fossilifères,allant de la partie supérieure du Turolien inférieur (MN11) à la fin du Turolien moyen (MN12). Les nouveautés taxinomiques sont la présence dugenre Protictitherium, découvert pour la première fois à Samos, la proposition du nouveau nom Skoufotragus pour Pachytragus Schlosser, avec lanouvelle espèce Skoufotragus zemalisorum, et la systématique modifiée des genres Pseudomeriones et Urmiatherium. En outre, six espècesd’Hipparion et quatre espèces de Gazella sont reconnues et une meilleure distinction morphométrique entre Samotherium boissieri et Samotheriummajor est réalisée. Un effort de corrélation des anciens sites fossilifères avec les nouvelles localités et la stratigraphie révisée du bassin de Mytiliniia permis de préciser l’âge des gisements. L’élaboration des données nouvelles et anciennes indique la présence à Samos de quatre assemblages de

Geobios 44 (2011) 237–251

§ Invited editor: Lorenzo Rook.* Corresponding author.

E-mail address: koufos@geo.auth.gr (G.D. Koufos).

0016-6995/$ – see front matter # 2011 Elsevier Masson SAS. All rights reserved.doi:10.1016/j.geobios.2010.08.004

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mammifères successifs, qui reflète un scénario en « quatre étapes évolutives ». Le paléoenvironnement de Samos au Turolien est déterminé commeétant un « bush » ouvert avec un étage herbeux à graminoïdes en C3. Une augmentation du caractère ouvert et sec du début à la fin du Turolienmoyen est également supposée. Les faunes de mammifères de Samos sont paléobiogéographiquement plus proches de celles d’Asie Mineure quede celles de Grèce continentale.# 2011 Elsevier Masson SAS. Tous droits réservés.

Mots clés : Turolien ; Samos ; Grèce ; Mammifères ; Stratigraphie ; Chronologie ; Paléoécologie

1. Introduction

The perfect preservation and the high abundance of fossilvertebrates, mainly mammals, in Samos Island (Greece)attracted the interest of scientists and amateurs since thesecond half of the 19th Century, when the expatriate Englishbotanist Charles Immanuel Forsyth Major visited the island andannounced his discoveries (Forsyth Major, 1888, 1894). Thenafter, several scientists and dealers (i.e., Stutzel, Sturz,Hentschel, Fraas, Acker, Brown, Skoufos, Melentis) worked

at Samos, and the great number of fossils unearthed wasdistributed in various museums and institutes of Europe and theUnited States. Solounias (1981) and recently Koufos (2009a)cited the chronicles and backstage stories of early and laterexpeditions dealing with Samos fossils.

Although vertebrate fossil collections from Samos arealmost everywhere across the world, most of them lack preciselocality indications. As a result, the correlation of fossils withthe local stratigraphy is limited if at all, and the old materialfrom different sites is frequently mixed, eliminating drastically

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Fig. 1. Geological map of the Mytilinii Basin with location of the old and new fossiliferous sites (geological map from Mountrakis et al., in press). Sm: Uppertectonic unit of Samos consisting of schists, sericite phyllites, quartzites and intercalations of marbles; Mi: Lower Neogene Group, including the Basal Fm, MavradzeiFm and Hora Fm; Lmi: Mytilinii Fm; Pl: Kokkarion Fm; sc1-sc2: scree deposits; al: alluvial deposits. Fossiliferous sites: 1. MLN – Q2 – Stefano; 2. MTN – ?Q6; 3.MYT– Q3 – ?Potamies; 4. MTL – 1 (A-D) – Q1 – Adriano; 5. Q4; 6. Q5; 7. Qx.

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the biostratigraphic significance and biochronologic use of theSamos fauna. Exceptions are: Forsyth Major’s collection inLondon and Lausanne that has some locality indications,referred to the local names of the areas where the fossils werefound (e.g., Adriano, Stefano, Potamies), and Barnum Brown’scollection of the American Museum of Natural History(AMNH) in which fossils are labeled as from seven distinctfossil sites or quarries (Qx, Q1-6) correlated in retrospect withthe local stratigraphy (Brown, 1924; Solounias, 1981;Kostopoulos et al., 2003). Yet, these correlations are sometimesdebated (Kostopoulos et al., 2009).

The puzzling taxonomy and time resolution of the Samosmammal faunas and the consequent doubts on their biochro-nological and palaeoecological signal was the irritant to a teamof palaeontologists from the Laboratory of Geology andPalaeontology of the Aristotle University of Thessaloniki, ledby Prof. G.D. Koufos, to undertake a new field campaign inSamos on 1993 that is still continued. The detailed results of15 years of field work are given in a special volume of Beiträgezur Paläontologie (Koufos and Nagel, 2009). This articleprovides a synopsis of that work with some new data andanalysis.

2. Stratigraphy of the Mytilinii Basin

The fossiliferous sites of Samos are placed within theNeogene deposits of the Mytilinii Basin (Fig. 1), a tectonicdepression, geotectonically correlated with the extended grabensystem of Asia Minor (Kostopoulos et al., 2009). According tothese authors the updated lithostratigraphy of the Neogenedeposits of the Mytilinii Basin consists of five successiveformations (Fig. 2).

Basal formation: red-brownish sands, alternating withgravel lenses, and intercalations of conglomerates andsandstones that represent alluvial fan deposits during theoriginal development of the basin, overlying unconformablythe pre-Neogene basement.

Mavradzei formation: it covers normally the Basal Fm andconsists of white-grey bituminous fresh-water limestones withPlanorbis, Lymnaea and Bithynia, alternated with greenish-yellowish clays and mudstones, as well as with lenticularintercalations of lignitic clays which pollen spectra indicateswarm and humid conditions (Ioakim and Koufos, 2009). Theformation ends at the top by a basalt flow, ca. 2 m-thick and anoverlying beige-green volcano-clastic bed of lachar-type; the

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Fig. 2. Litho-, bio- and chrono-stratigraphy of the Neogene deposits of the Mytilinii Basin, Samos, Greece. Data compiled from Kostopoulos et al. (2009), Ioakimand Koufos (2009) and Koufos et al. (2009b).

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basalt flow is dated at 11.2 � 0.7 Ma and the lachar at10.8 � 0.4 Ma (Weidmann et al., 1984). These datings suggestan age near the middle/late Miocene boundary for theMavradzei Fm.

Hora formation: the base of the Hora Fm consists of thick-bedded, massive, white-grey lacustrine limestones, marly-limestones of sub-littoral shallow water with intercalations ofgreenish tuffaceous clays of various thicknesses. The lime-stones are gradually thin-bedded, and in the upper parts of theformation they are laminated, representing deeper lacustrinedepositional environment, confirmed by microflora (Ioakimand Koufos, 2009; Kostopoulos et al., 2009).

Mytilinii formation: it consists of fluviolacustrine andvolcanoclastic deposits ending by reddish tuffaceous silts withmassive tuffs. Its stratigraphic position above the Hora Fm isdebated but our field observations suggest a continuoussuccession (Kostopoulos et al., 2009). The Mytilinii Fmcorresponds to a temporary desiccation of the Mytilinii Basinunder dry/warm temperate climatic conditions.

Kokkarion formation: shallow lacustrine thick-bedded tomassive, travertine-like limestones with Brotia cf. graecaalternate with green-brown clays and tuffaceous sands,outcropping mainly in the northern hilly ridge of the basin.

3. Vertebrate localities

Several fossiliferous localities with mainly mammals areknown from the Mytilinii Basin under various names(Appendix A). All of them are placed north to the Mytiliniivillage except Brown’s Q6 that is situated far to the north ofthe Mytilinii Basin, near the village of Kokkarion (Fig. 1).The fossiliferous sites we excavated are named with newnames in order to avoid confusion; their stratigraphic positionis given in Fig. 2. In the richest known fossiliferous area ofAdrianos ravine, we traced five fossil sites (pockets). Thelocality is named Mytilinii-1 (MTL) and the fossiliferoussites are characterized with the letters A, B, C, D, E. Twoadditional localities have been recognized in the Potamiesravine, named Mytilinii-3 (MYT) and Mytilinii-4 (MLN).The locality Mytilinii-5 (MTN) is located in the northernborders of the basin, near Kokkarion village (Fig. 1). The newcollection comes mainly from localities MLN, MYT andMTL.

One of the main goals of the Samos project was to correlatethe old fossiliferous sites with the new ones. Using allbibliographic and historical information, as well as our ownfield observations and updated taxonomic evidence, we propose

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Fig. 3. Taxomomic composition of the mammal samples from various Samos localities. NDS = Number of Determined Specimens.

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the correlation presented in Fig. 2 (Kostopoulos et al., 2003,2009). The locality Mytilinii-1 (MTL) and its sites (MTLA, B,C, D, E) can be correlated to the ‘‘Adriano’’ label of Forsyth-Major and to Brown’s Q1; the latter can be directly correlated tothe locality MTLD. The locality MYT can be correlated toBrown’s Q3 and possibly to the label ‘‘Potamies’’ of Forsyth-Major. The locality MLN can be correlated to Forsyth-Major’s‘‘Stefano’’ label and directly to Brown’s Q2. The MTN ispossibly identical to Brown’s Q6. The label ‘‘Vryssoula’’ ofForsyth-Major could be correlated to Brown’s Qx. Thestratigraphic correlation between old and new fossil localitiesof the Mytilinii Basin is detailed in Kostopoulos et al., 2009).

4. Mammal fauna

According to Solounias (1981) more than 30,000 specimenshave been collected from Samos between 1850 and 1950.Approximately 12,000 specimens are certainly registered inseveral museums and institutes but the sample of fullydescribed specimens does not exceed 1500. Furthermore, thenumber of mammalian taxa, their time distribution and theirage remain open to discussion. The new Samos collection isquite rich, including ca. 1200 determined specimens of severalmammalian groups (Fig. 3; Appendix B). The richest samplesare those from MTLA and MTLB, including respectively eight

Table 1Faunal list of the various new Samos mammal localities.

Species MLN MYT MTLA MTLB MTLC

Pseudomeriones pythagorasi + +‘‘Karnimata’’ provocator +Spermophilinus bredai cf. cf.Pliospalax sotirisi cf.Adcrocuta eximia +Hyaenictitherium wongii cf. + + cf.Machairodus giganteus +Metailurus parvulus +Parataxidea maraghana +Plioviverrops orbignyi +Protictitherium crassum +Choerolophodon pentelici +Zygolophodon turicensis +Ancylotherium pentelicum + + +‘‘Diceros’’ neumayri + + + +Dihoplus pikermiensis + +Hipparion brachypus + +Hipparion dietrichi + +Hipparion matthwei cf. cf. cf.Hipparion proboscideum aff. cf. cf. cf.Hipparion prostylum aff. +Hipparion forstenae cf. cf. cf.Pliohyrax graecus +Orycteropus gaudryi + +Microstonyx major +Helladotherium duvernoyi +Palaeotragus sp. sp.Palaeotragus rouenii + + +Samotherium boissieri +Samotherium major + + + +Gazella capricornis cf. cf. cf.Gazella mytilinii + +Gazella pilgrimi + + + +Miotragocerus vallencienesi + + +Miotragocerus sp.Majoreas sp.Palaeoryx majori + +Palaeoryx pallasi + +Palaeoryx sp.?Palaeoryx sp.Sporadotragus parvidens + +Tragoportax rugosifrons +Tragoportax sp.Skoufotragus zemalisorum +Skoufotragus laticeps + +Urmiatherium rugosifrons +

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and seven mammal groups of family to class level (Fig. 3(c,d)).Although the MYT sample is quite rich, its composition is thepoorest, including only four families (Fig. 3(b)). The equidspredominate in the MYT, MTLA and MTLB samples being themain group of the collection. Likely, the giraffids are abundantin MLN, MTLA and MTLC samples (Fig. 3(a,c,e)). The bovidsare also abundant and their frequency is almost constant in allsamples. The suids and proboscideans are relatively rare,represented by few remains in MTLA and MTLB, respectively(Fig. 3(c,d)). The carnivore remains are also uncommon, whilethe rhinocerotids are quite familiar. Aardwarks are quitecommon in the MTLA and MTLB samples, which alsoprovided some limited but important micromammalianremains. Besides mammals the Samos fauna also includessome reptiles: a small turtle has been found in the locality MLNand a skull of a Varanidae in MYT. The taxonomic compositionof the new fossil sites is given in Table 1 (see also Appendix B).As a total, 42 mammalian species have been recorded in thenew collection. Taxonomic novelties are the presence of thecarnivore genus Protictitherium that was traced for first time inSamos and the establishment of the new genus Skoufotragus,partly based on Pachytragus Schlosser, 1904 with a new speciesSkoufotragus zemalisorum.

4.1. Micromammals

Four out of the seven micromammalian taxa reported fromSamos by Black et al. (1980) are recognized in the newcollection (Vasileiadou and Sylvestrou, 2009). The question-able presence of Spermophilinus cf. bredai is confirmed byadditional material from MTLA and MTLB, whereas thehypodigm of the gerbilline Pseudomeriones pythagorasi Blacket al., 1980 is doubled in size, including dental specimenspreviously unknown. The poor original murine sample (Blacket al., 1980) is significantly enriched, confirming the initialidentification as ‘‘Karnimata’’ provocator and the presence ofthis species in MN12 of the Eastern Mediterranean. Finally, theoccurrence of a partial skull of Pliospalax cf. sotirisi furthersupports Black et al. (1980) opinion for the presence ofspalacids with simple dental morphology in Eastern Medi-terranean during pre-Pliocene times.

4.2. Carnivora

Fifteen carnivore species have been recorded in Samos fauna(Bernor et al., 1996b); among them Parataxidea maraghana,Plioviverrops orbignyi, Hyaenictitherium wongii, Adcrocutaeximia, Metailurus parvulus and Machairodus giganteus arepresent in the new collection. Although the hyaenids arecommon (Solounias, 1981), the small hyaenids (exceptPlioviverrops) are absent. A mandible discovered in MLN(Fig. 4) is characterized by its large size, relatively long andnarrow premolars and high cuspids in the carnassial. All thesecharacters are present in the holotype of Protictitheriumcrassum (Depéret, 1982) and in various samples of P. crassumfrom Greece and Eurasia (Koufos, 2009c). The genusProtictitherium is traced for first time in Samos mammal fauna.

4.3. Proboscidea

Among the collected material from Samos, the probosci-deans are relatively scarce. Solounias (1981) and Bernor et al.(1996b) reported Mammut borsoni (= Zygolophodon turicen-sis), Tetralophodon longirostris, Choerolophodon pentelici andDeinotherium cf. giganteum. The new proboscidean remains(Fig. 3) permit a full review of the Samos proboscideanassociation. Choerolophodon pentelici is described fromMTLB and Zygolophodon turicensis from MTLA (Konidarisand Koufos, 2009).

4.4. Rhinocerotidae

None of the three Chilotherium species reported by previousauthors in the old Samos collections has been recognized in thenew material. ‘‘Diceros’’ neumayri appears to be the mostabundant rhino in the new collection, recognized throughout theSamos sequence (Giaourtsakis, 2009). The Samos materialpreviously allocated to ‘‘Dicerorhinus’’ schleiermacheri is nowconsidered as belonging to Dihoplus pikermiensis, a lessabundant species in Samos, recognized mainly in the upperfossiliferous horizons (Giaourtsakis, 2009).

4.5. Equidae

The first detailed study of the Samos hipparionine horseswas given by Wehrli (1941) and Sondaar (1971). The latterauthor recognized four hipparion species: H. dietrichi, H.proboscideum, H. matthewi and Hipparion sp. Later, Forstén(1980, 1999) recognized one more species, H. brachypus, whileBernor et al. (1996b) added some more hipparion taxa to the

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Fig. 4. Protictitherium crassum, Mytilinii-4, MLN, Samos, middle Turolian,MN12. a, b. Left and right mandibular corpus, MLN-42, external view; c. Righttooth row, MLN-42, occlusal view.

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previous list. According to Bernor et al. (1996b) H. prostylum,H. gettyi and H. mediterraneum are present in Qx and Q6,whereas H. giganteum and H. nikosi are recognized in Q2(Fig. 2). The study of the new hipparion collection from Samosand their comparisons with the old material (Vlachou andKoufos, 2009) allow the recognition of eight to nine species,and permit an entire revision of the hipparionine associationprovided by Bernor et al. (1996b) (Fig. 5). H. prostylum hasbeen recognized in Q6 and probably in MLN and MYT.Although H. prostylum resembles H. dietrichi, it differs from itin having slightly smaller size, different morphology of thepost-orbital fossa (POF) and reduced metapodial length. Themorphometric characters of H. matthewi documented in Q1/MTLA-B and H. nikosi recognized in Q5 are clearlydistinguished. H. nikosi mainly differs from H. matthewi inhaving deeper nasal notch (placed above P3), longer muzzle,and moderately deep, subtriangular shaped POF that lacksanterior rim. Both species share the same skeleton pattern withelongated and slender bones. Furthermore, H. mediterraneumof Bernor et al. (1996b) and H. cf. mediterraneum of Forstén(1980) are synonymized with H. cf. forstenae known from theQ1/MTLA-B and probably from MYT and Q5. It differs fromH. prostylum in having different size and completely differentfacial morphology. H. cf. forstenae is smaller than H.mediterraneum and its facial morphology, alike to H. dietrichi,

has a strongly reduced POF, anteroventrally oriented, notposteriorly pocketed and situated far from the orbit. Thecomplete list of the hipparionine horses found in the Samoslocalities is given in Table 1, while their extensive descriptionand comparison is given by Vlachou and Koufos (2009).

4.6. Suidae

The suids are known from Samos by two species:Propotamochoerus sp. from an unknown stratigraphic level,and Microstonyx major. The latter taxon is recognized in all oldand new collections by several cranial and dental remains(Sylvestrou and Kostopoulos, 2009).

4.7. Giraffidae

Giraffids is one of the best represented mammal groups ofSamos, known by a considerable amount of material hold in theold and new collections. The study of the entire sample(Kostopoulos, 2009a) allows the recognition of Palaeotragusrouenii, Palaeotragus quadricornis, Samotherium boissieri,Samotherium major, and Helladotherium duvernoyi; thepresence of Bohlinia remains questionable but certainly ofminor importance. Among these taxa, Samotherium is veryimportant both because of its abundance, and biostratigraphic

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Fig. 5. Stratigraphic distribution of the Samos hipparionine horses. 1. H. prostylum. 2, 3. H. dietrichi. 4. H. proboscideum. 5, 6. H. brachypus. 7. H. cf. matthewi. 8. H.nikosi. 9, 10. H. cf. forstenae (data from Vlachou & Koufos, 2009).

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utility as a marker between the lower and upper fossiliferoushorizons of Samos. The postcranial elements of Samotheriumshow a gradual increase in time with the replacement ofSamotherium boissieri by Samotherium major around 7.4 Ma.Interestingly, Samotherium major and Helladotherium duver-noyi, two large giraffids of similar size, co-occur in the upperfossiliferous levels of Samos, implying that they had differentpalaeoecological profiles.

4.8. Bovidae

The comparison between the new and old bovid materialpermits a brave review of the Samos bovid’s association thatincludes 27 validated species (Kostopoulos, 2009b); 13 of themare recognized in the new material (Table 1). The review of theSamos bovids includes the recognition of four Gazella species,the formal attribution of Parurmiatherium rugosifrons toUrmiatherium, the further distinction of Palaeoryx pallasi fromPalaeoryx majori, the synonymy of Tragoportax curvicornisAndree, 1926 with Tragoportax punjabicus (Pilgrim, 1910) andthe establishment of a new generic name, Skoufotragusincorporating Pachytragus laticeps Andree, 1926, part of

Pachytragus crassicornis Andree, 1926 and the new speciesSkoufotragus zemalisorum (Fig. 6). The stratigraphic distribu-tion of the bovid taxa suggests four successive bovidassemblages through time.

4.9. Less common groups

The tubulidentates are known from Samos by the speciesOrycteropus gaudryi, recorded in MTLA and MTLB (Koufos,2009c). The pliohyraxes are rather rare in the Samos fauna,although two species are referred in the old collections:Pliohyrax kruppii and P. graecus; the latter was found in MTLC(Koufos, 2009d). The schizotheriine chalicothere Ancylother-ium pentelicum is the sole representative of this family inSamos, rarely documented in the old and new collections(Giaourtsakis and Koufos, 2009).

5. Chronology

The age of the Samos mammal fauna has been frequentlychallenged according to different sources of evidence(magnetostratigraphy, radiochronology, biochronology). The

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Fig. 6. Stratigraphic distribution of the most characteristic artiodactyls (Giraffidae and Bovidae) of Samos. 1. Samotherium boissieri, NHML M4215. 2.Samotherium major, female, NHMA MTLA540. 3. Samotherium major, male, AMNH22786. 4. Criotherium argalioides, NHML M4199. 5. Urmiatheriumrugosifrons, NHMW A571. 6. Tragoportax punjabicus, AMNH20566. 7. Skoufotragus zemalisorum, NHMA MYT1. 8. Skoufotragus laticeps, variety ‘‘long-braincase’’, PIM6. 9. Skoufotragus laticeps, variety ‘‘short-braincase’’, NHMA MTLB164. 10. Skoufotragus schlosseri, AMNH20567 (data from Kostopoulos,2009a,b).

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problem is certainly associated with the limited correlation ofthe old collections with the referred localities and theirstratigraphic succession within the Mytilinii Basin section. Theradiochronology of the volcanic tuffs and volcanoclasticsediments of the Mytilinii Fm befuddle more the problem(Koufos et al., 2009b and ref. herein). Summarizing previousoptions, the bulk of the Samos fauna appears either as a singleunit of Turolian age (Solounias, 1981; Weidmann et al., 1984;Bernor et al., 1996b), or as two successive units, an older oneincluding Brown’s Q1-4 and a younger one including Q5(Gentry, 1971; Sondaar, 1971; Heissig, 1975).

The composite magnetostratigraphic section of the MytiliniiBasin and its correlation with the GPTS suggests that theMytilinii Fm deposited between the Chrons C4n/C4r and C3Ar,corresponding to the time interval from �8.1 to �6.7 Ma(Kostopoulos et al., 2003). The study of the new mammalcollection, its comparison with the old material and the newmagnetostratigraphy of the Mytilinii Fm allowed a bettercorrelation of the old quarries with the local stratigraphy andthe new sites (Fig. 2). The new studies provided an improvedage determination for the fossiliferous deposits and thedistinction of different faunal associations through time(Koufos and Nagel, 2009). Combining all these data, fourmain fossiliferous horizons can be recognized in the MytiliniiFm (Koufos et al., 2009b).

5.1. Primary Mammal Assemblage of Samos (PMAS)

It incorporates data from the oldest known localities Qx andVryssoula, as well as from the old localities Stefano and Q2,and the new locality MLN. This part of the Mytilinii Fm can becorrelated to Chrons C4n.2n-C3Br.2n, corresponding to thetime interval from 8.0 to 7.4 Ma; an estimated age of�7.5 Ma issuggested for MLN. Important co-occurrences in PMASinclude Protictitherium crassum, Promephitis lartetii, Dicerosneumayri, Hipparion proboscideum, H. prostylum, Palaeotra-gus quadricornis, Samotherium boissieri, Gazella cf. ancyr-ensis, Criotherium argalioides and Tragoportax rugosifrons.Details on the age of the localities and the succession of themammalian faunas are given in Koufos et al. (2009b). The endof PMAS is apparently marked by the disappearance ofMajoreas woodwardi, Tragoreas oryxoides and Protoryxcapricornis. The MLN hipparions include H. prostylum, knownfrom early-middle Turolian (Bernor et al., 1996a) and H. aff.proboscideum, ranging from early to middle Turolian (Koufos,1987; Koufos and Kostopoulos, 1994; Sen et al., 1994). Thecombination of the giraffid Palaeotragus rouenii with the morerobust Palaeotragus sp. rather indicates an early Turolian age(Kostopoulos, 2009a). Although Samotherium boissieri isoriginally known from Samos, it is also known from the Turkishlocality Kemiklitepe-D, dated at�7.7 Ma (Sen et al., 1994). Thebovid Gazella pilgrimi is similar to the early Turolian forms fromContinental Greece and Palaeoryx is a common element in themiddle Turolian faunas of the Greek mainland (Kostopoulos,2009b). Thus, the coexistence of early and middle Turolianelements suggests a latest early Turolian age for the PMAS,fitting well with the magnetostratigraphic data (Fig. 2).

5.2. Intermediate Mammal Assemblage of Samos (IMAS)

It incorporates data from the old fossiliferous sites Q6 andQ3, as well as from the new MTN and MYT ones (Fig. 2). Themagnetostratigraphy of this part of the section suggests acorrelation with Chron C3Br.2r, ranging from�7.4 to�7.2 Maand indicating an estimated age of �7.3 Ma for MYT(Kostopoulos et al., 2003; Koufos et al., 2009b). Althoughseveral taxa from the previous assemblage continue to exist,IMAS, is characterized by a renewal of the fauna with thereplacement of Samotherium boissieri by S. major and the localoccurrence of Dihoplus pikermiensis, Ancylotherium penteli-cum, Hipparion cf. matthewi, H. cf. forstenae, Skoufotraguszemalisorum and several micromammals. Hipparion prostylumand Criotherium argalioides make their last appearance.Although the available material from MTN is scarce, thereplacement of Samotherium boissieri by S. major indicates anage younger than 7.4 Ma (Kostopoulos, 2009a). The newlocality MYT is identical to Q3 of Brown and probably to thePotamies of Forsyth Major (Fig. 2). The MYT material wasstudied extensively and provided useful biochronologicalinformation. The rhinocerotid assemblage of MYT (‘‘Diceros’’neumayri and Dihoplus pikermiensis) together with Ancy-lotherium pentelicum suggests a middle Turolian age(Giaourtsakis, 2009; Giaourtsakis and Koufos, 2009). Thebovids Palaeoryx and Sporadotragus parvidens are knownfrom the middle Turolian (MN12) in the Greek mainland(Kostopoulos, 2009b), even thought the latter species is alsodocumented in Kemiklitepe-D, dated at �7.7 My (Sen et al.,1994). The presence of Skoufotragus zemalisorum in MYTindicates an age older than MTL (Kostopoulos, 2009b). Themajority of the MYT fauna consists of middle Turolian (MN12)elements and the presence of some early Turolian ones suggestsa lower middle Turolian age, in agreement with themagnetostratigraphy (Fig. 2).

5.3. Dominant Mammal Assemblage of Samos (DMAS)

It incorporates data from the old localities Q4, Q1, Adrianoor Adrianos or Stefanidis ravine, as well as from the newlocality MTL (fossiliferous sites MTLA, MTLB and MTLC).The DMAS part of the Mytilinii Fm can be magnetostrati-graphically correlated to Chrons C3Br.1n-C3Bn, correspondingto the time interval from �7.2 to �6.9 Ma; an estimated age of�7.1 Ma is quite possible for the MTL fauna. The DMAS is anadvanced stage of IMAS; it is marked by the appearance ofHipparion brachypus, H. dietrichi and probably Chilotheriumschlosseri, and the replacement of Skoufotragus zemalisorumby S. laticeps. The species Metailurus parvulus, Helladother-ium duvernoyi, Gazella cf. capricornis, Urmiatherium rugosi-frons and Palaeoryx majori also occur. The majority of IMASelements continue in DMAS but usually with larger morphs.Promephitis larteti, Pliohyrax graecus, Hipparion probosci-deum, H. brachypus, Tragoportax rugosifrons and Palaeoryxpallasi make their last appearance in the Samos faunalsuccession. The rich MTL fauna (Table 1) provides otherarguments for the chronology of this locality: the rodents

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Pseudomeriones pythagorasi and ‘‘Karnimata’’ provocatorindicate a middle Turolian age (Vasileiadou and Sylvestrou,2009). The carnivores Metailurus parvulus and Machairodusgiganteus are similar to those from the Continental Greeklocalities of Pikermi, Chomateres, Vathylakkos, Halmyropo-tamos and Kerassia; all these localities are dated to middleTurolian (Koufos, 2009b). The rhinocerotids ‘‘Diceros’’neumayri, Dihoplus pikermiensis together with Ancylotheriumpentelicum suggest also a middle Turolian age (Giaourtsakis,

2009; Giaourtsakis and Koufos, 2009). The MTL hyracoidPliohyrax graecus resembles that from Pikermi and Halmyr-opotamos, suggesting also middle Turolian age (Koufos,2009d). The co-occurrence of Hipparion proboscideum andH. forstenae indicates an early-middle Turolian age; theaddition of H. brachypus, known from Pikermi (MN12), theBulgarian locality of Hadjidimovo (MN12) and the Turkishlocality of Akkasdagi (MN12; 7.0 � 0.1 Ma) gives a youngerfeature to the MTL fauna (Vlachou and Koufos, 2009).

[()TD$FIG]

Fig. 7. Faunal composition of the Samos mammal assemblages using taxonomy and Minimum Number of Individuals (MNI).

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The bovids Palaeoryx pallasi and Gazella capricornis are alsomiddle Turolian elements; moreover the occurrence of the moreevolved Skoufotragus laticeps suggests an age younger than�7.3 Ma (Kostopoulos, 2009b). Thus, DMAS seems tocorrespond better to the upper part of middle Turolian(MN12), fitting well with the magnetostratigraphy (Fig. 2).

5.4. Final Mammal Assemblage of Samos (FMAS)

It includes the old Brown’s Q5 (Gentry, 1971; Sondaar,1971; Vlachou and Koufos, 2009) and possibly locality L(Fig. 2). Although Q5 is relocated, our efforts to find fossilswere unsuccessful untill now – except some sporadic bones,possibly remains of the old excavations – we could not find thefossiliferous horizon. According to our magnetostratigraphicdata this part of the section can be correlated to Chrons C3Bn-C3Ar, corresponding to the time-span from �7.1 to �6.7 Ma.The age of the locality Q5 is estimated as being between 6.9 and6.7 Ma (Kostopoulos et al., 2003; Koufos et al., 2009b). TheFMAS is characterized by a decrease in the number of taxa inthe fauna. Hystrix primigenia, Hipparion nikosi and Trago-portax punjabicus appear for the first time, the more evolvedSkoufotragus schlosseri replaces the earlier S. laticeps, andOrycteropus gaudryi appears with a larger morph.

6. Palaeoecology

Several scientists dealt with the determination of the Samospalaeoenvironment based on the whole mammal assemblage oron part of it (Solounias and Dawson-Saunders, 1988; Boniset al., 1992a, 1999, 2004; Solounias et al., 1999; Strömberget al., 2007). In most of these analyses a key problem concernsthe lack of adequate time resolution due to mixing of materialfrom different horizons, representing different mammalassociations and potentially different environments. The newwell-dated collection with reliable correlation to the localstratigraphy permitted a better analysis of the fauna andprovided more precise results for the palaeoenvironment ofSamos during late Miocene. This includes palynological dataand comparison of the Samos mammal faunas with recent andfossil ones from known environments, the study of itscharacters (diversity, composition, guild structure) and thestudy of the dental meso- and micro-wear (Koufos et al.,2009a).

The pollen spectra of the upper part of Mytilinii Fm ischaracterized by the predominance of Caligonum, Lygeum,Olea, Cistus and Quercus ilex-type (Fig. 2), as well as of openvegetation plants and steppe elements (Ioakim and Koufos,2009). Such a wealthy herbaceous vegetal layer indicative of

[()TD$FIG]

Fig. 8. Correspondence Factor Analysis of late Miocene Eurasian mammal assemblages in comparison with modern ones from known environments. (see AppendixA for abbreviations).

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dry/warm-temperate climatic conditions is consistent with theanalysis of the late Miocene phytolith assemblages of theSamos fossiliferous sites and those from neighboring areas(Strömberg et al., 2007).

The faunal composition of the three basic new Samosmammal assemblages (MLN, MYT, MTL), using both thenumber of species per family or group category and theMinimum Number of Individuals (MNI) per family or groupcategory shows a clear predominance of bovids, giraffids andequids (Fig. 7); for more details see Koufos et al. (2009a).These families represent more than 75% of the number of taxa(72% of 21 individuals) in MLN, 79% (89%) in MYT and 57%(73%) in MTL (Fig. 7). In spite of the limited data for the poorMLN and MYT assemblages, an increasing trend of the bovids,equids and rhinos from the older MLN to the younger MTLassemblage is evident. The rhinos and chalicotheres showrelatively high frequency (> 10%), even with a low number ofoccurring taxa. The carnivores are quite diversified in MTL(18% of the taxa) but their number of individuals is low (9%).The other families or groups are represented by low frequenciesin both cases.

The Samos mammal assemblages are compared to a set ofEurasian fossil assemblages of Vallesian and Turolian age and aset of modern assemblages from known habitats including bothclosed and open faunas by Correspondance Factor Analysis(CFA), using the PAST software (Hammer et al., 2001) (Fig. 8).The CFA of the modern assemblages alone indicates cleardistinction between the closed and the open faunas (upper rightinset of Fig. 8). When fossil faunas are included, they form aseparate group from the modern ones along the axis-1. Alike themodern faunas, the late Miocene assemblages are split in twogroups along the axis-2: those of late Vallesian age from Westernand Central Europe match together and can be correlated to themodern closed ones; all late Miocene assemblages of EasternMediterranean form a group which can be generally correlated tothe modern open assemblages. The three mammal assemblagesof Samos are included into this latter group, suggestingchronogeographical and ecological relationships.

A detailed comparison of the taxonomic spectra (% numberof species per family or class) of the Samos assemblages withthose extracted from several types of modern environments(Fig. 9) cannot be direct, since the absence of primates in Samos

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Fig. 9. Taxonomic faunal composition (number of taxa per family or group category) of some modern mammal assemblages from known environments and MLN,MYT and MTL fossil mammal assemblages from Samos. Data for Umfolozi taken from Mentis (1970), for Birao from Felix (1953), for Lamto from Bourliere et al.(1974), for Transwall 1-9 from Rautenbach (1978), for Irangi from Rahim (1966), for Wilpatu from Eisenberg and Lockhart (1972), for Hluhluwe from Bourquin et al.(1971), for Gunong from Medway (1972) and for Serengeti from www.invista.com/science/ecology/natpark/serenget.htm.

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distinguishes it from almost all recent faunas in comparison.The absence of tapirids, cervids, tragulids, as well as theabundance of equids, bovids and giraffids separate clearly theSamos assemblages from the typical closed ones (Wilpattu,Gunong, Irangi, Hluhluwe) and suggest stronger resemblancewith the open ones, especially with Birao assemblage,representing a bushed-savannah and Transvaal 4-9, represent-ing a mixed-wooded savannah (Fig. 9).

The guild structure of the Samos carnivores based on thetaxa from the new collection, as well as for the wholecarnivoran assemblage (Nagel and Koufos, 2009) indicates theabsence of semifossorial, arboreal and semi-aquatic elements.Equally, small taxa (<3 kg) are missing, whereas the large-sized carnivores (>100 kg) are present (Indarctos, Machair-odus). The dietary characters of the Samos carnivoranassemblage includes insectivorous and hypercarnivorousspecies but also a bone/meat feeder (Adrocruta eximia; Nageland Koufos, 2009: fig. 2). The larger taxa, as well as the rarecursorial forms suggest an open habitat; on the other hand thegeneralists indicate the presence of more closed conditions.

The dental meso- and micro-wear of the Samos mammalassemblages, studied by G. Merceron (Koufos et al., 2009a)

indicates that the Samos herbivores match with the extantgrazers except for Skoufotragus zemalisorum from MYT(Koufos et al., 2009a: fig. 1). The combination of both meso-and micro-wear data suggests the predominance of inter-mediate feeders related to bushy/grassy vegetation. Indeedintermediate feeders represent 75% of the 21 individuals (80%of the taxa) in MLN, 41% of 27 individuals (39% of the taxa) inMYT, and 60% of 167 individuals (45% of the taxa) in MTL(Fig. 10). Grazers are also relatively abundant with 28% of 21individuals (20% of the taxa) in MLN, 33% of 27 individuals(31% of the taxa) in MYT, and 24% of 167 individuals (26% ofthe taxa in MTL.

Along the lineage Skoufotragus zemalisorum (MYT) –

Skoufotragus laticeps (MTL) – to Skoufotragus schlosseri (Q5)there is a shift towards more grazing feeding habits, as it isshown both by the cranial features (Kostopoulos, 2009b) andthe mesowear pattern (Koufos et al., 2009a). This hint isstrengthened by the relative abundance of rhinos: although inthe MYT assemblage ‘‘Diceros’’ neumayri, an open and drydweller and Dihoplus pikermiensis, a more closed andtemperated species are equally present, the former clearlypredominates in the MTL assemblage (Giaourtsakis, 2009).

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Fig. 10. Feeding preferences of the Samos mammal assemblages using taxonomy and Minimum Number of Individuals (MNI).

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Combining all available data, an open bush-land environ-ment with thick grassy-floor of C3 graminoids is proposed forthe late Miocene of Samos. An increase of the open and drycharacter of the environment is quite possible from MYT (�7.3Ma) to MTL (�7.1 Ma).

7. Biogeographic relationships

The biogeographic relationships of the Samos mammalassemblages were discussed among palaeontologists andvarious aspects have been formulated (Bonis et al., 1992a,2004; Bernor et al., 1996b; Solounias et al., 1999; Koufos et al.,2006). The two classical Turolian mammal faunas of Pikermiand Samos, although they are neighboring, differ significantlyfrom a palaeoecological point of view, implying a subdivisionof their common geographic domain [the so-called Greco-Iranian Province (Bonis et al., 1992b)] in smaller ecologicalclusters (Kostopoulos, 2009c). These data are further supportedby Koufos et al. (2009a: figs. 6-8) who showed that comparisonof the taxonomic composition of a set of Turolian assemblagesfrom the wider Greco-Iranian Province (by means of bothnumber of species and MNI per family or group category)allows the recognition of two main groups: a group of Turolianassemblages from the Greek mainland (PIK, VATH, PER,RZO, PXM, DYTI) and another one incorporating the Samos(MLN, MYT, MTL, Q5) and Anatolian faunas plus Maraghehin Iran (AKK, KTA, KTD, LMRG, MMRG, UMRG) (seeAppendix A). This grouping of the Samos Turolian mammalassemblages indicates that they were more similar to the WestAsian than to the Balkan faunas. According to the arrangementof the various mammal assemblages and the palaeoecologicalcharacter of each mammal group, the authors conclude thatthere is a Turolian environmental gradient across EasternMediterranean with more open and arid conditions towards theEast (Koufos et al., 2009a).

8. Conclusions

New field researches on Samos Island allowed a significantcollection of stratigraphically controlled fossil samples andimproved our knowledge on taxonomy, stratigraphy, chronol-ogy, palaeoecology and palaeobiogeography of the lateMiocene mammal faunas. The Mytilinii Fm is dated to theTurolian, covering a time-interval from the upper part of earlyTurolian (MN11) to the lower part of late Turolian (MN13). Thecombination of the biochronological and magnetostratigraphicdata suggests an estimated age of �7.5 Ma for MLN, �7.3 Mafor MYT and �7.1 Ma for MTL.

The study of the fauna and its evolutionary relationshipsindicates four successive mammal associations that cover themiddle part of the Turolian in Eastern Mediterranean. Thepalaeoenvironmental analysis of the Samos fauna suggests anopen and dry bushy environment with a thick grass-floor,similar to a modern bushed-savannah and possibly a gradualstrengthening of the open/arid character during the middleTurolian. The Samos mammal faunas are biogeographically

closer to those of Asia Minor and different from those ofcontemporaneous faunas from Continental Greece.

Acknowledgements

We are grateful to Dr J. Agustí and L. Rook for inviting us toparticipate to this volume in honour of Dr P. Mein. Many thanksare due to Dr R.L. Bernor and Dr Sevket Sen for their usefulcomments on the manuscripts. Numerous persons helped usduring our field work in Samos; we thank all very much. Thefield campaigns in Samos were financially supported by thePrefecture of Samos, the ‘‘Konstantinos and Maria ZimalisFoundation’’ and the former Community of Mytilinii.

Appendices A-B. Supplementary data

Supplementary data associated with this article can befound, in the online version, at doi:10.1016/j.geobios.2010.08.004.

References

Bernor, R.L., Koufos, G.D., Woodburne, M., Fortelius, M., 1996a. The evolu-tionary history and biochronology of European and southeastern Asian lateMiocene and Pliocene hipparionine horses. In: Bernor, R.L., Fahlbusch, V.,Mittmann, H.W. (Eds.), The Evolution of Western Eurasian NeogeneMammal Faunas. Columbia University Press, New York, pp. 7–46.

Bernor, L.R., Solounias, N., Swiser, C.C., Van Couvering, A.J., 1996b. Thecorrelation of the classical ‘‘Pikermian’’ mammal faunas-Maragheh, Samosand Pikermi, with the European MN unit system. In: Bernor, R., Fahlbusch,V., Mittmann, H.W. (Eds.), 1. The evolution of the Western EurasianNeogene Mammal Faunas. Columbia University Press, New York, pp.137–154.

Black, C.C., Krishtalka, L., Solounias, N., 1980. Mammalian fossils of Samosand Pikermi. Part 1. The Turolian rodents and insectivores of Samos. Annalsof Carnegie Museum 49, 359–378.

Bonis, L., de Bouvrain, G., Koufos, G.D., 1999. Palaeoenvironments of thehominoid primate Ouranopithecus in the late Miocene deposits of Mace-donia, Greece. In: Agustí, L., Rook, L., Andrews, P. (Eds.), HominoidEvolution and climatic change in Europe. 1. The evolution of the Neogeneterrestrial ecosystems in Europe. Cambridge University Press, London, pp.205–237.

Bonis, L., de Bouvrain, G., Koufos, G.D., 2004. Evolution of late Miocenefaunas and palaeoenvironment in the northern and eastern Mediterraneanrealms. Proceedings 5th International Symposium on Eastern Mediterra-nean Geology, Thessaloniki, Greece, 14-20 April 2004, Abstracts 447–450.

Bonis, L., de Bouvrain, G., Geraads, D., Koufos, G.D., 1992a. Diversity andpalaeoecology of Greek late Miocene mammalian faunas. Palaeogeography,Palaeoclimatology, Palaeoecology 91, 99–121.

Bonis, L., de Brunet, M., Heintz, E., Sen, S., 1992b. La province Greco-irano-afghane et la répartition des faunes mammaliennes au Miocène supérieur.Paleontologia i Evolucio 24/25, 103–112.

Brown, B., 1924. Samos-Romantic Island of the Aegean. Natural History 27,1178–1181.

Bourliere, F., Minner, E., Vattoux, R., 1974. Les grands mammifères de larégion de Lamto, Côte d’Ivoire. Mammalia 38, 433–447.

Bourquin, O., Vicent, J., Hitchins, P., 1971. The Vertebrate of the Hluhluwegame reserve-corridor (State-land)-Umfolozi game reserve complex. TheLammengeyer 14, 5–58.

Eisenberg, J., Lockhart, M., 1972. An ecological reconnaissance of WilpattuNational Park, Ceylan. Smithsonian Contributions to Zoology 101, 1–118.

Felix, A., 1953. Notes sur la faune de Birao. Mammalia 17, 55–66.Forstén, A.M., 1980. How many Hipparion species at Samos? Neues Jahrbuch

für Geologie und Paläontologie, Monatschefte 7, 391–396.

G.D. Koufos et al. / Geobios 44 (2011) 237–251250

Author's personal copy

Forstén, A.M., 1999. Snout proportions in some Eurasian hipparions (Mam-malia, Equidae): taxonomic and functional implications. Geodiversitas 21,255–278.

Forsyth Major, C.J., 1888. Sur un gisement d’ossements fossiles dans l’île deSamos contemporain de l’âge de Pikermi. Compte Rendus Hebdomadairesdes Séances de la Société géologique de France 107, 1178–1181.

Forsyth Major, C.J., 1894. Le gisement ossifère de Mytilinii et catalogued’ossements fossiles recueillis à Mitylini, île de Samos, et déposés auCollège Galliard à Lausanne. Éditions Georges Bridel and Cie, Lausanne,pp. 1–15.

Gentry, A.W., 1971. The earliest goats and other antelopes from SamosHipparion Fauna. Bulletin of the British Museum (Natural History),Geology 20, 229–296.

Giaourtsakis, I., 2009. Rhinocerotidae. In: Koufos, G.D., Nagel, D. (Eds.), TheLate Miocene Mammal Faunas of the Mytilinii Basin, Samos Island,Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 157–187.

Giaourtsakis, I., Koufos, G.D., 2009. Chalicotheriidae. In: Koufos, G.D., Nagel,D. (Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin,Samos Island, Greece: new Collection. Beiträge zur Paläontologie, 31. pp.189–205.

Hammer, C., Harper, D.A.T., Ryan, P.D., 2001. PAST: Paleontological StatisticsSoftware Package for Education and Data Analysis. Palaeontologia Elec-tronica 4, 1–9.

Heissig, K., 1975. Rhinocerotidae aus dem Jungtertiars Anatolien. Geolo-gisches Jahrbuch B 15, 145–151.

Ioakim, C., Koufos, G.D., 2009. Palynology. In: Koufos, G.D., Nagel, D.(Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 27–35.

Konidaris, G.E., Koufos, G.D., 2009. Proboscidea. In: Koufos, G.D., Nagel, D.(Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 139–155.

Kostopoulos, D.S., 2009a. Giraffidae. In: Koufos, G.D., Nagel, D. (Eds.), TheLate Miocene Mammal Faunas of the Mytilinii Basin, Samos Island,Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 299–343.

Kostopoulos, D.S., 2009b. Bovidae. In: Koufos, G.D., Nagel, D. (Eds.), TheLate Miocene Mammal Faunas of the Mytilinii Basin, Samos Island,Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 345–389.

Kostopoulos, D.S., 2009c. The Pikermian Event: temporal and spatial resolu-tion of the Turolian large mammal fauna in SE Europe. Palaeogeography,Palaeoclimatology, Palaeoecology 274, 82–95.

Kostopoulos, D.S., Koufos, G.D., Sylvestrou, I.A., Syrides, G.E., Tsompa-chidou, E., 2009. Lithostratigraphy and fossiliferous sites. In: Koufos,G.D., Nagel, D. (Eds.), The Late Miocene Mammal Faunas of theMytilinii Basin, Samos Island, Greece: new Collection. Beiträge zurPaläontologie, 31. pp. 13–26.

Kostopoulos, S.D., Sen, S., Koufos, D.G., 2003. Magnetostratigraphy andrevised chronology of the late Miocene mammal localities of Samos,Greece. International Journal of Earth Sciences 92, 779–794.

Koufos, G.D., 1987. Study of the Turolian hipparions of the lower Axios valley(Macedonia, Greece). 1. Locality ‘‘Ravin des Zouaves-5’’ (RZO). Geobios20, 293–312.

Koufos, G.D., 2009a. History of the Samos fossil mammals. In: Koufos,G.D., Nagel, D. (Eds.), The Late Miocene Mammal Faunas of theMytilinii Basin, Samos Island, Greece: new Collection. Beiträge zurPaläontologie, 31. pp. 1–12.

Koufos, G.D., 2009b. Carnivora. In: Koufos, G.D., Nagel, D. (Eds.), The LateMiocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece:new Collection. Beiträge zur Paläontologie, 31. pp. 57–105.

Koufos, G.D., 2009c. Tubulidentata. In: Koufos, G.D., Nagel, D. (Eds.), TheLate Miocene Mammal Faunas of the Mytilinii Basin, Samos Island,Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 107–125.

Koufos, G.D., 2009d. Hyracoidea. In: Koufos, G.D., Nagel, D. (Eds.), The LateMiocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece:new Collection. Beiträge zur Paläontologie, 31. pp. 127–137.

Koufos, G.D., Kostopoulos, D.S., 1994. The late Miocene mammal localities ofKemiklitepe (Turkey). 3. Equidae. Bulletin du Muséum national d’Histoirenaturelle (4) 16C, 41–80.

Koufos, G.D., Merceron, G., Kostopoulos, D.S., Vlachou, T.D., Sylvestrou, I.,2006. Palaeoecology and palaeobiogeography. In: Koufos, G.D. (Ed.), Thelate Miocene Vertebrate locality of Perivolaki, Thessaly, Greece. Palaeon-tographica Abt, A 276. pp. 201–221.

Koufos, G.D., Nagel, D., 2009. The Late Miocene Mammal Faunas of theMytilinii Basin, Samos Island, Greece: new Collection. Beiträge zurPaläontologie 31, 1–438.

Koufos, G.D., Kostopoulos, D.S., Merceron, G., 2009a. Palaeoecology-palaeo-biogeography. In: Koufos, G.D., Nagel, D. (Eds.), The Late MioceneMammal Faunas of the Mytilinii Basin, Samos Island, Greece: newCollection. Beiträge zur Paläontologie, 31. pp. 409–428.

Koufos, G.D., Kostopoulos, D.S., Vlachou, T.D., 2009b. Chronology. In:Koufos, G.D., Nagel, D. (Eds.), The Late Miocene Mammal Faunas ofthe Mytilinii Basin, Samos Island, Greece: new Collection. Beiträge zurPaläontologie, 31. pp. 397–408.

Medway, L., 1972. The distribution and altitudinal zonation of birds andmammals on Gunong Benom. Bulletin of the British Museum naturalHistory (Zoology) 23, 105–154.

Mentis, M.T., 1970. Estimates of natural biomasses of large herbivores in theUmfolozi game reserve area. Mammalia 34, 363–393.

Mountrakis, D., Kilias, A., Vavliakis, E., Psilovikos, A., Karakaisis, G.,Papazachos, C., Thomaidou, E., Seitanidis, G., in press. Neotectonicmap of Samos Island, scale 1:100.000. Earthquake Planning and ProtectionOrganization of Greece.

Nagel, D., Koufos, G.D., 2009. Carnivore guild structure. In: Koufos, G.D.,Nagel,D. (Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 391–396.

Rahim, U., 1966. Les mammifères de la forêt équatoriale de l’Est du Congo.Annals Museum Royal Africa 149.

Rautenbach, I.L., 1978. Ecological distribution of the mammals of the Trans-vaal. Annals Transvaal Museum 31, 131–156.

Sen, S., Bonis de, L., Dalfes, N., Geraads, D., Koufos, G.D., 1994. Lesgisements de mammifères du Miocène supérieur de Kemiklitepe, Turquie.1. Stratigraphie et magnétostratigraphie. Bulletin du Muséum nationald’Histoire naturelle (4) 16C, 5–17.

Solounias, N., 1981. The Turolian fauna from the island of Samos, Greece.Contributions to Vertebrate Evolution 6, 1–232.

Solounias, N., Dawson-Saunders, B., 1988. Dietary adaptations and paleoecol-ogy of the late Miocene ruminants from Pikermi and Samos in Greece.Palaeogeography, Palaeoclimatology, Palaeoecology 65, 149–172.

Solounias, N., Plavcan, J.M., Quade, J., Witmer, V., 1999. The paleoecology of thePikermian biome and the savanna myth. In: Agustí, L., Rook, L., Andrews, P.(Eds.), Hominoid Evolution and climatic change in Europe. 1. The evolutionof the Neogene terrestrial ecosystems in Europe. Cambridge University Press,London, pp. 427–444.

Sondaar, P.Y., 1971. The Samos Hipparion. Proceedings Koninklijke Neder-landse Akademie Wetenschappen B 74, 417–441.

Strömberg, C.A.E., Werdelin, L., Friis, E.M., Sarac, G., 2007. The spread ofgrass-dominated habitats in Turkey and surrounding areas during theCenozoic: phytolith evidence. Palaeogeography, Palaeoclimatology,Palaeoecology 250, 18–49.

Sylvestrou, I.A., Kostopoulos, D.S., 2009. Suidae. In: Koufos, G.D., Nagel, D.(Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 283–

297.Vasileiadou, K., Sylvestrou, I.A., 2009. Micromammals. In: Koufos, G.D.,Nagel,

D. (Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: new Collection. Beiträge zur Paläontologie, 31. pp. 37–55.

Vlachou, T.D., Koufos, G.D., 2009. Equidae. In: Koufos, G.D., Nagel, D.(Eds.), The Late Miocene Mammal Faunas of the Mytilinii Basin, SamosIsland, Greece: New Collection. Beiträge zur Paläontologie, 31. pp. 207–

281.Wehrli, H., 1941. Beitrag zur Kenntnis der ‘‘Hipparionen’’ von Samos.

Palaeontologische Zeitschrift 22, 321–386.Weidmann, M., Solounias, N., Drake, R.E., Curtis, J., 1984. Neogene

stratigraphy of the Mytilinii Basin, Samos Island, Greece. Geobios17, 477–490.

G.D. Koufos et al. / Geobios 44 (2011) 237–251 251