Ioannis X. Giaourtsakis Ludwig-Maximilians-Universität, München

Late Neogene Rhinocerotidae of Greece: distribution, diversity and stratigraphical range

Giaourtsakis, I.X., 2003 - Late Neogene Rhinocerotidae of Greece: distribution, diversity and stra- tigraphical range - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 235- 253 [ISSN 0923-9308] Published 1 December 2003

The present contribution provides a comprehensive overview of the Neogene of Greece and evaluates their affinities with the European and Asian species. The distribution, diver- sity and dispersal of the family in the Eastern Mediterranean are discussed in detail. Knowledge of rhinos during the Middle in Greece is still very limited, because of the scarce mate- rial. During the Late Miocene the family was diverse and abundant in Greece. The dominant form, , was probably an African immigrant that invaded Southeastern Europe, Anatolia and the Middle East during the Late Miocene. This rather subhypsodont tandem-horned rhino has already achieved most of the apomorphic characters of the living Dicerotina. The con- temporary species Dihoplus pikermiensis is also common, but the origin and evolutionary rela- tionships among its Miocene and Plio-Pleistocene relatives are still controversial. The Greek Late Miocene representatives of the subfamily are obviously closer related to the Asian genera and , than to the typical aceratheres , Alicornops and Hoploaceratherium found in Central and Western Europe at the same time. Chilotherium and Acerorhinus migrated into Greece from Central Asia trough Anatolia at the beginning of the Vallesian. Chilotherium dominated Samos, where it was represented by at least three species: Ch. samium, Ch. kowalevskii and Ch. schlosseri. In continental Greece, the richest acerathere material comes from the recently discovered locality of Pentalophos-1, where the species Aceratherium kiliasi has been described. Later, this material was assigned partly to Chilotherium and partly to Acerorhinus. The Late Miocene rhino assemblages of Greece are in conformity with the ones from Southeastern Europe and the Eastern Mediterranean, especially with the ones from Anatolia. By the end of the Turolian, all these species became extinct in Greece, unable to survive the paleoen- vironmental changes and the faunal turnover at the Miocene- boundary. Pliocene rhino- ceroses are poorly documented in Greece. Stephanorhinus cf. etruscus has been reported with more or less certainty from several localities, but the available material is not sufficient for useful comparisons.

Correspondence: Ioannis X. Giaourtsakis, Institut für Paläontologie und Historische Geologie, Ludwig-Maximilians-Universität München, Richard-Wagner-Str. 10, D-80333 Munich, Germany, [email protected]

Keywords: Rhinocerotidae, Dihoplus, Greece, Eastern Mediterranean, Miocene, Pliocene

INTRODUCTION Wagner (1848) described and illustrated the The study of fossil Rhinocerotidae in Greece first rhino specimens from Pikermi introdu- has already begun at the middle of the last cing a new species, pachygnathus. century. The first remains of a fossil rhinoce- In subsequent studies, Roth & Wagner (1854) ros from Greece were mentioned by Goldfuss and Wagner (1857) recognised the presence (1841) in a brief note announcing the disco- of a second species at Pikermi, Rhinoceros very of fossil mammals at a new locality schleiermacheri. Gaudry’s (1862-67) monu- north of Athens, Pikermi. A few years later, mental monograph on the Pikermi fauna

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documented adequately both species. In addi- created the species Aceratherium kiliasi. tion, Gaudry described an adult mandible These two papers mark also the last signifi- with well developed incisors as Aceratherium cant contributions on the Neogene fossil sp. Weber (1904, 1905) was the first one who Rhinocerotidae of Greece. studied thoroughly rhino material from More details on the research chronicle as Samos, although the island was known long well as on the geological setting and the ago for its wealth of Neogene mammals. In faunal history of each locality (Fig. 1) are his first contribution, Weber studied the given in the author’s Diploma Thesis, which remains of Rhinoceros pachygnathus and has been accomplished at the University of Rhinoceros schleiermacheri, while in his Athens (Giaourtsakis 1999). A Ph.D. study second work he published a remarkable dealing with an extensive systematic revision sample of acerathere material and described of the fossil rhinoceroses of Greece, inclu- two new species, Aceratherium schlosseri ding old and new unpublished material, is and Aceratherium samium. Andree (1920) currently in progress at the University of later created two additional species based on Munich. The essence of this contribution is to material from Samos, Aceratherium wegneri provide a comprehensive updated overview and Aceratherium angustifrons. Arambourg & of the present knowledge on the Neogene rhi- Piveteau (1929) documented the presence of noceroses of Greece and to evaluate their Rhinoceros pachygnathus and Aceratherium affinities with the European and Asian record. sp. at the Neogene deposits of the Axios Errors often repeated in the past are emended, Valley, near Thessaloniki. Naturally, modern while some new aspects on old controversies workers have gradually modified the generic are introduced. and some of the specific names used by these During the Neogene we can recognise three first authors. successive rhino assemblages in Greece and During the years after the Second World the Eastern Mediterranean, which are inter- War and up to the beginning of the 1980’s rupted by major faunal turnovers at the very little systematic work was undertaken on Middle/Late Miocene and at the Miocene/ the fossil rhinoceroses of Greece. However, Pliocene boundary. during this period quite a few occasional papers and many references in faunal lists of MIDDLE MIOCENE preliminary reports revealed in silence the Middle Miocene localities with large mam- presence of the family in numerous Neogene mals are poorly documented in Greece (De localities of Greece. The rhinoceroses of Bonis & Koufos 1999). The richest and best Pikermi and Samos continued to be mentio- known site from this time period is the locali- ned frequently in major reviews and global ty of Thymiana on the island of Chios. From works on the family, but in most cases neither the older excavations on Chios, the presence new material was introduced nor was the old of Rhinocerotidae indet. has been reported one properly revised. A bright exception to only in a preliminary report (Tobien 1968). A the aforementioned was the important contri- recent Greek-French field campaign has yiel- bution of Geraads (1988), who reviewed a lot ded many new specimens, but no rhinoceros of the known Rhinocerotinae material of was present (De Bonis & Koufos 1999). In an Pikermi and Samos. Geraads established overview of the family Rhinocerotidae, excellent criteria for distinguishing cranial Heissig (1989) stated the presence of "early and dental remains of Ceratotherium neumay- Dicerotini specimens from the Middle ri and Dihoplus pikermiensis. Two years Miocene of Anatolia and Chios". The author later, Geraads & Koufos (1990) studied in considered these specimens as being earlier detail the rich material discovered at the loca- even than the first named species of this line- lity Pentalophos-1, near Thessaloniki, and age, the African Paradiceros mukirii HOOIJER,

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1968. During a conversation, Prof. K. Heissig constricted protocones and the wider, oblique explained that the referred specimens from ectometalophe in M3, often observed in Chios belong to the collections of Prof. H. Chilotherium. The affinities of this material Tobien, but their present status is uncertain recall the morphology of the "West European" (K. Heissig, pers. comm. 2001). Thus these aceratheres, which were present in the references will be considered here as Eastern Mediterranean region at that time Rhinocerotidae indet. (Heissig, 1976). This work was published Two lower premolars from a lignite mine before the modern revision that separated the near the village of Chrysavgi (Mygdonia various "Aceratherium" species in several basin, N. Greece) were described as "Diceros new genera and therefore Symeonidis (1974) pachygnathus" by Psarianos (1958). Dimo- used provisionally the name "Aceratherium poulos (1972) studied also these teeth sp." for the remains of Atalanti. together with three upper molars and some Currently three genera (Aceratherium ribs and placed them to "Dicerorhinus orien- KAUP, 1832; Alicornops GINSBURG & GUÉRIN, talis". Both authors implied a Late Miocene 1979; Hoploaceratherium GINSBURG & age for the fossils. In our opinion, these few HEISSIG, 1989) with about six species are teeth remains are insufficient to establish an considered valid from the later Middle and accurate taxonomic ascription at the present. Late Miocene of Western Europe (MN6 – De Bonis & Koufos (1999), in their review of MN13). Cerdeño (1996) proposed a synony- the Miocene large succession in my between Hoploaceratherium and Greece, considered also this material as Acerorhinus KRETZOI, 1942, but we believe Rhinocerotidae indet., but they proposed a this view needs further investigation. late Middle Miocene age (MN7-MN8) for Regarding the Eastern Mediterranean region, Chrysavgi. This age is based on a micromam- Heissig (1976) reported the presence of malian association, which has been collected Hoploaceratherium aff. tetradactylum and much later and studied by Koliadimou Alicornops simorrense in several Middle (1996). It is not clear however, if the rhino Miocene localities of Turkey. Ginsburg & remains and the micromammals come from Guérin (1979) argued that some of the teeth the same stratigraphic level. of A. simorrense belong to H. aff. tetradacty- The last reference of a Middle Miocene rhi- lum, but Cerdeño & Sánchez (2000) partly noceros from Greece is better documented, supported Heissig’s determinations. This case but until now was overlooked. Symeonidis demonstrates the difficulty of ascribing isola- (1974) described some interesting remains ted teeth to these genera. Another unresolved from a small lignite pit near the city of problem among these taxa is the association Atalanti as Aceratherium sp. The presence of of postcranial material from different locali- a thin, hornless nasal fragment and portions ties with the dental remains and the few avai- of two well-developed second lower incisors, lable skulls. A simple comparison between one of a male and one of a female, leave no the measurement tables of recent publications doubt that the rhino from Atalanti belongs to (Guérin 1980, 1988; Cerdeño & Alcalá 1989; the subfamily Aceratheriinae. The few dental Cerdeño 1989, 1997; Cerdeño & Sánchez remains with strong lingual cingula in the 2000; Heissig in press) reveals that there is premolars, combined with marked paracone no common approach among the specialists. ribs and constricted protocones speak also for This issue became more evident, when two this opinion. The morphology and the size of nearly complete acerathere skeletons from the the teeth exclude an ascription to Brachypo- Early Vallesian locality of Höwenegg (Hegau, therium, too. Moreover, these specimens lack Germany) were described as Aceratherium features such as the prominent antecrochet, incisivum by Hünermann (1989). The fact the weaker paracone rib combined with more that Eppelsheim, the type locality of

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Aceratherium incisivum, may actually com- time period, we find representatives of both promise more than one acerathere species has subfamilies Rhinocerotinae and Aceratheriinae. been previously discussed by Heissig (1972a, 1996b), but not further investigated. Subfamily Rhinocerotinae Moreover, Guérin (1980) has also reported Two species of the subfamily Rhinocerotinae the presence of two different aceratheres in are present during the Late Miocene in Höwenegg. Revision of old and new material Greece: Ceratotherium neumayri (OSBORN, from Eppelsheim and Höwenegg confirms the 1900), formerly known as Diceros pachyg- presence of two different acerathere types in nathus, and Dihoplus pikermiensis (TOULA, these two important localities (Giaourtsakis, 1906). Both species suffer from complicate under study). In this case, an interesting, but nomenclature problems due to false determi- puzzling question may arise: which remains nations and repeated errors in the past, but would correspond to the type skulls of mainly because the holotype of Rhinoceros Aceratherium incisivum (Kaup 1832, 1834)? pachygnathus WAGNER, 1848, a fragmentary Keeping these circumstances in mind, we juvenile mandible housed at the Paleonto- retain provisionally the rhinoceros of Atalanti logical Museum of Munich, belongs actually as Aceratheriini indet. to Dihoplus pikermiensis. Heissig (1975), The Eastern Mediterranean record of who discovered the mistake, proposed to Middle Miocene Rhinocerotidae is better switch the names of the taxa (‘Stephano- documented in Anatolia. Apart from the ace- rhinus pachygnathus’ for ‘Dicerorhinus ratheres Hoploaceratherium aff. tetradacty- pikermiensis’) and abandon the specific name lum and Alicornops simorrense mentioned ‘pikermiensis’, but this brought more chaos above, Heissig (1976) documented the pre- to the already confused literature. To avoid sence of one brachypothere, further confusion we will follow provisional- brachypus and two primitive elasmotheres, ly the concept used by Geraads (1988): Hispanotherium grimmi and ‘Beliajevina’ Ceratotherium neumayri (OSBORN, 1900) for tekkayai (but for different opinions on the Diceros pachygnathus (WAGNER, 1848) and elasmotheres compare Cerdeño 1996 and ‘Dicerorhinus’ pikermiensis (TOULA 1906) for Antoine 2000). Brachypotherium and elas- ‘Dicerorhinus’ orientalis (SCHLOSSER, 1921), motheres have not been recorded in Greece, a younger synonym often used in the literatu- probably due to the poor fossil record during re. However, we prefer to place ‘Dicero- this time period. The end of the Middle rhinus’ pikermiensis within the genus Miocene marks an important faunal turnover Dihoplus BRANDT, 1878 and argue for this caused by climatic changes that favoured the decision. expansion of more open and arid habitats Ceratotherium neumayri was probably an (Janis 1989). These events affected also the African immigrant that invaded Southeastern rhino assemblages of the region. The elas- Europe, Anatolia and the Middle East during motheres, together with Brachypotherium and the Late Miocene. C. neumayri was widespre- the aforementioned acerathere genera disap- ad in many Greek localities and is the best peared from the Eastern Mediterranean and known species from this time period. This were gradually replaced by new immigrants subhypsodont tandem-horned rhino has alre- from Central Asia and Africa at the beginning ady achieved most of the apomorphic charac- of the Late Miocene. ters of the living Dicerotina. A large number of skulls, sometimes with associated mandi- LATE MIOCENE bles, as well as ample postcranial material have been described from Pikermi (Wagner During the Late Miocene rhinoceroses were 1848; Gaudry 1862-67; Guérin 1980; Geraads diverse and abundant in Greece. During this 1988), Samos (Weber 1904; Thenius 1955;

238 GIAOURTSAKIS: Neogene Rhinocerotidae of Greece

Figure 1 Distribution of the family Rhinocerotidae in Greece during the Neogene.

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Geraads 1988), Vathylakkos (Arambourg & hus, based on a few dental differences found Piveteau 1929), Ravin de Zouaves (Koufos in a skull from Maragheh. Thenius (1955) re- 1980), and Pentalophos (Geraads & Koufos examined carefully the type skull from 1990). Some of the references about Pikermi Maragheh and questioned the value of and Samos are partly incorrect and currently, Osborn’s dental differences. He introduced together with new unpublished material, however some new features and proposed the under revision. The presence of this species existence of two separate subspecies: Diceros has been reported at a few more Greek locali- pachygnathus pachygnathus at Pikermi and ties, as well (see Figure1 and Appendix). Thessaloniki and Diceros pachygnathus neu- C. neumayri is very common in the Eastern mayri at Maragheh and Samos. Geraads Mediterranean. Its occurrence is very well (1988) rejected Thenius’ opinion with the documented in Anatolia (Heissig 1975; argument that the proposed differences disap- Geraads 1994; Kappelman et al. 1996). The pear, when additional skulls from the several published record of the species in other localities were taken into consideration. Balkan countries (Bulgaria, former In our opinion, a certain degree of variation Yugoslavia) is sufficient only to confirm its is expected among the various populations presence and has not been studied in detail. due to temporal evolution and/or local envi- C. neumayri has been also reported in a few ronmental factors. The question is how good localities of the former USSR, but the availa- we can document these habitual differences ble data are inadequate. Tsiskarishvili (1987) and if they are enough to establish distinct described the new species Diceros gabuniae species or subspecies. Heissig (1975) obser- from a Vallesian locality in Caucasus as ved a progressive size increasing as well as being a regional variant. A few poorly docu- some morphological specialisations for C. mented and somewhat doubtful references neumayri at the Late Miocene sequences of from Hungary, Italy and Austria may outline Anatolia (e.g. diminution of the fibular facet the Western margin of the species. From at astragalus, increase of gracility at metapo- these, only Thenius (1956) described and illu- dials etc.). The advantage of Heissig’s obser- strated properly some mandibular fragments vations is that they are based on stratigraphi- from the Vienna basin, Austria, but this assig- cally well calibrated localities. On the other nment must be re-examined. Still, this does hand, the number of the available specimens not exclude the possibility that some popula- from these localities is too limited to test the tions of C. neumayri might have wandered in variability and constancy of the suggested these regions. Eastern elements have been features (Heissig, unpublished data; author’s often reported in some West European locali- personal observations). For instance, a small, ties. A few Spanish remains reported as but anatomical almost complete sample from Diceros pachygnathus by Guérin (1980) were the Turkish locality of Mahmutgazi housed at later assigned to Dihoplus schleiermacheri by Museum für Naturkunde Karlsruhe shows a Cerdeño (1989, 1992). In the Middle East, C. remarkable stability near the maximum valu- neumayri is undoubtedly present at Maragheh, es observed in much greater samples from Iran (Thenius 1955), but the rhino material Pikermi. Only a few interesting morphologi- from this locality still lacks its dedicate cal differences could be observed between the monograph. This occurrence marks the two localities. Unfortunately the great bulk of Eastern margin of the species. the fossils from the classical locality of Several authors have discussed the occur- Pikermi has no stratigraphic data and it is not rence of different geographic or temporal clear yet, if it occurs in significantly distinct subspecies of C. neumayri. Osborn (1900) horizons (Theodorou & Nicolaides 1988, originally created "Atelodus" neumayri as a Bernor et al. 1996). Such ‘contaminated’ separate species from "Atelodus" pachygnat- samples may tolerate a greater variability

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when used for statistical purposes. In any be discussed further here. In any case, the case, only careful investigations on both synapomorphies within the group of the morphological and metrical variability Dicerotina are remarkable constant and clear. between well defined population samples can It is enough to mention that the monophyly clarify, whether some of the expected evolu- of this group is probably the only one within tionary trends are sufficient to support the the family which is unequivocally supported distinction of geographic or temporal subspe- by all modern phylogenetical hypotheses pro- cies. Until then, it is better to avoid the appli- posed so far (Heissig 1973, 1989; Guérin cation of different specific or subspecific 1982, 1989; Groves 1983; Prothero et al. names. 1986; Prothero & Schoch 1989; Geraads The oldest and most plesiomorphic repre- 1988, Cerdeño 1995; McKenna & Bell 1997). sentative of the subtribe Dicerotina is The contemporary species Dihoplus piker- Paradiceros mukirii HOOIJER, 1968 from the miensis is also common in Greece and has Middle Miocene of Fort Ternan, Kenya. The been described in Pikermi (Roth & Wagner species has also been reported in North- 1854; Gaudry 1862-67; Geraads 1988), eastern Africa at Beni Mellal, Morocco Samos (Weber 1904; Geraads 1988) and (Guérin 1976) and in the Kisegi Formation in Halmyropotamos (Melentis 1968, 1969). Its Ouganda (Guérin 1994). As stated earlier, presence in other sites, such as the well Heissig (1989) cited the presence of "early known Graecopithecus locality of Tour la Dicerotini specimens from the Middle Reine (Paraskevaidis 1977), is not sufficient- Miocene of Anatolia and Chios", but this ly documented. This large tandem-horned unpublished material appears to be inade- rhino recalls in many features the smaller quate for any useful comparison at the pre- living Sumatra rhino Dicer orhinus sumatren- sent (K. Heissig, pers. comm. 2001). Late sis. In the past they have been often treated Miocene representatives of Dicerotina are as congeneric. This was also the case for all poorly documented in Africa compared to the fossil tandem-horned species apart from richness of the Eastern Mediterranean locali- Coelodonta and the Dicerotina. In time, quite ties. Only three African sites have yielded a few new generic names were introduced for some interesting Dicerotina material and in some of these species. In most cases though, each case a new species has been described. a suprageneric arrangement for the group has The first species comes from the Vallesian of been retained, without always a general Algeria and was originally described as agreement. Currently the origin and evolutio- Dicerorhinus primaevus by Arambourg nary relationships among the Miocene and (1959), but it was later recognised as Diceros Plio-Pleistocene representatives of this line- primaevus by Geraads (1986). The second age are extremely controversial among the species was found in the Turolian site of specialists and even their suprageneric status Duaria (Tunisia) and was described as has been challenged. In the following discus- Diceros douariensis by Guérin (1966). sion, only the most recent opinions will be Recently, Guérin (2000) assigned the rhino reviewed with emphasis on the Miocene spe- remains from Arrisdrift (Sperrgebiet, cies. We believe this discussion is necessary Namibia) to a third species, Diceros in order to establish some stability, at least at australis. All three species appear to be more generic level. primitive than Ceratotherium neumayri. Guérin (1980, 1982, 1989) follows the ‘tra- Due to the existing complicate nomenclatu- ditional’ view for the Miocene and Early re issues, which require a formal request at Pliocene species without an ossified nasal the International Commission, the generic septum and includes them within the genus status of these species as well as their rela- Dicerorhinus GLOGER, 1841. For the rest of tionships with the two living genera will not the Plio-Pleistocene species with partly ossified

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nasal septum he creates the new subgenus smaller than both species. A mandible, origin- Brandtorhinus. The generic names ally figured as Rhinoceros pachygnathus by Lartetotherium GINSBURG, 1974 and Gaudry (1862-67, Pl. xxviii, fig. 1, 2), as well Stephanorhinus KRETZOI, 1942 are considered as adequate material under study, shows that as synonyms. All the species are ranked ‘S.’ pikermiensis has a weak, but still func- together with Coelodonta BRONN, 1831 under tional anterior dentition. Stephanorhinus the Subfamily Dicerorhininae. etruscus lacks completely this feature. The Prothero & Schoch (1989), following basi- appearance of internal cingulum at the pre- cally the phylogeny proposed by Prothero et molars is not always constant in ‘S.’ piker- al. (1986), include the genera Dicerorhinus, miensis and in no case as strong as in Lartetotherium, Stephanorhinus and Coelo- Stephanorhinus etruscus. Moreover, ‘S.’ donta within the subtribe Dicerorhinina, but pikermiensis lacks many of the apomorphic they don’t discuss the specific context of characters shared by the species included in each genus. Alternative phylogenetical hypo- Stephanorhinus (Fortelius et al. 1993), espe- theses for some species were also proposed cially the absence of a partially ossified nasal by Heissig (1981), Groves (1983), Geraads septum. The affinities of ‘S.’ pikermiensis are (1988) and Fortelius et al. (1993). Groves indeed closer to Dihoplus schleiermacheri (1983) suggests that the genus Dicerorhinus than to Stephanorhinus etruscus. Some mor- must be restricted only to the living Sumatra phological and metrical differences are rhino. He supports the use of the generic expected, since Dihoplus schleiermacheri is names Lartetotherium and Stephanorhinus for better documented in the Vallesian of Western some fossil species and states the availability Europe, while Dihoplus pikermiensis in the of the name Dihoplus for D. schleiermacheri. Turolian of Eastern Mediterranean, but these Heissig (1989, 1996a, 1999) regards the can be treated at specific or even subspecific medium sized Middle Miocene Larteto- level. The same is also true for the larger therium sansaniense as clearly plesiomorphic Dihoplus ringstroemi (ARAMBOURG, 1959) of and recognises two evolutionary lineages Eastern Asia. during the Late Miocene and Pliocene: The Cerdeño (1995) presents a cladistic analysis first evolves successively through the species of the whole family Rhinocerotidae and intro- Dihoplus schleiermacheri, ‘Dicerorhinus’ duces some new ideas. The author (Cerdeño megarhinus and ‘Dicerorhinus’ jeanvireti, 1995, 1998) ranks all the Miocene species while the other one through Stephanorhinus into a single genus, Lartetotherium, while the pikermiensis and Stephanorhinus etruscus. Plio-Pleistocene species are all placed within Heissig’s arguments uniting ‘Stephanorhinus’ the genus Stephanorhinus (including also the pikermiensis with Stephanorhinus etruscus species with no partially ossified nasal sep- and separating it from Dihoplus schleierma- tum, such as the Early Pliocene ‘Dicero- cheri are the following: the ‘medium size’ of rhinus’ megarhinus). Apart from the strati- ‘S.’ pikermiensis (compared to the ‘large’ D. graphic separation there are not many argu- schleiermacheri), the narrower skull, the loss ments offered to support this arrangement, of all the enlarged incisors and the strong since the analysis considers most of the taxa interior cingula in the upper premolars. The at generic level. Another interesting result of size of the many skulls from Pikermi and Cerdeño’s analysis is that the subtribe Samos that we know is about the same as the Dicerorhinina was not supported. Instead of size of the only known complete skull of this, Stephanorhinus and Coelodonta were Dihoplus schleiermacheri, the lectotype from placed together with Elasmotherium and Eppelsheim. The differences in the postcra- Ninxiatherium (=Parelasmotherium) under nial skeleton are also insignificant. the subtribe Elasmotheriina. It is obvious Stephanorhinus etruscus is considerably (Cerdeño 1995: Tab. 4, node 20; Tab. 5, node

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24) that the only character practically suppor- horn. Skulls found in Sandelzhausen ting this ‘monophyly’ was the presence of a (Bavaria, Germany) unequivocally have only partially or totally ossified nasal septum, a one nasal horn (Heissig 1972b; Groves 1983; fact also admitted by the author, since the rest author’s personal observations). of the apomorphic characters gathered at The subject requires certainly further study these nodes were also shared by other groups. and discussion. Regarding the Miocene taxa Out of the 45 taxa, only these four genera had concerned here, we can note the following states 1 or 2 for character 4, i.e. partially or conclusions: We follow Heissig (1972b, totally ossified nasal septum (ibid. Tab. 2, 3). 1989, in press) and Groves (1983) in conside- Erroneously, this character state was interpre- ring Lartetotherium sansaniense as a primiti- ted by the analysis as a synapomorphy ve Rhinocerotinae, separate, and near the (Cerdeño 1995: p. 20), although it is clear base of the subfamily. The phyletic position from the paleontological record that the of more primitive forms, such as acquisition of an ossified nasal septum has ‘Dicerorhinus’ abeli in Southeast Asia is been achieved at least twice independently poorly documented. The same is true for the within the groups of Stephanorhinus-Coelo- Spanish ‘Dicerorhinus’ montesi and the small donta and Elasmotherium-Ninxiatherium as a ‘Dicerorhinus’ steinheimensis in Europe. If result of parallelism. This view is also sup- the African ‘Dicerorhinus’ leakeyi or the ported by another cladistic analysis (Antoine Chinese ‘Dicerorhinus’ cixianensis belong 2000), in which the presence and the deve- also to Lartetotherium or to Dihoplus cannot lopment of an ossified nasal septum were be decided at the present. We believe though coded as two separate binary characters inste- that the large Late Miocene species: ad of a single multiple one. Cerdeño (1995) ‘Dicer orhinus’ schleiermacheri in Central discussed also the possibility of an indepen- and Western Europe, ‘D.’ pikermiensis in dent acquisition of the ossified nasal septum Southeastern Europe and the Eastern but did not adopt it in her final classification. Mediterranean and ‘D.’ ringstroemi in For these reasons, we do not favour the clas- Eastern Asia are morphologically and metri- sification of Stephanorhinus and Coelodonta cally very closely related. We propose there- within the Elasmotheriina (sensu Cerdeño fore the generic name Dihoplus BRANDT, 1995). 1878, originally erected for Rhinoceros The genus Lartetotherium has been erected schleiermacheri KAUP, 1832, to include all of by Ginsburg (1974) for the species Rhinoceros them. Dihoplus has in any case the priority sansaniensis LARTET, 1851 from the Middle over all the alternative generic names mentio- Miocene locality of Sansan in France. There ned above (Stephanorhinus, Lartetotherium) are many primitive and autapomorphic char- and may be convincingly extended to include acters found in this species (Ginsburg 1974, other Neogene species, as well. A stable and Groves 1983, Heissig in press), but the pre- persuading phylogeny for the whole group, sence or not of a frontal horn tends to be the from the Miocene forms to the highly specia- most significant and controversial one. lised Coelodonta and the living Sumatran Ginsburg (1974) did not discuss the horn rhino, is yet to be given. arrangement in his generic diagnosis, while the original specific diagnosis (Lartet 1851) Subfamily Aceratheriinae was also somehow undecided, because of the The Greek representatives of the subfamily craniocaudal distortion of the type skull. Aceratheriinae during the Late Miocene are Guérin (1980) and Cerdeño (1989) favour the obviously closer related to the Asian genera presence of a frontal horn in the type material Chilotherium and Acerorhinus than to the from Sansan. Heissig (in press) believes that typical aceratheres Aceratherium, Alicornops the French specimens could lack a frontal and Hoploaceratherium found in Central and

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West Europe at the same time. Chilotherium 1913). and Acerorhinus migrated into Greece from The smaller and less hypsodont Ch. sami- Central Asia trough Anatolia at the beginning um is the most primitive of the three species. of the Vallesian. Chilotherium has dominated In Anatolia, Ch. samium first appears in Samos, where it was represented by at least Vallesian (MN10/11) about at the same time, three species: Ch. samium, Ch. schlosseri and but not together, with the more hypsodont Ch. kowalevskii. The richest acerathere mate- Ch. habereri (Heissig 1975). Acerorhinus, rial in continental Greece comes from the another acerathere genus that lacks the spe- recently discovered locality of Pentalophos-1, cialization of Chilotherium, was also present where the species Aceratherium kiliasi has at that time in Anatolia with the species been described. Later, this material was partly Acerorhinus zernowi (BORISSIAK, 1914). assigned to Chilotherium and partly to Unfortunately, many of Samos’ classical col- Acerorhinus. The few references on lections are missing accurate stratigraphic ‘Aceratherium sp.’ from Pikermi, Vathylakos data for comparison with Anatolia (compare and Chomateri that are based on scanty finds Solounias 1981a,b and Bernor et al. 1996). could also belong to these genera. Ch. kowalevskii with semimolariform molars The island of Samos has been known for its was found on material from Grebeniki wealth of Miocene mammals since the middle (Ukraine). It is probably a later immigrant of the last century. Weber (1905) studied a (MN 12) that represents a divergent evolutio- remarkable sample of acerathere material nary lineage. The larger and high crowned including several skulls with associated man- Ch. schlosseri is the typical Turolian chilot- dibles as well as partly articulated limb bones here. The advanced stage of Ch. schlosseri is and described two new species, Aceratherium very similar to the contemporary Chinese schlosseri and Aceratherium samium. Andree genotype Ch. andersoni and both species can (1920) created later two additional species be possibly traced back to Ch. habereri based on material from Samos: Aceratherium (Heissig 1975, 1996a). All these chilothere wegneri and Aceratherium angustifrons. forms indicate diverse ecological preferences, Unfortunately, Weber’s entire material as well which explain their parallel existence. as many specimens of Andree’s material have Additional studies on the unpublished mate- been destroyed during the Second World War. rial from Samos are necessary to establish At present, only a skull housed at the clear affinities for all species. Geological Institute of Padova and referred to In continental Greece, the richest acerathere Ch. wegneri by Leonardi (1947) as well as material comes from the recently discovered another one with associated mandible at the locality of Pentalophos-1, where the species Paleontological Institute of Hamburg descri- Aceratherium kiliasi has been described by bed as Ch. schlosseri by Lehman (1984) Geraads & Koufos (1990). The material comprise the intact published chilothere comprises of several skulls and mandibles, material from Samos. However, a wealth of but no postcranials. Heissig (1996a, 1999) unpublished material is expected to be availa- designated the material from Pentalophos ble at the numerous museums where fossils partly to Chilotherium kiliasi and partly to from Samos are stored (Solounias 1981a,b). Acerorhinus zernowi, with the remark that the Ringström (1924) placed rightfully all these first one could be specifically identical with species to his new genus Chilotherium. Chilotherium samium. Heissig did not provi- Heissig (1975) published a preliminary report de any arguments for this separation. Judging on the Miocene rhinoceroses of Anatolia. In from the description and the illustrations of this report, Heissig considered Ch. wegneri ‘Aceratherium’ kiliasi, part of the material synonymous with Ch. schlosseri and Ch. reveals many similarities with Chilotherium angustifrons with Ch. kowalevskii (PAVLOW, samium. This point has been also evaluated

244 GIAOURTSAKIS: Neogene Rhinocerotidae of Greece

by Geraads & Koufos (1990), but they consi- at a time where no other generic name was dered Chilotherium samium as Aceratherium. available, because it clearly differed from the The low and broad holotype skull with mandible of Aceratherium incisivum from flattened dorsal profile and elevated orbita Eppelsheim. Since then, this reference has (upper border at the same level as the frontal) been frequently repeated in numerous publi- shows more affinities to the Chilotherium cations, sometimes also as Aceratherium inci- cranial type (Ringström 1924) than to the sivum and others as Aceratherium cf. incisi- Vallesian ‘Aceratherium’ skulls form Eppels- vum. It appears that only Geraads & Koufos heim (Kaup 1832, 1834) and Höwenegg (1990) have examined the mandible again, (Hünerman 1989). The upper dentition of the but they also considered the material too poor type skull (Geraads & Koufos 1990, pl. 3, for a specific determination. This is indeed a fig. 4 ) appears also to be identical with the puzzling specimen. Gaudry’s description (destroyed) type material of Ch. samium from offers some interesting characters but the Samos figured by Weber (1904, Pl. II, Fig. 5) accompanying figure is not very informative and the unpublished material from Turkey (ibid., pl. 33, fig. 6). The short distance housed at the Paleontological Museum of between the second lower incisors, mentioned Munich and referred to the same species by also by Geraads & Koufos, excludes an Heissig (1975). Some important diagnostic ascription to Chilotherium. The large size of characters are based on the mandible, espe- the mandible, the short diastema combined cially on the symphysial part. The absence of with a long and upraised symphysis could an anteriorly broadened mandibular symphy- speak for an Acerorhinus determination. sis, a synapomorphy of the true chilotheres, However it is not clear if the symphysis is led Geraads & Koufos (1990) to exclude an labially concave. The small metapodials iden- ascription to Chilotherium (only one of the tified by Gaudry as "Rhinocéros indéterminé three mandibles was "slightly broadened"). de petite taille" may actually be more useful. On the other hand, a narrower symphysis They belong also to an acerathere and proba- often combined with a concave labial side is bly to the same species. Their size and graci- considered as diagnostic for the genus lity index correspond well to the values found Acerorhinus. This condition, unknown to the for Acerorhinus (Bohlin 1937, Cerdeño three ‘West European’ acerathere genera dis- 1996). Unpublished material under study sup- cussed earlier, is documented among others ports these observations, but a re-examination in Acerorhinus paleosinensis (RINGSTRÖM, of Gaudry’s collection is required to establish 1924), Acerorhinus tsaidamensis (BOHLIN, final conclusions. For this reason these speci- 1937), Acerorhinus fuguensis (DENG, 2000) mens are temporary considered as Acera- and the genotype Acerorhinus zernowi theriini indet. (Borissiak 1914; Heissig 1975; Cerdeño Arambourg & Piveteau (1929) described a 1996). Geraads & Koufos (1990) offer many strong second lower incisor from Vathilakos 3 other interesting characters in their detailed (Axios Valley, N. Greece) as ‘Aceratherium description and it is quite possible that at sp.’. An acerathere mandible from the locality least some part of the material belongs indeed of Chomateri near the classical Pikermi was to a new species. Since a close re-examina- cited as ‘Aceratherium sp.’ by Marinos & tion of the original material from Pentalophos Symeonidis (1974) in a preliminary faunal is necessary for final conclusions, we prefer list. Both specimens are here referred to as to keep the acerathere remains from this site Aceratheriini indet., since they could belong temporary under their original name to any of the aforementioned genera. ‘Aceratherium’ kiliasi. The present status of the tribe Aceratheriini in Gaudry (1862-67) described an acerathere Southeastern Europe and Asia is extremely mandible from Pikermi as Aceratherium sp. complicated and in fact badly needs a revi-

245 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

sion. Three genera, Chilotherium, Subchilo- Europe, only the less specialized Dicero- therium and Acerorhinus are currently recog- rhinina managed to resist into the Pliocene. nised during the terminal Middle and the Late Miocene, but their context and even their PLIOCENE suprageneric affinities are still debatable and During the Early Pliocene (Ruscinian), the not very well understood (Heissig 1975, localities with large mammals in Greece are 1989, 1996a; Qiu et al. 1988; Bayshashov extremely rare (Koufos & Kostopoulos 1993; Cerdeño 1996; Deng 2000). More than 1997). The presence of Rhinocerotidae indet. twenty species have been assigned to these has been cited in the faunal lists of three genera, ranging from Portugal to Japan. The localities (Apollakia, Nea Silata, Alatini), but limits within these forms are not always evi- no specimens were published. The Apollakia dent and the distinguishing characters Formation, placed near the south-west coast designated are rather confusing than convin- of the island Rhodes, is the best known of cing. Many of the newly introduced species these three. Apollakia features a typical lack a differential diagnosis from older spe- Ruscinian micro- and macromammalian local cies. In a preliminary report, Heissig (1975) fauna, which has been described by van de presented a table summarising many of the Weerd et al. (1982). The presence of known species together with an initial phylo- Rhinocerotidae indet. is referred only in the genetical hypothesis for the group, but not faunal list of the site. Syrides (1990) noted enough arguments were presented to support the presence of Rhinocerotidae fragments this. The number of new species has been among the few remains of large mammals doubled since Heissig’s summary. It is clear found at Nea Silata (Chalkidiki). The micro- that a basic phylogenetical framework, not to mammals of this site were originally dated to speak for a solution, can be discussed only basal Pliocene (Syrides 1990). New studies when regional revisions in Greece, Turkey on the rich micromammalian fauna of Nea and the rest of the Balkans as well as in the Silata indicate a transitional Turolian/ former USSR, the Middle East and especially Ruscinian character for the locality (K. in China and Mongolia are completed. Vassiliadou, pers. comm. 2001). Marinos (1965) mentioned in a short comment the Conclusions presence of several mammals including The Late Miocene rhino assemblages of ‘Rhinoceros sp.’ at the locality of Alatini, Greece are in conformity with the ones from near Thessaloniki. A few years later, Southeastern Europe and the Eastern Sickenberg (1972) described the carnivore Mediterranean, especially with the ones from Nyctereutes donnezani from the same locality Anatolia. From a biogeographical aspect, suggesting a Ruscinian age. In all three cases, these results correspond to the Eastern we can assume that the available material regional block of Fortelius et al. (1996) or, in was too scarce to support any further investi- a closer approach, to the Greco-Iranian or gation. sub-Paratethyan mammalian province of The scantiness of large mammal faunas Bernor (1984) and De Bonis et al. (1993, during the Ruscinian characterises the whole 1999). By the end of the Turolian, all these Eastern Mediterranean region. The most com- species became extinct in Greece, unable to mon rhino in Western Europe during this survive the paleoenvironmental changes and period is the large ‘Dicerorhinus’ the faunal turnover at the Miocene-Pliocene megarhinus. Only recently Guérin & Sen boundary. The aceratheres disappeared com- (1998) documented adequately the presence pletely from Europe and Western Asia, while of this species at the Early Pliocene locality the subtribe Dicerotina was restricted to of Çalta, near Ankara (Turkey). ‘Dicerorhinus’ Africa, where it survived until nowadays. In megarhinus probably represents the terminal

246 GIAOURTSAKIS: Neogene Rhinocerotidae of Greece

evolutionary stage of the large Miocene E. Georgiadou, E. Velitzelos and M. Dicerorhinina, here placed under the genus Dermitzakis for their kind interest and advice Dihoplus. As discussed above, the origin and during and after my studies at the University evolutionary relationships among the of Athens. I am deeply indebted to Dr. J. Miocene and Plio-Pleistocene representatives Franzen and Prof. F. Steininger for their hos- of this subtribe are still controversial. pitality and invaluable assistance during my Middle and Late Pliocene localities with long-term visit at the Naturmuseum und large mammals are rather frequent in Greece. Forschungsinstitut Senckenberg. Prof. F. Despite the numerous localities, knowledge Schrenk provided access to material housed of rhinos during this time period is also very at the Hessisches Landesmuseum Darmstadt. limited, because of the scarce material. It is Dr. R. Frey and D. Schreiber are kindly ack- notable that the published material exceeds nowledged for the permission to study the the number of five specimens in one site collections of the Staatliches Museum für only, Sesklo (Thessaly, Central Greece). Naturkunde Karlsruhe. For valuable help and Symeonidis (1992), in a brief description, discussion I am further indebted to Drs. A. assigned the few postcranial materials from Athanasiou, T. Deng, U. Göhlich, Ch. Hetler, Sesklo to Dicerorhinus cf. etruscus. Later, H.W. Mittmann, M. Moser, G. Rössner, S. Athanasiou (1998) re-examined the Sesklo Roussiakis, G. Storch, C. Theocharopoulos, fauna and observed some significant differen- U. Uhlig as well as to Profs. V. Fahlbusch ces from the standards established for and N. Solounias. I am very grateful to Dr. E. Stephanorhinus etruscus by Guérin (1980) Cerdeño and Dr. J. de Vos for the critical rea- and Fortelius et al. (1993). Therefore he ding of the manuscript and their useful sug- referred the material temporarily to gestions. Early stages of this work (1999) Stephanorhinus sp. Further isolated remains have been partially supported by the National of Rhinocerotidae have been reported with Scholarship Foundation of Greece (IKY). My more or less certainty as Stephanorhinus sp. current studies on the fossil rhinoceroses of or Stephanorhinus cf. etruscus from several Greece are generously supported by the other Late Pliocene sites (Fig. 1, Appendix), Deutsche Akademische Austauschdienst but the available material is not sufficient for (DAAD). Finally, I would like to take this useful comparisons. opportunity to thank Dr. H. de Bruijn for his lifetime contribution to the study of the fossil ACKNOWLEDGEMENTS faunas of the Eastern Mediterranean region. I wish to thank Prof. C. Doukas (University of Athens) for inviting me to participate at REFERENCES the conference on Migration and Distribution Andree, J., 1920 - Rhinocerotiden aus dem Unterpliozän of Tertiary Mammals in Eurasia, as well as von Samos - Paläontologische Zeitschrift 20: 189-212 for his continuous interest in, and support of, Antoine, P.O., 2000 - Origine et differenciation des my research. I would also like to thank Prof. Elasmotheriina parmi les Rhinocerotidae (Mammalia, K. Heissig for sharing with me his passion Perissodactyla): Analyse cladistique et implications and his knowledge of fossil rhinoceroses and biostratigraphiques et paléobiogéographiques - Ph.D. for providing liberal access to the collections Thesis, Muséum National d’Histoire Naturelle, Paris of the Bayerische Staatssammlung für Arambourg, C., 1959 - Vertebres continentaux du Paläontologie und Historische Geologie, miocene superieur de l'Afrique du nord - Publication München. I am indebted to Prof. G. Service Carte Géologique d’Algérie, Paléontologie 4: Theodorou for facilitating my work at the 1-159 Geological Institute and the Paleontological Arambourg, C. & Piveteau, J., 1929 - Les vertébrés du Museum of the University of Athens. I also Pontien de Salonique - Annales de Paléontologie 18: wish to acknowledge Profs. N. Symeonidis, 57-140

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Athanassiou, A., 1998 - Contribution to the study of the Forschungsinstitut Senckenberg 197: 187-203 fossil mammals of Thessaly - Gaia 5: 1-354 [in Cerdeño, E., 1998 - Diversity and evolutionary trends of Greek] the Family Rhinocerotidae (Perissodactyla) - Bartsiokas, A., 1998 - Paleontology of Kythira Island - Palaeogeography, Palaeoclimatology, Palaeoecology Publications of the Society of Kytherian Studies 9: 141: 13-34 1-96 [in Greek] Cerdeño, E. & Alcalá, L., 1989 - Aceratherium Bayshashov, B.U., 1993 - Neogene rhinoceroses of alfambrense sp. n., nuevo rinocerótido del Vallesiense Kazakhstan - Izdatel'stvo Gylym Almaty, Kazakhstan Superior de Teruel (España) - Revista Española de [in Russian] Paleontología 4: 39-51 Bernor, R.L., 1984 - A zoogeographic theater and bio- Cerdeño, E., & Sánchez, B., 2000 - Intraspecific varia chronologic play: The time/biofacies phenomena of tion and evolutionary trends of Alicornops simorrense Eurasian and African Miocene mammal provinces - (Rhinocerotidae) in Spain - Zoologica Scripta 29: Paléobiologie Continentale 14: 121-142 275-305 Bernor, R.L., Solounias, N., Swisher III, C.C. & Van De Bonis, L., Bouvrain, G. & Koufos, G.D., 1999 - Couvering, J. A., 1996 - The corellation of three Palaeoenviroments of Late Miocene primate localities classical "pikermian" mammal faunas - Maragheh, in Macedonia, Greece - in: Augustí, J., Rook, L., & Samos and Pikermi- with the European MN Unit Andrews, P. (eds.) - Hominoid Evolution and climate Systm - in: Bernor, R. L., Fahlbush, V., & Mittman, Change in Europe vol. 1: The Evolution of Neogene H.-W. (eds.) - The Evolution of Western Eurasian Terrestrial Ecosystems in Europe - pp. 415-435, Neogene Mammal Faunas - pp. 137-154, Columbia Cambridge University Press University Press, New York De Bonis, L., Brunet, M., Heintz, E. & Sen, S., 1993 - Bohlin, B., 1937 - Eine Tertiäre Säugetier-Fauna aus La province gréco-irano-afghane et la répartition des Tsaidam - Palaeontologia Sinica 14 (1): 1-111 faunes mammaliennes au Miocène supérieur - Borissiak, A.A., 1914 - Mammifères fossiles de Paleontologia i Evolució 24-25: 103-112 Sébastopol, I. - Mémoires du Comité Géologique, De Bonis, L. & Koufos, G.D. 1999 - The Miocene large Petrograd 137: 1-154 mammals succesion in Greece - in: Augustí, J., Rook, Brandt, J., 1878 - Tentamen synopseos rhinocerotidum L. & Andrews, P., (eds.) - Hominoid Evolution and viventium et fossilium - Mémoires l’Académie climate Change in Europe vol. 1: The Evolution of Impériale des Sciences de St. Pétersbourg 7e Série 26 Neogene Terrestrial Ecosystems in Europe - pp. 205- (5): 1-66 237, Cambridge University Press Bronn, H., 1831 - Über die fossilen Zähne eines neuen Deng, T., 2000 - A new species of Acerorhinus Geschlechtes aus der Dickhäuter-Ordnung Coelodonta, (Perissodactyla, Rhinocerotidae) from the Late Höhlenzahn - Neues Jahrbuch für Mineralogie, Miocene in Fugu, Shaanxi, China - Vertebrata Geologie und Paläontologie, Jahrgang 1831: 51-61 Palasiatica 38(3):203-217 Cerdeño, E., 1989 - Revisión de la Sistemática de los Dimopoulos, G., 1972 - Dicer orhinus orientalis, aus dem rinocerontes del Neógeno de España - Colección Obermiozän (Pont) des Beckens von Lagadas Tesis Doctorales Universidad Complutense de Madrid (Mazedonien/ Griechenland) - Folia Biochaemica et 306/89: 1-429 Biologia Graeca 9: 47-60 Cerdeño, E., 1992 - Spanish Neogene rhinoceroses - Fortelius, M., Mazza, P. & Sala, B., 1993 - Stephano- Palaeontology 35(2): 297-308 rhinus (Mammalia, Rhinocerotidae) of the western Cerdeño, E., 1995 - Cladistic analysis of the Family European Pleistocene, with a revision of Rhinocerotidae - American Museum Novitates 3143: Stephanorhinus etruscus (Falconer, 1868) - 1- 25 Palaeontographia Italica 80: 63-155 Cerdeño, E., 1996 - Rhinocerotidae from the Middle Fortelius, M., Werdelin., L., Andrews, P., Bernor, R.L., Miocene of the Tung-gur formation, Inner Mongolia Gentry, A., Humphrey, L., Mittmann W. & Viranta, (China) - American Museum Novitates 3184: 1- 43 S., 1996 - Provinciality, diversity, turnover and paleo- Cerdeño, E., 1997 - Rhinocerotidae from the Turolian ecology in land mammal faunas of the later Miocene site of Dorn-Dürkheim 1 (Germany) - Courier of western Eurasia - in: Bernor, R. L., Fahlbush, V. &

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GIAOURTSAKIS: Neogene Rhinocerotidae of Greece

Mittman, H.-W. (eds.) - The Evolution of Western Documents des Laboratoires de Géologie de la Eurasian Neogene Mammal Faunas - pp. 414-448, Faculté des Sciences de Lyon 16: 1-50 Columbia University Press, New York Guérin, C., 1976 - Les restes de rhinocéros du gisement Gaudry, A, 1862-67 - Animaux fossiles et géologie de Miocène de Beni Mellal, Maroc - Géologie l'Attique - F. Savy (ed.), Paris Méditerranéenne 3: 105-108 Geraads, D., 1986 - Sur les relations phylétiques de Guérin, C., 1980 - Les rhinoceros (Mammalia, Dicerorhinus primaevus Arambourg, 1959, rhinocéros Perissodactyla) du Miocene terminal au Pleistocene du Vallésien d'Algérie - Comptes Rendus de superieur en Europe occidentale. Comparaison avec l'Académie des Sciences Série II 302: 835-837 les especes actuelles - Documents des Laboratoires de Geraads, D., 1988 - Révision des Rhinocerotinae Géologie de la Faculté des Sciences de Lyon 79: (Mammalia) du Turolien de Pikermi. Comparaison 1-1185 avec les formes voisins - Annales de Paléontologie Guérin, C., 1982 - Les Rhinocerotidae (Mammalia, 74: 13-41 Perissodactyla) du Miocène terminal au Pléistocène Geraads, D., 1994 - Les gisements de mammifères du supérieur d'Europe occidentale compares aux espèces Miocène supérieur de Kemiklitepe, Turquie: 5. actuelles: Tendances évolutives et relations Rhinocerotidae - Bulletin du Muséum National phylogénétiques - Géobios 15: 599-605 d'Histoire Naturelle Section C 16(1): 81-95 Guérin C., 1988 - Contributions a l'étude du gisement Geraads, D. & Koufos, G.D., 1990 - Upper Miocene Miocène supérieur de Montredon (Hérault). Les Rhinocerotidae (Mammalia) from Pentalophos-1, grands Mammifères VI: Les Périssodactyles, Macedonia, Greece - Palaeontographica (A) 210: Rhinocerotidae - Palaeovertebrata, Mémoire extraor- 151-168 dinaire 1988: 97-134 Giaourtsakis, I.X., 1999 - The fossil rhinoceroses of Guérin, C., 1989 - La famille des Rhinocerotidae Greece: localites and literature - Unpublished (Mammalia, Perissodactyla): systématique, histoire, Diploma Thesis, University of Athens [in Greek] évolution, paléoécologie - Cranium 6(2): 3-14 Ginsburg, L., 1974 - Les rhinocérotidés du Miocène de Guérin, C., 1994 - Les Rhinocéros (Mammalia, Sansan (Gers) - Comptes Rendus de l'Académie des Perissodactyla) du Neogene de l'Ouganda - in: Senut, Sciences Série D 278: 597-600 B. & Pickford, M. (eds.) - Geology and palaeobiolo Ginsburg, L. & Guérin, C., 1979 - Sur l'origine et l'ex gy of the Albertine rift valley, Uganda-Zaire; Volume tension stratigraphique du petit Rhinocérotidé II, Palaeobiology - pp. 263-279, Centre International Miocène Aceratherium (Alicornops) simorrense Pour la Formation et les Echanges Geologiques, (Lartet, 1851), nov. subgen. - Compte Rendu Occasional Publication 29, Paris Sommaire des Séances de la Société Géologique de Guérin, C., 2000 - The Neogene rhinoceroses of Namibia France 3: 114-116 - Palaeontologia Africana 36: 119-138 Ginsburg, L. & Heissig, K., 1989 - Hoploaceratherium, a Guérin, C. & Sen, S., 1998 - La gisement de vertébrés new generic name for "Aceratherium" tetradactylum pliocènes de Çalta, Ankara, Turquie. 6. - in: Prothero, D.R. & Schoch, R.M. (eds.) - The evo- Rhinocerotidae - Geodiversitas 20 (3): 397-407 lution of Perissodactyls - Oxford Monographs on Heissig, K., 1972a - Paläontologische und geologische Geology and Geophysics 15: 399-417 Untersuchungen im Tertiär von Pakistan 5: Gloger C.L., 1841 - Gemeinnütziges Hand- und Rhinocerotidae (Mammalia) aus den unteren und Hilfsbuch der Naturgeschichte - Breslau mittleren Siwalik-Schichten - Abhandlungen der Goldfuss, G.A., 1841 - Brief an Geologisches Jahrbuch - Bayerischen Akademie der Wissenschaften, Neues Jahrbuch für Mineralogie, Geologie und Mathematische Naturwissenschaftliche Klasse 152: Paläontologie, Jahrgang 1841: 357-358 1-112 Groves, C.P., 1983 - Phylogeny of the living species of Heissig, K., 1972b - Die obermiozäne Fossil-Lagerstäte rhinoceros - Zeitschrift für Zoologische Systematik Sandelzhausen 5.: Rhinocerotidae (Mammalia), und Evolutionsforschung 21: 293-313 Systematik und Ökologie - Mitteilungen der Guérin, C. 1966 - Diceros douariensis nov. sp., un Bayerischen Staatssammlung für Paläontologie und rhinocéros du Mio-Pliocène de Tunisie du nord - Historische Geologie 12: 57-81

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Heissig, K., 1973 - Die Unterfamilien und Tribus der cahier - Heyer Ed., Darmstadt. rezenten und fossilen Rhinocerotidae (Mammalia) - Kappelman, J., Sen, S., Fortelius, M., Duncan, A., Säugetierkundliche Mitteilungen 21: 25-30 Alpagut, B., Crabaugh, J., Gentry, A., Lunkka, J.-P., Heissig, K., 1975 - Rhinocerotidae aus dem Jungtertiär McDowell, F., Solounias, N., Viranta, S. & Werdelin, L., Anatoliens - Geologisches Jahrbuch (B) 15: 145-151 1996 - Chronology and biostratigraphy of the Heissig, K., 1976 - Rhinocerotidae (Mammalia) aus der Miocene Sinap Formation of Central Turkey - in: Anchitherium-Fauna Anatoliens - Geologisches Bernor, R. L., Fahlbush, V. & Mittman, H.-W. (eds.) - Jahrbuch (B) 19: 3-121 The Evolution of Western Eurasian Neogene Heissig, K., 1981 - Probleme bei der cladistischen Mammal Faunas - pp. 78-95, Columbia University Analyse einer Gruppe mit wenigen eindeutigen Press, New York Apomorphien: Rhinocerotidae - Paläontologisches Koliadimou, K.K., 1996 - Paleontological and biostrati Zeitschrift 55: 117-123 graphical study of the Neogene-Quaternary micro Heissig, K., 1989 - The Rhinocerotidae - in: Prothero, mammals from Mygdonia basin - Ph.D. Thesis, D.R. & Schoch, R.M. (eds.) - The evolution of University of Thessaloniki [in Greek] Perissodactyls - Oxford Monographs on Geology and Koufos, G.D., 1980 - Paleontological and stratigraphical Geophysics 15: 399-417 study of the Neogene continental deposits of Axios Heissig, K., 1996a - The stratigraphical range of fossil Valley (Macedonia, Greece) - Scientific Annals of the rhinoceroses in the Late Neogene of Europe and Faculty of Physics and Mathematics, Aristotelian Eastern Mediterranean - in: Bernor, R. L., Fahlbush, Univercity of Thessaloniki 19: 1-322 [in Greek] V. & Mittman, H.-W. (eds.) - The Evolution of Koufos, G.D. & Kostopoulos, D. S., 1997 - Western Eurasian Neogene Mammal Faunas - pp. Biochronology and succesion of the Plio-Pleistocene 339-347, Columbia University Press, New York macromammalian localities of Greece – in: Aguilar, Heissig, K., 1996b - Ein Schädel von Hoploaceratherium J.-P., Legendre, S. & Michaux, J. (eds.) - Actes du aus dem Obermiozän Bayerns - Mitteilungen der Congres BiochroM '97 - Mémoires et Travaux, Ecole Bayerischen Staatssammlung für Paläontologie und Pratique des Hautes Itudes, Institut de Montpellier Historische Geologie 96: 145-155 21: 619-634 Heissig, K., 1999 – Family Rhinocerotidae – in: Rössner, Kretzoi, M., 1942 - Bemerkungen zum System der nach G. & Heissig, K. (eds.) – The Miocene Land miozänen Nashorn Gattungen - Földtani Közlöny 72 Mammals of Europe – pp. 175-188, Verlag Dr. F. (4-12): 309-318 Pfeil, München Lartet, E., 1851 - Notice sur la colline de Sansan - Heissig, K., (in press) - The fossil rhinoceroses of Portes édit., Auch Rudabanya. Lehmann, U., 1984 - Notiz über Säugetierreste von der Hooijer, D.A., 1968 - A rhinoceros from the Late Insel Samos in der Sammlung des Geologisch- Miocene of Fort Ternan, Kenya - Zoologische Palaontologischen Instituts und Museums Hamburg - Mededelingen 43(6): 77-92 Mitteilungen aus dem Geologischen-Paläonto- Hünermann, K.A., 1989 - Die Nashornskelette logischen Institut der Universität Hamburg 57: (Aceratherium incisivum Kaup 1832) aus den 147-156 Jungtertiär vom Höwenegg im Hegau (Südwest- Leonardi, P., 1947 - Resti fossili inediti di rinoceronti deutschland) - Andrias 6: 1-116 conservati nelle collezioni dell' Istituto Geologico Janis, C.M., 1989 - A climatic explanation for patterns of dell’ Università di Padova - Memoire dell’Instituto diversity in ungulate mammals - Palaeontology 32: Geologico dell’Università di Padova 15: 3-30 463-481 Marinos, G., 1965 - Beiträge zur Kenntnis der Kaup, J., 1832 - Über Rhinoceros incisivus Cuv., und Verbreitung des Pleistozäns in Mazedonien - eine neue Art, Rhinoceros schleiermacheri - Isis Scientific Annals of the Faculty of Physics and von Oken 1832: 898-904 Mathematics, Aristotelian University of Thessaloniki Kaup, J., 1834 - Description d'ossements fossiles de 9: 95-111 mammifères inconnus jusqu'à présent qui se trouvent Marinos, G. & Symeonidis, N. K., 1974 - Neue Funde au Muséum grand-ducal de Darmstadt. Troisième aus Pikermi (Attica, Griechenland) und eine allge

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meine geologische Übersicht dieses Paläonto- Knochenüberreste von Pikermi in Griechenland - logisches Raumes - Annales Géologiques des Pays Abhandlungen der Bayerischen Akademie der Helléniques 26: 1-27 Wissenschaften 7: 371-464 McKenna M.C. & Bell, S.K. 1997 - Classification of Sakellariou-Mane, H., Psilovikos, A. & Koufos, G. D., mammals above the species level - Columbia 1979 - Contribution to the study of Villafranchian in University Press, New York N. Chalkidiki - Scientific Annals of the Faculty of Melentis, J.K., 1968 - Die Pikermifauna von Physics and Mathematics, Aristotelian Univercity of Halmyropotamos (Euboea - Griechenland) 1. Teil: Thessaloniki 19: 279-296 Odontology and Craniology - Annales Géologiques Schlosser, M., 1921 - Die Hipparionenfauna von Veles in des Pays Helléniques 19: 283-341 Mazedonien - Abhandlungen der Bayerischen Melentis, J.K., 1969 - Die Pikermifauna von Akademie der Wissenschaften, 29 (4): 1-55 Halmyropotamos (Euboea - Griechenland) 2. Teil: Schmidt-Kittler, N., Bruijn de, H. & Doukas, C., 1995 - Osteologie - Annales Géologiques des Pays The vertebrate locality Maramena (Macedonia, Helléniques 21: 217-306 Greece) at the Turolian/Ruscinian boundary Mitzopoulos, M.K., 1960 - Die Hipparionfauna von (Neogene). 1, General Introduction - Münchner Tanagra bei Theben - Annales Géologiques des Pays Geowissenschaftliche Abhandlungen A 28: 9-18 Helléniques 12: 301-314 Sickenberg, O. 1968 - Die Pleistozänen Knochenbrezien Osborn, H.F., 1900 - Phylogeny of the rhinoceroses of von Volax (Griech.-Mazedonien) - Geologisches Europe - Bulletin of the American Museum of Jahrbuch 85: 33-54 Natural History 12: 229-267 Sickenberg, O., 1972 - Ein Unterkiefer des Caniden Paraskevaidis E., 1977 - Säugetierreste aus Griechenland Nyctereutes donnezani (Dep.) aus der Umgebung von - Proceedings of the VIth Colloquium on the Geology Saloniki (Griech. Mazedonien) und seine biostrati- of the Aegean Region 3: 1143-1154 graphische Bedeutung - Annalen des Naturhistorischen Pavlow, M.W., 1913 - Mammifères tertiaires de la Museums in Wien 76: 499-513 Nouvelle Russie, 1. Partie: Artiodactyla, Solounias, N., 1981a - The Turolian fauna from the Perissodactyla (Aceratherium kowalevskii n.s.) - Island of Samos, Greece - with special emphasis on Nouveaux Mémoires de la Société impériale des the hyaenids and the bovids - Contributions to Naturalistes de Moscou 17: 1-68 Vertebrate Evolution 6: 1-232 Prothero, D.R., Manning, E. & Hanson, B.C., 1986 - The Solounias, N., 1981b - Mammalian fossils of Samos and phylogeny of the Rhinocerotoidea (Mammalia, Pikermi. Part 2. Resurrection of a classic Turolian Perissodactyla) - Zoological Journal of the Linnean fauna - Annals of Carnegie Museum 50: 231-270 Society 87: 341-366 Steininger, F.F., Berggren, W.A., Kent, R.L., Bernor, Prothero, D.R. & Schoch, R. M., 1989 - Classification of R.L., Sen, S. & Agusti, J., 1996 - Circum- the Perissodactyla - in: Prothero, D.R. & Schoch, Mediterranean Neogene (Miocene and Pliocene) R.M. (eds.) - The evolution of Perissodactyls - marine-continental chronologic correlations of Oxford Monographs on Geology and Geophysics 15: European Mammal Units - in: Bernor, R. L., 530-537 Fahlbush, V. & Mittmann, H.-W. (eds.) - The Psarianos, P., 1958 - Neue Rhinocerotidenfunde aus dem Evolution of Western Eurasian Neogene Mammal Tertiär und Quertär von Mazedonien (Griechenland) - Faunas - pp. 7-46, Columbia University Press, New Proceedings of Athens Academy 33: 303-312 York Qiu, Z., Xie, J. & Yan, D., 1988 - A new chilothere skull Symeonidis, N.K., 1974 - Ein bemerkenswerter from Hezheng, Gansu, China - with special reference Wirbeltierfund aus dem Lignit von Atalanti (Fthiotis, to the Chinese "" - Scientia Sinica (B) Griechenland) - Annales Géologiques des Pays 31: 493-502 Helléniques 26: 306-313 Ringström, T.J., 1924 - Nashörner der Hipparion Fauna Symeonidis, N.K., 1992 - Fossil mammals of Lower Nord Chinas - Palaeontologia Sinica, Serie C 1 (4): Pleistocene (Villafranchian) age from the Sesklo 1-156 Basin (Volos) - Annales Géologiques des Pays Roth, J., & Wagner, A., 1854 - Die fossilen Helléniques 35: 1-41 [in Greek]

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Symeonidis, N.K., Theodorou, G.E., Schütt, H. & Mainz" 7: 51-58 Velitzelos, E., 1985-86 - Paleontological and strati Toula, F., 1906 - Das Gebiss und Reste der Nasenbeine graphical observations in the area of Achaea and von Rhinoceros (Ceratorhinus Osborn) hundschei- Aetolia-Acarnania (W. Greece) - Annales mensis - Abhandlungen der k.k. Geologischen Géologiques des Pays Helléniques 33: 329-365 Reichsanstalt, Wien 20 (2): 1-38 Symeonidis, N.K. & Vos de, J., 1976 - Großsäugerfunde Tsiskarishvili, G.V., 1987 - Late Tertiary rhinoceroses aus den Pleistozänen Spaltenfüllungen von (Rhinocerotidae) of the Caucasus - AN GruzSSR, Turkovunia in Athen - Annales Géologiques des Pays Gosudarstvennyy Muzey Gruzii. Izdatel'stvo Helléniques 28: 135-144 ''Metsnierba'', Tbilisi [in Russian] Syrides, G., 1990 - Lithostratigraphical, Biostarti- Van de Weerd, A., Reumer, J.W.F. & de Vos, J., 1982 - graphical and Paleogeographical study of the Pliocene mammals from Apolakkia Formation Neogene-Quaternary sentiments of Chalkidiki (Rhodes, Greece) - Proceedings of the Koninklijke Peninsula - PhD Thesis, University of Thessaloniki Nederlandse Akademie van Wetenschappen B 85: [in Greek] 89-112 Thenius, E., 1955 - Zur Kenntniss der Unterpliozänen Wagner, A., 1848 - Urweltliche Säugetiere-Überreste aus Diceros-Arten - Annalen des Naturhistorischen Griechenland - Abhandlungen der Bayerischen Museums in Wien 60: 202-209 Akademie der Wissenschaften 5: 335-378 Thenius, E., 1956 - Ueber das Vorkommen von Diceros Wagner, A., 1857 - Neue Beiträge zur Kenntniss der pachygnathus (Wagner) im Pannon (Unter-Pliozaen) fossilen Säugthier-Überreste von Pikermi - des Wiener Beckens - Neues Jahrbuch für Geologie Abhandlungen der Bayerischen Akademie der und Palaeontologie, Monatshefte 1: 35-39 Wissenschaften 8: 109-158 Theodorou, G.E. & Nicolaides, S.N., 1988 - Weber, M., 1904 - Über tertiäre Rhinocerotiden von der Stratigraphic horizons at the classic mammal locality Insel Samos I. - Bulletin de la Société imperériale des of Pikermi, Attica, Greece - Modern Geology 13: Naturalistes in Moscou 17: 477-501 177-181 Weber, M., 1905 - Über tertiäre Rhinocerotiden von der Theodorou, G.E., Roussiakis, S. & Athanasiou, A., 1995 Insel Samos II. - Bulletin de la Société imperériale - Contribution to the study of the terrestial neogene des Naturalistes in Moscou 18: 345-363 of Greece. Ardiodactyla and Rhinocerotidae from the Woodward, A.S., 1901 - On the bone-beds of Pikermi, Kerasia and Chalkoutsi localities - Romanian Journal Attica, and on similar deposits in northern Euboea - of Stratigraphy 76: 129-130 Geological Magazine 8 (4): 481-486 Tobien, H., 1968 - Paläontologischen Ausgrabungen nach jungtertiären Wirbeltieren auf der Insel Chios Received 24 July 2001 (Griechenland) und bei Maragheh (NW Iran) - Accepted 16 September 2002 Jahrbuch der Vereinigung "Freunde der Universität

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APPENDIX Neogene localities with rhinoceroses in Greece. (fl)= cited only in a faunal list (no description/specimens published). Chronostratigraphic units after Steininger et al. (1996).

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DEINSEA - ANNUAL OF THE NATURAL HISTORY MUSEUM ROTTERDAM P.O.Box 23452, NL-3001 KL Rotterdam The Netherlands

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