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Zeitschrift/Journal: Beiträge zur Paläontologie

Jahr/Year: 2009

Band/Volume: 31

Autor(en)/Author(s): Giaourtsakis Ioannis X., Koufos George D.

Artikel/Article: The Late Faunas of the Mytilinii Basin, Samos Island, : New Collection - 10. Chalicotheriidae 189-205 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Beitr. Paläont., 31:189-205, Wien 2009

The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection 10. Chalicotheriidae

by

Ioannis X. Giaourtsakis1* & George D. Koufos2*

G iaourtsakis, I.X. &K oufos, G.D., 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 10. Chalicotheriidae. — Beitr. Palaont., 31:189-205, Wien.

Abstract (Griechenland). Einige der postcranialen Elemente wer­ den zum ersten Mal aus dieser Fundstelle beschrieben. Die In the present study, we describe several postcranial neuen Funde werden mit dem bisher bekannten Material elements ofAncylotherium pentelicum (Chalicotheriidae, dieser A rt, sowie mit dem anderer Chalicotherien-Arten ) from the island of Samos (Greece), some verglichen. Die miozänen Fundstellen mit Chalicothe­ of which are recorded for the first time from this site. riinae- und Schizotheriinae-Funden aus dem östlichen The material is compared to the known hypodigm of the Mittelmeerraum und benachbarter Regionen werden , as well as with other . The known zusammengefasst und ausgewertet. Potenzielle paläo- Miocene record of the two subfamilies, ökologische Präferenzen beider Unterfamilien werden and Schizotheriinae, in the Eastern kurz besprochen. Die Anwesenheit des Schizotheriinen Mediterranean and adjacent regions is briefly reviewed A. pentelicum und das Fehlen von Chalicotheriinae in den and evaluated. The potential paleoecological preferences turolischen Wirbeltierfaunen von Samos und Anatolien ot both subfamilies are briefly discussed. The presence of deuten auf die Verbreitung von mehr offenen Lebensräu­ the schizotheriine A. pentelicum and the absence of any men in der Umgebung dieser Fundstellen hin, anders als chalicotheriine species in the Turolian faunal assemblages im Vergleich zum kontinentalen Griechenland, FYR of of Samos and Anatolia indicate the expansion of more Macedonia und Bulgarien. open habitats in these localities compared to continental Greece, FYR of Macedonia and Bulgaria. Schlüsselworte: Turolium, Samos, Griechenland, Mam­ malia, Perissodactyla, Chalicotheriidae, Systematik. Keywords: Turolian, Samos, Greece, Mammalia, Peris- sodactyla, Chalicotheriidae, Systematics. 1. Introduction

Zusammenfassung pentelicum is a characteristic but rare element of the Late Miocene faunas in the Eastern Mediterranean In der vorliegenden Untersuchung beschreiben wir neu­ and adjacent regions. The first specimens from the island of es postcraniales Material von Ancylotherium pentelicum Samos were mentioned in a short report byM ajor (1888). (Chalicotheriidae, Schizotheriinae) von der Insel Samos A few years later,M ajor (1894) recorded several specimens of A. pentelicum in the collections of M CGL, Lausanne, and drew attention to an almost complete forelimb be­ longing to one individual. Schlosser (1921) mentioned Dipl.-Geol. IoannisX . G iaourtsakis , Ludwig-Maximilians- the presence ofA. pentelicum specimens among the Samos University of Munich, Department of Geo- and Environmen­ collection of the BSPG in Munich. D ietrich (1928:Abb. tal Sciences, Section of Paleontology, Richard-Wagner-Str. 1) illustrated a left juvenile mandibular fragment from 10, D-80333 Munich, Germany, e-mail: i.giaourtsakis@lrz. the collections of the MNHB in Berlin. Schaub (1943) tmi-muenchen.de documented in detail the forelimb material of the M CGL reported by M ajor (1894) together with some additional 2) Prof. George D. K o u f o s , Aristotle University ofThessaloniki. specimens from BSPG, MNHN and BMNH. Schaub’s Department of Geology, Laboratory of Geology and Palaeon- (1943) analysis on the osteology and functional anatomy tology, GR-54124 Thessaloniki, Greece, e-mail: georgeko@ of the forelimb ofA. pentelicum has been used ever since as geo.auth.gr; [email protected] a standard reference for numerous subsequent studies on ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

190 Beitr. Paläont., 31, Wien, 2009

chalicotheres. Recently, G eraads et al. (2006) recorded ing institutions: Athens Museum of Paleontology and several upper deciduous tooth series from Samos housed Geology, University of Athens (AMPG); Laboratory of at the collections of M CG L and illustrated two of them Geology and Palaeontology, University of Thessaloniki (Geraads et al., 2006:P1. 1, figs. 2, 3). In the present (LGPUT); Bayerische Staatssammlung für Paläontologie study, we describe new material of postcranial elements und Geologie, München (BSPG); Staatliches Museum für of A. pentelicum from the island of Samos. In particular, Naturkunde, Stuttgart (SMNS); Hessisches Landesmu­ elements of the hind limb are recorded for the first time seum, Darmstadt (HLMD); Museum für Naturkunde der from this site. The studied material was unearthed from Humboldt Universität zu Berlin (MNHB); Musée Can­ two localities, Mytilinii-3 (MYT) and Mytilinii-1 (MTL). tonal de Géologie, Lausanne (MCGL); Naturhistorisches The locality M Y T is situated in Potamies ravine and is Museum, Wien (NHMW ); Muséum National d’Histoire dated to early middle Turolian (early MN 12) at ~7.3 Naturelle, Paris (MNHN); British Museum of Natural Ma. The locality M TL is located in Adrianos ravine and History (= Natural History Museum), London (BMNH). consists of several fossiliferous sites; the chalicotheres have Comparisons with North American schizotheriines are been traced in the sites M TL A and MTLB, dated to the based primarily on C oombs (1978, 1979, 2004), as well uppermost middle Turolian at ~7.1 Ma. Detailed infor­ as Holland & Peterson (1914). mation about the geological and stratigraphical setting of the different fossiliferous sites of Samos is provided by Abbreviations: Kostopoulos et al. (this volume) and Koufos et al. (this L: length; DT: transversal diameter; DAP: anteroposterior (dor- volume). Historical data about the excavations in Samos soplantar) diameter; Prox: proximal; Dist: distal; Dia: diaphysis; is given by Koufos (this volume). Art: articular surface; max.: maximum.

The generaAncylotherium, , and are 2. Material and Methods abbreviated, while the generaAnisodon, Metaschizotherium, and Kalimantsia are not abbreviated. The specimens ofAncylotherium pentelicum described in the present article are stored in the collections of the Natural History Museum of the Aegean, Mytilinii, Samos 3. Systematic Palaeontology (NHMA). Methodology of postcranial measurements fol­ lows Roussiakis & T heodorou (2001), including a few Order Perissodactyla O wen, 1848 additional measurements. Measurements ranging 0-150 Family Chalicotheriidae G ill, 1872 mm were taken with a digital caliper precise to 0.01 mm Subfamily Schizotheriinae Holland and rounded to the nearest 0.1 mm. For larger measure­ & P eterson, 1914 ments, a vernier caliper with a precision of 0.1 mm was applied. A ll measurements are given in millimeters (mm). Ancylotherium, G audry, 1863 For specimens from Pikermi and Halmyropotamos, the values o f Roussiakis & T heodorou (2001) were in most Ancylotherium pentelicum (Gaudry & Lartet, 1856) cases preserved, as no significant interobserver difference (Plates 1, 2) was found. In the present study, Ancylotherium and Metaschizoth- Localities: Mytilinii-1 A, IB, (MTLA, MTLB), Adri­ erium are treated as two separate genera(K oenigswald, anos ravine, Mytilinii Basin, Samos, Greece; Mytili­ 1932; Butler, 1965; Roussiakis & T heodorou, 2001; nii-3 (MYT), Potamies ravine, Mytilinii Basin, Samos, G eraads et al., 2007). A subgeneric treatment between Greece. Ancylotherium and Metaschizotherium, as suggested by Age: Middle Turolian, MN 12 (late Miocene). Zapfe (1967) and C oombs (1974, 1989), may be equally M ytilinii-lA (M TLA, B): ~7.1Ma. reasonable. However, since the available record ofMetas­ Mytilinii-3 (MYT): ~7.3 Ma. chizotherium limited at present, and no cranial and suf­ Material: ficient postcranial evidence has been reported, we prefer MTLA: Right Mt-III, MTLA-277; ungual phalanx, to keep them separate at generic level. A discussion on M TLA-485. the inapplicability of other generic names{Macrotherium, MTLB: Left radius and ulna, M TLB-173; right radius Colodus, ), previously used forAncylotherium and ulna, M TLB-328; left femur, MTLB-338. pentelicum, is provided by Roussiakis & Theodorou MYT: Proximal phalanx, MYT-15; right Mt-III, MYT- (2001). Systematic recommendations for Asian and North 64. American Schizotheriinae follow C oombs (1989, 2004). Measurements: The measurements are given in Tables Chalicotheriinae nomenclature and follow the 1-3. recent analysis byA nquentin et al. (2007), but for dif­ ferent perspectives compare alsoB onis et al. (1995) and Description: G eraads et al. (2001, 2006). Radius and ulna. Radii and ulnae of A. pentelicum are Comparisons with Eurasian and African chalicotheres presently known from Pikermi and Samos.G audry (1863) have been carried out in the collections of the follow­ described a pair from Pikermi, which is housed in the col- ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

G iaourtsakis, I.X. Sc K oufos, G.D., Chalicotheriidae. 191

Samos Pikermi NHMA NHMA MCGL AMPG AMPG MNHN M TLB-173 M TLB-328 Schaub,1943 1972/22 PG 95/522 PIK 3549a R-L — ca. 565.0 557.0 519.3 521.0 556.0 U-L — — 703.0 661.0 — ca. 660.0 R-DT prox. max 154.0 156.0 157.0 141.0 159.0 148.0 R-DT prox. art 139.2 140.0 137.0 125.0 140.8 132.0 U-DT prox. art 135.2 134.5 127.0 126.4 131.0 (128.0) U-H scav 82.5 80.5 76.0 72.0 81.9 — U-L olecranon — — — 173.0 — — U-L olecr. + inc. — — 194.0 199.0 — — R-DT dia. min 62.8 — 59.0 (54.0) — 59.3 R-DAP dia. min 54.1 — — (57.0) — 56.4 DT dist. max — 193.0 199.0 193.2 199.2 194.0 DT dist. art — 166.0 162.0 164.0 165.2 167.0 R-DT dist. art — 124.8 123.0 118.0 128.0 124.3 U-DT dist. art — (S0.0) 52.0 (52.0) 48.0 48.4 DAP dist. max — 98.2 — — 97.8 104.8

Table 1: Comparative measurements (in mm) of radii and ulnae ofAncylotherium pentelicum (G a u d r y & L a r t e t , 1856) from the Eastern Mediterranean.

lections ofMNHN in Paris. Roussiakis & T heodorou the ulna, which is roughly divided into three confluent por­ (2001) documented seven additional specimens from tions. The medial portion extends along the caudal border Pikermi in the collections of AM P G in Athens. Schaub of the medial concavity of the articular fovea. It faces (1943) documented a complete radius and ulna from Samos caudally and is rather flattened transversally. A t the level stored in the collections of MCGL, Lausanne. During the where the medial and lateral concavities of the articular recent excavations on Samos, two new pairs have been fovea join, it bends proximally and blends with the median recovered from the locality MTLB. M TLB-173 is a left portion. The median portion has a moderately depressed adult specimen, missing the distal epiphyses and part of surface and faces proximally, like the two concavities of the distal shaft of both bones, and the olecranon of the the articular fovea. Approximately at the midpoint of the ulna (PI. 1, Fig. 3). MTLB-328 is a right adult specimen. It caudal border of the lateral concavity, the median portion retains the distal epiphysis, but is preserved in two portions of the articular surface for the ulna blends with the lateral (PI. 1, Figs. 1, 2). A segment of missing bone in the middle one. The lateral portion of the articular surface for the ulna of the diaphysis does not allow a secure contact between faces caudolaterally and is slightly concave transversally. the proximal and distal portions, but the total length of None of the ulnae preserves the olecranon. The anconeal the radius could be approximately estimated; the olecranon process of the ulna is strong and prominent. Below the of the ulna is not preserved. The state of preservation and anconeal process, the trochlear notch for the articulation the texture ofM TLB-173 and M TLB-328 are somewhat with the humeral trochlea is arched. The medial and lateral different, but the metrical and morphological similarities, coronoid processes are asymmetrical. as well as their taphonomic proximity, may indicate that The shaft of the radius is long and appears quite slender both specimens could belong to the same individual. with respect to the broadened proximal and distal epi­ In proximal view, the articular fovea of the radius that physes. A t mid-shaft, the transverse and anteroposterior articulates with the humeral trochlea is divided into two (craniocaudal) diameters become nearly subequal. The asymmetrical concavities, which are separated by an in­ ulna is characterized by a more slender shaft with a sub- cipient, concave ridge at the level of the radial coronoid triangular cross-section. The interosseous space between process. The lateral suboval concavity is larger and deeper radius and ulna is small and was probably filled by a dense than the subcircular medial one. On the cranial side of interosseous membrane during lifetime, as indicated by the the proximal epiphysis of the radius, there is no prominent presence of bilateral coarse surfaces. radial tuberosity, but a coarse surface for the attachment The distal epiphyses of the radius and ulna are co-ossified, of the biceps tendon. In lateral aspect, the tuberosity for prohibiting any relative movements between the two the collateral ligament is particularly strong and promi­ bones. The distal articular facets for the scaphoid, luna- nent. In caudal aspect, the arrangement between the tum and pyramidal form a continuous arched cavity (PI. articular facets for the ulna cannot be observed, because 1, Fig. 2). Contrary to extant perissodactyls, all articular both specimens are preserved in natural contact. Isolated surfaces for the carpals are quite smooth; none is mark­ radii from Pikermi (AM PG: PG 95/522, PG 95/27, PG edly projecting or depressed. This adaptation enables 95/549) show a transversally widened articular surface for enhanced flexion capabilities in the joint, as suggested by ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

192 Beitr. Palaont., 31, Wien, 2009

Schaub (1943). On the lateral half of the cranial surface the third trochanter is projecting more prominently than of the distal epiphysis of the radius, a pronounced massive the lesser one, a condition quite similar to the extant peris­ tuberosity is developed, most likely for the accommodation sodactyls (Coombs, 1978:Fig. 13b; 1979: Fig. 17b). of strong ligamentous attachments to support the flexion The distal epiphysis of the femur is partly broken off and and extension of the hand. The medial (radial) and the missing its lateral half. In caudal view, the well-preserved lateral (ulnar) styloid processes are formed as rounded medial condyle is large and ovoid (PI. 1, Fig. 5). The eminences; the medial one is stronger and extends more intercondyloid fossa is moderately depressed and has a distally than the lateral one. coarse surface. It served as the attachment point for the Femur. Very few femora ofA. pentelicum are known. anterior cruciate ligament that prevents inward rotation G audry (1863) briefly described a juvenile femur from and forward displacement of the tibia towards the femur. Pikermi. Schlosser (1921) referred to another specimen A distinct depression in the inner surface of the medial from Veles. Recently,R oussiakis 8 c Theodorou (2001) femoral condyle probably served as the attachment point documented a complete femur from Pikermi, as well as for the posterior cruciate ligament that prevents backward two fragmentary distal portions. MTLB-338 is the first displacement of the tibia away from the femur. In medial specimen reported from Samos and is a moderately well- view, the medial epicondyle forms a large, rough, convex preserved left femur (PI. 1, Figs. 4-6). eminence, to which the tibial collateral ligament of the The proximal epiphysis, which the femoral head and knee-joint must have been attached. In cranial view, the the greater trochanter, is completely missing. Due to its articular surface for the patella retains only a small portion condition, the only measurable dimensions of the specimen of the medial condyle, which is rather short. Complete are the minimum transversal and anteroposterior diam­ specimens from Pikermi (Roussiakis 8 c Theodorou, eters of the shaft (DTdia. = 95.4; DAPdia. = 62.0), which 2001:Fig. 26) have a shallow and rather symmetrical are similar to the specimens from Pikermi (Roussiakis patellar facet that distinguishes the distal epiphysis of 8 c Theodorou, 2001:Tab. 11). The shaft of the femur ancylothere femurs from sympatric rhinocerotids and gi- is rather flattened and broad; its transverse diameter is raffids (isolated distal fragments of femurs are frequent in always greater than the anteroposterior one throughout collections). In the and giraffes, the patellar its length. The shaft is broader in its proximal half, due surface is deeper and markedly asymmetrical, because its to the presence of the lesser and third trochanters. Distal medial condyle is markedly larger than the lateral one. to the third trochanter, the shaft narrows, though less Third Metatarsal. Third metatarsals ofA. pentelicum have abruptly than in the North American Moropus elatus and been described from Pikermi (Gaudry, 1883; Zapfe, Tylocephalonyx skinneri (Coombs, 1978, 1979), and then 1979; Roussiakis 8 cTheodorou, 2001), Veles (Schloss­ widens again towards the distal epiphysis. The cranial er, 1921), and Salihpa$alar (Sara

G iaourtsakis, IX. Sc Koufos, G.D., Chalicotheriidae. 193

Samos Pikermi Veles Salihpasalar NHMA NHMA AMPG AMPG MNHN Schlosser, Saraç et al., M TLA-227 M YT-64 PG 95/513 PG 95/515 PIK 3540d 1921 2002 Lmax — — 169.0 175.0 171.0 170.0 140.0 DT prox 57.6 60.7 58.5 62.9 59.3 60.0 46.0 DAP prox 52.5 53.0 60.3 54.1 56.7 — 43.0 DT dia 46.2 48.8 47.9 47.2 46.5 — 39.0 DAP dia 28.8 27.5 28.8 28.7 28.0 — 29.0 DT dist.max — — 59.3 (60.5) 57.2 59.0 — DAP dist.art — — 64.8 64.5 60.8 — 54.0

Table 2: Comparative measurements (in mm) of Mt-III ofAncylotheriutnpentelicum (G a u d r y &c L a r t e t , 1856) from the Eastern Mediterranean. narrow Mt-II facet on the medial side of the Mt-III from from M. elatus by the fact that the dorsal and plantar Mt-IV Salihpaçalar. C oombs (1978:Figs. 17a, 19a, 27e; 1979: Fig. facets are broadly confluent (Coombs 2004:Fig. 15.6b), a 23a) documented the presence of a bipartite Mt-II facet condition also observed in several terminal taxa of dif­ for M. elatus, M. hollandi, and T. skinneri, consisting of a ferent rhinocerotid lineages Brachypotherium,( Coelodonta dorsal and a plantar portion that may be either separated antiquitatis). In specimens of T. skinneri from the Sheep by a shallow groove or variably connected by a narrow Creek Formation in , including the holotype strip of articular surface. A n Mt-III from northern Eagle skeleton, the dorsal Mt-IV facet is concave(C oombs, 1979: County, , questionably referred toTylocephalonyx 23b, 24d), but not as deeply as inA. pentelicum. It usually by C oombs (1979), exhibits a single, narrow, band-like remains separated from the smaller plantar facet by a shal­ Mt-II articular facet (Coombs, 1979:Fig. 29f). A similar low groove, as in M. elatus. The proximal Mt-III fragment variation in the arrangement of the articular surface for of a partial skeleton from the Split Rock vertebrate fauna in the Mt-II is documented among chalicotheriine taxa by , which can be referred toT. skinneri, differs from Butler (1965) and Zapfe (1979). In lateral view, the the Sheep Creek specimens in having the concave dorsal articular surface for the M t-IV consist of a large, deeply and the flattened plantar portions of the M t-IV articular concave, dorsal portion that occupies most of the lateral surface confluent (Coombs, 1979:Fig. 28f). In the latter side of the proximal end, and a smaller, flattened, plantar specimen, a notable articular facet for the cuboid is also portion that is limited to the lateroplantar corner and is present. Other postcranial elements from the Split Rock to some extent obscured by postmortem abrasion (PI. 2, partial skeleton also exhibit some morphological variation Fig. 2, 5). Better preserved specimens from Pikermi that with respect to the typical population ofT. skinneri from we have examined (AM PG: PG 95/513, PG 95/515, PG Sheep Creek, indicating that it may represent a different 95/551), indicate that these two portions are either in wide subspecies (Coombs, 1979). The development of a large contact, or perhaps slightly separated by a very narrow and deeply concave dorsal M t-IV facet on the Mt-III of incipient groove. The large, deeply concave, facet is evident A. pentelicum must be a derived feature that enhances the in the Mt-III from Salihpajalar figured by Saraç et al. interlocking between the third and fourth metatarsals, (2002:Fig. 3.3b), but the status of the plantar flattened thus more stabilizing the tarsometatarsal joint and reduc­ portion is not clear in the illustration. Most likely, based ing the abductive and adductive movements in-between on this specimen, G eraads et al. (2007:464) argued that these metapodials. only a single dorsal facet between Mt-III and Mt-IV occurs The shaft of the bone is straight and dorsoplantarly in A. pentelicum, a statement that must be amended based compressed with respect to the proximal epiphysis (PI. on the present evidence. G uérin &c Pickford (2005) 2, Figs. 2, 5). In dorsal view, the most striking feature reported “un bord lateral déprimé en dessous de la facette is a marked longitudinal depression on the medial side articulaire répondant au Mt IV” for the Mt-III of the of the dorsal surface of the shaft (PI. 2, Figs. 1, 4). A African Ancylotherium cheboitense from the terminal Late similar depression is also developed on the lateral side Miocene locality of Cheboit, Baringo District, , of the dorsal surface of the shaft, but it is less markedly but they did not discuss or depict the arrangement of the expressed. These depressions are well-developed in the Mt-IV facets. In the North AmericanM. elatus, the dorsal adult specimens from Pikermi, AM PG : PG 95/513, PG Mt-IV facet is flat to faintly concave. The smaller plantar 95/515 and MNHN: PIK 3540d (compare also G audry, facet is better delimited than inA. pentelicum. Both Mt-IV 1883:P1.20, Fig. 3; Roussiakis & T heodorou, 2001:Fig. facets are slightly slanted towards the center of the lateral 29.3). In the juvenile specimen AM PG : PG 95/551, both surface and remain clearly distinct from one another; either depressions are faintly expressed. In plantar view, the separated by a marked groove, or by being directly adjacent. surface of the shaft is flattened to slightly depressed. The This morphology of the M t-IV facets inM. elatus is appar­ medial and lateral aspects of the shaft show longitudinal ently more primitive and quite similar to the arrangement rugose surfaces for the attachment of the interosseus liga­ observed in and rhinocerotids. M. merriami differs ments connecting the Mt-III with the adjacent metatarsals. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

194 Beitr. Paläont., 31, Wien, 2009

Samos Pikermi Halmyropotamos Pinaryaka NHMA Schaub AMPG AMPG AMPG AMPG Saraç et al., M YT-15 1943 PG 95/523 PG 95/524 1969/128 1969/151 2002 L max 127.0 114.0 116.5 125.4 121.0 115.5 125.0 D T prox. max 75.6 79.0 74.7 76.3 69.4 69.7 75.0 D AP prox. max 59.2 — 57.6 64,0 (65.2) 59.9 62.0 D T prox. art 65.2 — (61.8) (63.6) (61.9) 61.6 — DAP prox. art 49.6 — (43.6) (44.7) (47.8) (46.0) — DT dist. max 57.5 56.0 54.3 58.0 — 52.6 — DT dist. art 47.7 — 49.2 51.8 — 45.2 —

Table 3: Comparative measurements (in mm) of large-sized proximal phalanges ofAncylotherium pentelicum (G a u d r y Ôc L a r t e t , 1856) from the Eastern Mediterranean that may be potentially attributed to the third digit of the manus. For dimension of smaller- sized proximal phalanges also compare Ro u s s ia k is ôc T h e o d o r o u (2001:Tab. 21).

Phalanges. Due to their peculiar and easily recogniz­ ridges form two coarse surfaces separated by a shallow able morphology, phalanges ofA. pentelicum are among but broad longitudinal depression. The distal articulation the most common findings of this species and have been for the middle phalanx is displaced almost entirely to the reported from almost all of its known localities. Particu­ palmar aspect of the bone. Its bilateral keels and median larly remarkable is the duplex bone (fused proximal and trochlear depression are in line with the marginal ridges middle phalanges) of the enlarged second digit of the and the longitudinal depression of the shaft. The bilateral manus, which is followed by a massive ungual phalanx. keels appear somewhat asymmetrical, but their detailed The presence of smaller duplexes, which belong to the morphology is obscured by surface abrasion. second digit of the pes, has been documented inM etas- MTLA-485 is a well-preserved ungual phalanx (PI. 2, chizotherium , Moropus and Tylocephalonyx, but remains Figs. 7, 8 ). The specimen maybe tentatively assigned to the currently undocumented and prospective inA. pentelicum pes, due to its smaller size with respect to the specimens (Coombs & Rothschild, 1999). The precise position of assigned to the manus by Schaub (1943). Its maximum isolated phalanges from other digits is difficult to be as­ length is circa 88.0 mm; the length of the plantar surface certained. Generally, the larger specimens are tentatively is 67.7 mm; the maximum transversal diameter is 42.5 assigned to the manus, and the smaller ones to the pes, mm; the transversal diameter of the articular surface is primarily based on the substantial proportional differ­ 28.2 mm, and the anteroposterior one 48.8 mm. The tip ences observed between metacarpals and metatarsals. The of the dorsal process is broken off. The non-articular sur­ topographical definition of intermediate specimens cannot faces of the bone are very coarse, bearing multiple small be granted, also taking into account the potential pres­ furrows. The articular surface for the medial phalanx is ence of considerable sexual dimorphism in chalicothere curved in an arch and consists of two gently concave facets taxa (Coombs, 1975, 1978). Functional interpretations separated by a median ridge. The claws are sharp, slightly for the phalangeal adaptations of chalicotheres have been asymmetrical, and deeply cloven dorsally, but not plantarly thoroughly debated by K oenigswald (1932), Schaub (PI. 2, Figs. 7, 8 ). The subungual process is evident as a (1943), B orissiak (1945), Butler (1965), Zapfe, (1979), coarse tubercle on the plantar side and is not particularly and C oombs (1983, 1989). strong, such as in the ungual of the enlarged digit II of M YT-15 is a well-preserved proximal phalanx of an adult the manus (Roussiakis & T heodorou, 2001:Fig. 30.6; individual (PI. 2, Fig. 9). The size of the bone, in particu­ Sara9 et al., 2002:Fig. 4.4). C oombs (1978) noted that lar its length, indicates that it most probably represents a pes unguals of M. elatus have a very reduced subungual proximal phalanx of the digit III of the manus (Tab. 3). process with respect to the manus. The rest of the plantar Morphologically, M YT-15 does not differ from the known surface is extremely coarse. specimens of the species(S chaub, 1943; M elentis, 1969a, 1969b; Roussiakis Sc Theodorou, 2001; Sara£ et al., 2002). The articular facet for the corresponding metapod 4. Discussion is ovally heart-shaped, bearing a small but marked incision in the middle of its proximoplantar border. Its articular Within the family Chalicotheriidae, two well-documented surface is gently concave and proximodorsally oriented, subfamilies can be distinguished by a unique combination occupying almost one third of the bone in dorsal view. of several features (Coombs, 1989). Chalicotheriinae Above the proximoplantar border of the metapodial facet, retain a conservative low-crowned dentition, but attain the two small bilateral tubercles are slightly asymmetri­ highly modified postcranials with remarkably shortened cal and separated by a notch. Distally to the metapodial hindlimbs. Schizotheriinae retain more conservative facet, the shaft of the phalanx narrows. In dorsal view, the postcranial proportions and morphology, but they develop shaft is transversally convex and longitudinally slightly a more derived dentition with more elongated and higher- concave. In palmar view, the medial and lateral marginal crowned molars. Based on the available evidence, the place ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Giaourtsakis, I.X. &K oufos, G.D., Chalicotheriidae. 195

Figure 1: Late Miocene localitieswithAncylotheriumpentelicum (Gaudry ÔcLartet, 1856) in the Eastern Mediterranean and adjacent regions. 1. Strumyani; 2. Gorna Sushitsa; 3. Hadjidimovo; 4. Kalimantsi ; 5. Veles; 6. Pentalophos; 7. Kerassia; 8. Halmyropotamos; 9. Pikermi; 10. Samos; 11. Karaburun; 12. Giilpmar; 13. Kemiklitepe; 14. Salihpajalar; 15. Konya-Kiziloren; 16. Pinaryaka; 17. Akka^dagi; 18. Maragheh; 19. Novoukrainka.

and time of divergence of the Chalicotheriinae from the from the Seyitomer coal basin (Kiitahya) to M oropus elatus, Schizotheriinae cannot be precisely determined (C oom bs, primarily due to the presence of an articular facet for the 1989). In the present discussion, we will focus on the East­ ectocuneiform. However, based solely on this evidence, ern Mediterranean schizotheriine record and lay particular the major extension of the geographic range of the North emphasis on its well-documented late Miocene representa­ American species seems ambiguous, given that Mt-IVs of tive Ancylotherium pentelicum . We will also briefly discuss M etaschizotherium are not well known (R innert, 1956). the Eastern Mediterranean chalicotheriine record, as its The last and most recent reference to a middle Miocene synchronic and sometimes sympatric occurrence may offer schizotheriine in the Eastern Mediterranean is comprised significant palaeoecological and biogeographical impli­ of a quite fragmentary set of dental and postcranial remains cations (T h enius, 1953; Z a p f e , 1979; C oo m bs, 1989; from the coal-mine Bursa-Orhaneli (Burmu) in Western G eraad s et ah, 2001; S chulz et ah, 2007). Anatolia (O nar & Y ild iz, 2005). The most prominent specimens, a well-preserved duplex and the proximal half of a second metacarpal, demonstrate characteristic schizother­ 4.1. Middle Miocene Record iine features (O nar & Y ild iz, 2005:Fig. 3). These authors considered the duplex as belonging to the second digit of the The oldest remains of a schizotheriine chalicothere in manus. The reported dimensions of the Bursa-Orhaneli du­ the Eastern Mediterranean were reported from the late plex (O nar & Y ildiz, 2005:Tab. 1) are significantly smaller Orleanian/early Astaracian mammalian fauna of Kultak than the manus duplexes of A. pentelicum. In particular, (Milas-Mugla), in southwestern . A single left its total length is reported to be less than half the length proximal phalanx referable to digit IV of the manus was documented for A. pentelicum specimens (Sch au b, 1943; recovered (K aya et ah, 2001:Fig. 3.2a, b) and attributed R ou ssiakis & T heodorou, 2001; Sa r a

196 Beitr. Palaont., 31, Wien, 2009

rectly attributed to the pes byC oombs (1974), have a very crowned molars with respect to its potential Middle M i­ similar size and morphology to the specimen from Bursa- ocene forerunnerMetaschizotherium fraasi. The inadequate Orhaneli. A schizotheriine duplex from the Aragonian site Vallesian record does not yet clarify yet the exact evolu­ of Bunol (Valencia, Spain) assigned to the digit II of the tionary relationship between these two species(C oombs, pes, also has similar proportions to the Anatolian specimen 1989). The oldest evidence of A. pentelicum is known from (Belinchon 8 cM ontoya, 1990:Lam. 1, Figs. 7a, b). The the Vallesian locality of Pentalophos in Greece, but the size difference observed between well-documented manus material is scanty (Bonis et al., 1999). During the Turo­ and pes duplexes ofMoropus (Coombs, 1978:Figs. 19c, d) lian, the geographic range ofA. pentelicum covers the entire and Tylocephalonyx (Coombs, 1979:Fig. 25) also justify the Subparatethyan (Bernor, 1983, 1984) or Greco-Iranian smaller size of the Bursa-Orhaneli duplex. (Bonis et al., 1992a, 1992b) zoogeographic province All three middle Miocene Anatolian schizotheriine (Fig. 1). It has been firmly documented in the localities of references seem to be related withMetaschizotherium, but Pikermi (W agner, 1857; H ensel, 1862; G audry, 1863; additional material is necessary to allow a reliable deter­ Thenius, 1953; Roussiakis 8 cTheodorou, 2001), Samos mination. Metaschizotherium is a rare element in middle (M ajor, 1894; Schaub, 1943), Halmyropotamos (M e- Miocene faunas of Western and Central , where it lentis, 1969a, 1969b), Kerassia (Theodorou et al., 2004), probably includes more than one species(K oenigswald, and Thermopigi (Geraads et al., 2007) in Greece; Veles 1932; Zapfe, 1967, 1974, 1979; C oombs, 1974, 1989). in FYR of Macedonia (Schlosser, 1921; G arevski, 1974; In particular, the ongoing study of the significant M eta­ G arevski & Zapfe, 1983); Gorna Sushitsa, Kalimantsi, schizotherium material from Sandelzhausen, Germany, Hadjidimovo, and Strumyani-1 in Bulgaria (Bakalov, (M. C oombs, in prep.) will further clarify its systematic 1955; Bakalov & Nikolov, 1962; G eraads et al., 2001, affinities with respect to its potential late Miocene suc­ 2006); Giilpinar (Kaya, 1986), Kemiklitepe (Kaya, 1988; cessor Ancylotherium pentelicum. W e must note that a Sen, 1994), Salihpa^alar and Pinaryaka (SARA 9 et al., schizotheriine DP3 from the early Turolian Bulgarian 2002), Karaburun (Kaya et al., 2005), Konya-Kiziloren locality of Kalimantsi-1, initially described as an M2 and Akka^dagi (Sara£ 8 c Sen, 2005) in Turkey; and of Metaschizotherium fraasi by Nikolov (1972:Fig. 1), is Maragheh in Iran (M ecquenem, 1924; G eraads et al., properly referred toA. pentelicum by C oombs (1974) and 2006). The northern range ofA.pentelicum maybe slightly G eraads et al. (2001, 2006). extended by some specimens documented at Novoukrainka The available Middle Miocene record of chalicotheriines in Ukraine (Pidoplichko, 1959; K orotkevich, 1988), in the Eastern Mediterranean is presently as scanty as the and the eastern range by some unpublished specimens one of schizotheriines. The few recovered specimens from reported from Molayan, Afghanistan (Brunet et al., Anatolia were referred to as comparable withAnisodon 1984; C oombs, 1989). The Chinese schizotheriine taxa grande, a well-documented taxon from several synchronous Huanghotherium anlungense Tung, Huang 8 c Qiu, 1975, Western and Central European localities (Zapfe, 1979; from Shanxi Province and Gansuodon pingliangense W u A nquentin et al., 2007). Fortelius (1990) documented 8 c C hen, 1976, from Gansu Province, both based on a several isolated teeth from Pajalar, G eraads 8 c Sara£ few high-crowned and elongated molars, as well as an (2003) described a third metacarpal and some phalanges astragalus from Yushe (Bohlin, 1936), may be related with from Qandir, and Becker-Platen et al. (1975) reported Ancylotherium, but the available material is too scanty to its presence in Sofca and Catakbagyaka. Few dental determine precise affinities(C oombs, 1989). The same is specimens from the locality of Plevlja in Montenegro true for some schizotheriine specimens reported asAn­ were also referred to as comparable withAnisodon grande cylotherium aff. A. pentelicum from Spanish localities such (Petronijevic, 1957; Bonis et al., 1995). as Nombrevilla, Los Valles de Fuentiduena, and Concud Summarizing, the Middle Miocene record of chalicotheres (A lberdi et al., 1981; H eissig, 1999). remains still scanty and very incomplete in the Eastern Contrary to the schizotheriines, the Turolian chalico- Mediterranean. It appears to include representatives of theriine record is restricted only to the western margin both schizotheriines and chalicotheriines, but a reliable of the Greco-Iranian/Subparatethyan zoogeographic specific or even generic attribution is hazardous at present, province. Bonis et al. (1995) attributed a complete skull as additional material is required to establish unambigu­ from the locality of Dytiko 3, Greece, to a new species, ous systematic affinities with respect to the Western and Anisodon macedonicum. Chalicotheriine specimens from Central European Middle Miocene chalicothere taxa. Pikermi (Butler, 1965; Symeonidis, 1973) and Vathy- lakos (A rambourg 8 c P iveteau, 1929) in Greece, and Veles in FYR of Macedonia (Garevski 8 c Zapfe, 1983) 4.2. Late Miocene Record may be temporarily referred to asAnisodon sp., until more material becomes available for accurate specific affinities The available Late Miocene schizotheriine and chalicoth- (Bonis et al., 1995; A nquetin et al., 2007). G eraads eriine record in the Eastern Mediterranean and adjacent et al. (2001) described a well preserved cranium as a new regions is primarily restricted to the Turolian, and Valle- genus and species, Kalimantsia bulgarica, from the locality sian evidence remains extremely rare at present. of Kalimantsi, Bulgaria, and suggested that other Balkan Ancylotherium pentelicum is the typical Turolian schizoth­ chalicotheriines maybe included in this genus.A nquetin eriine, demonstrating significantly larger size and higher et al. (2007) argued that the skull ofKalimantsia shows ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

G iaourtsakis, I.X. &c Koufos, G.D., Chalicotheriidae. 197

several plesiomorphic features that suggest it should be excluded from the cladeAnisodon and noted that further and even with some grazers. The abrasive mesowear signa­ comparative studies are required to establish precise ture of the chalicotheriines was convincingly resolved as a generic affinities. Chalicotheriine specimens from the result of the consumption of twigs, barks, and branches in Bulgarian localities of Hadjidimovo, Gorna Sushitsa, addition to the leaf browsing, forming a dietary combina­ Kromidovo and Batishnitsa (Bakalov & N ikolov, 1962; tion similar to the extant “mabra” western tree hyrax,D en- Nikolov, 1975; G eraads et al., 2001,2006) are temporar­ drohyrax dorsalis. This interpretation is further supported ily best referred to as Chalicotheriinae indet. Given the by the extensive surfaces for the masticatory muscles on scant but evident presence of Chalicotheriinae in Anatolia the mandible of the chalicotheriines, and is in accordance during the Middle Miocene(F ortelius, 1990; G eraads with their shortened hindlimbs and protractible, clawed & SARAf, 2003), their striking absence from the well- forelimbs (Schulz et al., 2007). sampled Turolian localities of Samos (Greece), Turkey, of Schizotheriinae, in particular Ancylotherium, and Maragheh (Iran) is apparently related to provincial are commonly associated with more open woodlands. paleoenvironmental changes and constraints. Schizotheriines have higher-crowned and more elon­ gated molar teeth compared to chalicotheriines. Among , this is typically viewed as a selective response 5» Paleoecological and Biogeographical to a coarser diet (Koenigswald, 1932; Thenius, 1953; Remarks Butler, 1965; Zapfe, 1976,1979; C oombs, 1983,1989; H eissig, 1999). However, a mesowear analysis employed Dietary and paleoecological inferences of chalicotheres on Metaschizotherium by S chulz et al. (2007) suggested have been based on the morphological and physical a less abrasive diet with respect to the lower-crowned properties of their dentition, as well as on their peculiar chalicotheriines. The obtained mesowear signature clas­ postcranial adaptations (A bel, 1920; D ietrich, 1928; sified Metaschizotherium within the spectrum of extant Koenigswald, 1932; Schaub, 1943; Borissiak, 1945; intermediate feeders. Nevertheless, as demonstrated by Butler, 1965; Zapfe, 1976,1979; C oombs, 1983,1989; the example of the chalicotheriines that were plotted close Schultz et ah, 2007). to extant grazing antelopes, the mesowear signature alone Fossils of Chalicotheriinae are commonly associated with may be not sufficient to resolve the dietary preferences of denser woodland fauna assemblages. Chalicotheriines had the schizotheriines. The abrasive component in the diet of remarkably short hindlimbs that were capable of ­ chalicotheriines was attributed to twigs and barks, and a ing their body weight in an erect posture, resulting in a similar abrasive component might have contributed to the clinograde stance comparable to extant gorillas. Due to diet of schizotheriines, perhaps at a different ratio. A grass their low-crowned dentition, a primarily leaf browsing diet component in the diet of schizotheriines can be excluded. has been frequently suggested, assisted by the employment Schizotheriines developed more high-crowned teeth in of their elongated forelimbs with a protractable humerus comparison to the chalicotheriines, but their dentition and hooked clawed digits (Koenigswald, 1932; Zapfe, still remained functionally low-crowned brachydont (sensu 1976, 1979; C oombs, 1982, 1983, 1989; H eissig, 1999). Fortelius, 1985). The premolars are never noticeably A recent dental microwear study byS chulz et ah (2007) molariform, the occlusal surface of the molars is concave, supported the presence of a strong browsing component no cement is developed, the vertical styli on the ectoloph in the diet of the chalicotheriines, which is different are strong, their intermediate relief is high and the cusp from that observed in extant browsing ruminants. But a apexes are subjectively sharp or rounded, but never blunt. dental mesowear analysis employed by the same authors These morphological properties of the dentition indicate yielded a substantial abrasive component in the diet that two distinct chewing phases during the occlusal stroke, was similar in abrasiveness to that consumed by modern shearing and crushing, a combination associated with the grazing antelopes. Schulz et ah (2007) extensively dis­ cominuition of a primarily browsing diet (Fortelius, cussed the apparent inconsistency between the microwear 1985; Fortelius & Solounias, 2000). W e suggest that and meso-wear signals, combining the results of both the higher-crowned and more elongated molars of the methods. They concluded that the abrasive component, functionally brachydont dentition of the schizotheriines indicated by the mesowear signature, is not related to a reflect an increase in the quantity and abrasiveness of the grass component, due to the fact that the low numbers of browse components of their diet, as a response to the ex­ scratches documented by the microwear analysis exclude a pansion of more open and/or seasonal habitats and their substantial amount of graminoids. Owing to the very low nutritionally inferior forage. The less specialized postcra­ scratch density of the microwear signature, frugivory, in­ nial anatomy of the schizotheriines would also tolerate cluding hard seeded fruits was also excluded as the abrasive more evolutionary flexibility towards a modification of component. A comparison with the mesowear signature of their dietary requirements, in order to cope more efficiently some extant species classified as “minute abraded with the environmental changes than the chalicotheriines. brachydonts (mabra)” by Fortelius ScSolounias (2000) New postcranial adaptations, such as longer necks and provided a close resemblance. Similar to chalicotheriines, limbs, were evolved to assist these dietary adjustments. the mesowear signature of “mabra” species shares a high Schizotheriines were able to stand erect on the hindlimbs frequency of high occlusal reliefs with browsers, but also a while feeding, in order to increase the browsing height ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

198 Beitr. Palàont., 31, Wien, 2009

and to allow the forelimbs to hook branches within reach 7. References (Coombs, 1983:Fig. 9). Schaub (1943) suggested that the development of a massive forelimb inA. pentelicum might A bel, O., 1920. Studien über die Lebensweise von Chali- have been used in pulling down tree branches and other cotherium. — Acta Zoológica, 1:21-60, Stockholm. vegetation in a manner similar to the trunk and tusks of A lberdi, M.T., G insburg, L. 8 c M orales, J., 1981. extant elephants. In A. pentelicum, the toes of the manus Rhinocerotidae del yacimiento de los Valles de Fuenti- were held back against the dorsal sides of the metacarpals; dueña (Segovia). — Estudios Geológicos, 37:439-465, the elevation and retraction of the claws was interpreted Madrid. as an adaptation to avoid blunting during walking on A nquetin, J., A ntoine, P.-O., Tassy, P., 2007. Middle hard ground, as is the case in more open and arid habitats Miocene Chalicotheriinae (Mammalia, Perissodac- (Schaub, 1943; Butler, 1965) tyla) from France, with a discussion on chalicotheriine The relative occurrence of schizotheriines and chalicother- phylogeny. — Zoological Journal of the Linnean iines in the Turolian localities of the Eastern Mediterranean Society, 151:577-608, London. and adjacent regions seems thus to reflect environmentally- Bakalov, R, 1955. Chalicotheriidae in Siidwestbulgar- controlled provincial differences (Thenius, 1953; Zapfe, ien. — Izvestija Geologiceskogo Instituta Bolgarskoj 1979; C oombs, 1989). Localities in the western margin of Akademii Nauk, 3:151-159, Sofia. the Greco-Iranian/Subparatethyan zoogeographic province Bakalov, P. ôcNikolov, I., 1962. Les Fossiles de Bulgarie still provided niches with denser tree coverage and more X. Mammifères Tertiaires. — Académie des Sciences temperate conditions favored by the chalicotheriines. Their de Bulgarie, pp. 162, Sofia. absence from the central and eastern parts of the Greco- Becker-Platen, J.D., Sickenberg, O. 8 c Tobien, H., Iranian/Subparatethyan zoogeographic province (Samos, 1975. Vertebraten-Lokalfaunen der Türkei und Turkish localities, Maragheh) is associated with the more ihre Altersstellung. — Geologisches Jahrbuch, B, rapid expansion of open and dry habitats in these areas 15:45-100. (Koufos et al., this volume). On the other hand, the den­ B elinchón, M. 8 c M ontoya, R, 1990. Presencia de tal and postcranial adaptations of A. pentelicum allowed a Phyllotillon naricus PILGRIM, 1910 (Chalicother­ greater environmental tolerance and thus the ability to suc­ iidae, Perissadactyla, Mammalia) en el Aragoniense cessfully dwell in more open and perhaps seasonal habitats. de Buñol (Valencia). — Paleontología i Evolució, 23: Analogous evidence is provided by the relative distribution 171-180, Sabadell. of other herbivores in the region, including rhinocerotids, Bernor, R.L., 1983. Geochronology and zoogeographic equids, bovids, and giraffids (Koufos et al., this volume). relationships of Miocene Hominoidea. — [in:] Rus­ In the cases of sympatry, the dietary and habitual differ­ sell, L.C. 8c Robert, S.C., (eds.). New interpreta­ ences betweenAncylotherium and the chalicotheriines could tions of ape and human ancestry. — 21-64, New York suggest limited intraspecific competition, as they may have (Plenum Press). occupied different niches. Based on the available evidence, B ernor, R.L., 1984. A zoogeographic theater and the major biotic turnover at the Miocene/ bound­ biochronologic play: The time/biofacies phenomena ary led both subfamilies to disappear from the Eastern of Eurasian and African Miocene mammal prov­ Mediterranean and adjacent regions. inces. — Paléobiologie Continentale, 14:121-142, Montpellier. B ohlin, B., 1936. Notes on some remains of 6. Acknowledgements from China and Mongolia. — Bulletin of the Geological Society of China, 15(3):321-330, The excavations on Samos have been supported by the Pre­ Beijing. fecture of Samos and the “Konstantinos and Maria Zimalis” Bonis, L. de, Bouvrain, G., G eraads, D. & Koufos, Foundation. The Municipality of Mytilinii provided generous G.D., 1992a. Diversity and paleoecology of Greek assistance too. The Natural History Museum of the Aegean late Miocene mammalian faunas. — Palaeogeography offered us the premises for the preparation and storage of the Palaeoclimatology Palaeoecology, 91:99-121. fossils. For facilitating access to material under their care, we Bonis, L. de, Bouvrain, G., K oufos, G.D. ScTassy, R, are indebted to K. Heissig (BSPG); E. Heizmann (SMNS); O. 1995. Un crâne de chalicothère (Mammalia, Perisso- Sandrock, C. Kurz (HLMD); O. Hampe, N. Klein (NHMB); dactyla) du Miocène supérieur de Macédoine (Grèce): G. Hock (NHMW); M. Dermitzakis, G. Theodorou, S. remarques sur la phylogénie des Chalicotheriinae. — Roussiakis (AMPG); A. Currant, J. Hooker (BMNH); and Palaeovertebrata, 24(1-2):135-176, Montpellier. P. Tassy, C. Sagne (MNHN). We are obliged to K. Heissig Bonis, L. de, Bouvrain, G. 8c K oufos, G.D., 1999. for his useful remarks on the manuscript. Financial support Palaeoenvironments of late Miocene primate locali­ for comparative studies was provided to I. Giaourtsakis (EU- ties in Macedonia, Greece. — [in:]A gustí, J., Rook, SYNTHESYS: GB-TAF-574, FR-TAF-1226, FR-TAF-2545) L. & A ndrews, P. (eds.). Hominoid Evolution and and to G. Koufos (EU-SYNTHESYS: AT-TAF-702, FR- climatic change in Europe, Volume 1: The evolution TAF-3102, GB-TAF-1842) by the European Commission’s of Neogene terrestrial ecosystems in Europe — 1:413- Research Infrastructure Action, and the Research for Human 436, Cambridge (Cambridge University Press). Origin Initiative (RHOI, under NSF award BCS-0321893). Bonis, L. de, Brunet M., Heintz, E. &S en, S., 1992b. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

G iaourtsakis, I.X. & K 0UFOS, G.D., Chalicotheriidae. 199

La province gréco-irano-afghane et la répartition des Fortelius, M., 1985. Ungulate cheek teeth: develop­ faunes mammaliennes au Miocène supérieur. — Pale­ mental, functional, and evolutionary interrelations. ontología i Evolució, 24-25:103-112, Sabadell. — Acta Zoologica Fennica, 180:1-76, Helsinki. B o r is s ia k , A., 1945. The Chalicotheres as a biological Fortelius, M., 1990. Less common ungulate species type. — American Journal of Science, 243:667-679, from Pajalar, middle Miocene of Anatolia (Turkey). New Haven. — Journal of Human Evolution, 19:479-487, New B runet, M., H e in t z , E. ScBattail, B., 1984. Molayan York. (Afghanistan) and the Khaur Siwaliks of Pakistan: an Fortelius, M . & S olounias, N., 2000. Functional example of biogeographic isolation of Late Miocene characterization of ungulate molars using the abrasion mammalian faunas. — Geologie en Mijnbouw, -attrition wear gradient: a new method for recon­ 63:31-38, Utrecht. structing paleodiets. — American Museum Novitates, B u t l e r , P.M., 1965. East African Miocene and Pleis­ 3301:1-36, New York. tocene Chalicotheres. Fossil mammals of , 18. G arevski, R., 1974. Beitrag zur Kenntnis der Pikermifau- — Bulletin of the British Museum (Natural History) na Mazedoniens: Fossilreste der Chalicotheriiden. — Geology, 10:163-237, London. Fragmenta Balcanica Musei Macedonici Scientiarum C o o m b s, M.C., 1974. Ein Vertreter von Moropus aus dem Naturalium, 9:201-209, Skopje. europäischen Aquitanien und eine Zusammenfassung G arevski, R. & Zapfe, H., 1983. Weitere Chalicothe- der europäschen postoligozänen Schizotheriinae riiden-Funde aus der Pikermi-Fauna von Titov Veles (Mammalia, Perissodactyla, Chalicotheriidae). — (Mazedonien, Jugoslawien). — Acta Musei Macedo­ Sitzungsberichte der Österreichischen Akademie der nici Scientiarum Naturalium, 17:1-20, Skopje. Wissenschaften, Mathematisch-N aturwissenschaftli- G audry, A., 1862-67. Animaux fossiles et géologie de che Klasse, I, 182:273-288, Wien. lAttique. [The chapter onAncylotherium was published C o o m b s, M.C., 1975. Sexual dimorphism in chalicotheres in 1863] — pp. 474, Paris (Savy). (Mammalia, Perissodactyla). — Systematic Zoology G audry, A. & L artet, E., 1856. Sur les résultats de 24(l):55-62, London. recherches paléontologiques entreprises dans lAttique C o o m b s, M.C., 1978. Re-evaluation of early Miocene sous les auspices de lAcadémie. — Comptes Rendus North American Moropus (Perissodactyla, Chalico­ des Seances de lAcadémie des Sciences, 43:271-274, theriidae, Schizotheriinae). — Bulletin of Carnegie Paris. Museum of Natural History, 4:1-62, Pittsburgh. G eraads, D., S passov, N. & K ovachev, D., 2001. C o o m b s, M.C., 1979. Tylocephalonyx, a new genus of New Chalicotheriidae (Mammalia) from Bulgaria. North American dome-skulled Chalicotheres (Mam­ — Journal of Vertebrate Paleontology, 21:596-606, malia, Perissodactyla). — Bulletin of the American Northbrook. Museum of Natural History, 164:1-64, New York. G eraads, D., Spassov, N. &c Kovachev, D., 2006. The C o o m b s, M.C., 1982. Chalicotheres (Perissodactyla) as Bulgarian Chalicotheriidae (Mammalia): an update. large terrestrial mammals. — Third North American — Revue de Paléobiologie, 25(2):429-437, Genève. Paleontological Convention, Proceedings, 1:99-103, G eraads, D., Tsoukala, E. & Spassov, N., 2007. A skull Montreal. ofAncylotherium (Chalicotheriidae, Mammalia) from C o o m b s, M.C., 1983. Large mammalian clawed her­ the Late Miocene of Thermopigi (Serres, N. Greece) bivores: A comparative study. — Transactions of and the relationships of the genus. — Journal of Ver­ the American Philosophical Society, 73(7):l-96, tebrate Paleontology, 27(2):461-466, Northbrook. Philadelphia. G ill, T , 1872. Arrangement of the families of mammals C o o m b s, M.C., 1989. Interrelationships and diversity in with analytical tables. — Smithsonian Miscellaneous the Chalicotheriidae. — [in:] P r o t h e r o , D.R. &c Collections, 11:1-98, Washington. S c h o c h , R.M. (eds.). The evolution of Perissodactyls. G uérin, C. & P ickford, M ., 2005. Ancylotherium — Oxford Monographs on Geology and Geophysics, cheboitense nov. sp., nouveau Chalicotheriidae (Mam­ 15:438-457, New York (Oxford University Press). malia, Perissodactyla) du Miocène supérieur des C o o m b s, M.C., 2004. Moropus merriami in the Early Tugen Hills (Kenya). — Comptes Rendus Palevol, Barstovian Lower Snake Creek Fauna of Nebraska, 4:225-234, Paris. with comments on biogeography of North American H eissig, K., 1999. Family Chalicotheriidae. — [in:] Röss- Chalicotheres. — Bulletin of the American Museum NER, G.E. & H eissig K. (eds), The Miocene Land of Natural History, 285:191-208, New York. Mammals of Europe — 189-192, München (Verlag C o o m b s, M.C. & R o t h s c h il d , B.M., 1999. Phalan­ Dr. Friedrich Pfeil). geal fusion in schizotheriine chalicotheres (Mam­ H ensel, R.F., 1862. Über die Reste einiger Säugetier­ malia, Perissodactyla). — Journal of Paleontology, arten von Pikermi in der Münchener Sammlung. 73:682-690. — Monatsberichte der königlich-preußischen Akad­ D ie t r ic h , W.O., 1928. Lassen sich und emie der Wissenschaften zu Berlin, 1862:560—569, verwandte Gattungen schon rekonstruieren? — Zen­ Berlin. tralblatt für Mineralogie, Geologie und Paläontologie, Holland, W.J. & P eterson, O.A., 1914. The osteology 1928:366-376, Berlin. of the with special reference to ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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a mounted skeleton ofMoropus elatus M a r s h , now M ecquenem, R. de, 1924. Contribution à l'étude des installed in the Carnegie Museum. — Memoirs of the fossiles de Maragha. — Annales de Paléontologie, Carnegie Museum, 3:189-406, Pittsburgh. 13:135-160, Paris. K a y a , T., 1986. Çanakkale ve çevresi Perissodactyla M elentis, J.K., 1969a. Studien über fossile Vertebraten fosilleri (paleontoloji, biyostratigrafi, paleoekoloji ve Griechenlands 28: Die Pikermifauna von Halmyro- paleocografya). — Ph.D. thesis, Ege University Sci­ potamos (Euböa - Griechenland), 2. Teil: Osteolo­ ence Faculty, pp. 239, Izmir. gie. — Annales Géologiques des Pays Helléniques, K a y a , T., 1988. Kemiklitepe (U^ak, Esme) faunasindaki 21:217-306, Athens. Perissodactyla buluntulari. — 9th National Biology M elentis, J.K., 1969b. Ancylotheriumpentelicum G audry Congress, 1:355-361, Sivas. & L artet aus dem Pont von Halmyropotamos K a y a , T., 1993. First record of Moropus elatus (Chali- (Euböa, Griechenland). — Archives des études cotheriidae - Perissodactyla) in Turkey (Seyitömer d’Eubée, 15:1-18, Chalkida. Kütahya). — Turkish Journal of Earth Sciences, Nikolov, L, 1972. On the stratigraphic level of M eta- 2:189-194, Ankara. schizotherium fraasi K oenigswald. [in Bulgarian], K a y a , T , G e r a a d s , D. & T u n a , V , 2005. A new late — Bulgarian Academy of Sciences, Bulletin of the Miocene mammalian fauna in the Karaburun Penin­ Geological Institute, Series Paleontology, 21:129-131, sula (W. Turkey). — Neues Jahrbuch für Geologie Sofia. und Paläontologie, Abhandlungen, 236(3):321-349, Nikolov, L, 1975. Neue Fossilienfundevon Familie Chali- Stuttgart. cotheriidae (Mammalia) aus Bulgarien. — Review K a y a , T , T u n a , V. & G e r a a d s , D., 2001. A new late of the Bulgarian Geological Society, 36:311-313, Orleanian / early Astaracian Mammalian fauna from Sofia. Kultak (Milas-Mugla), southwestern Turkey. — Géo­ O nar, V. & Yildiz, B., 2005. New Chalicotheriidae bios, 34(6):673-680, Lyon. (Perissodactyla-Mammalia) fossil from Miocene of K o e n ig s w a l d , G.H.R. von, 1932. Metaschizotherium Turkey (Bursa-Orhaneli). — Uludag University, Jour­ fraasi n.g. n.sp., ein neuer Chalicotheriide aus dem nal of the Faculty of Veterinary Medicine, 24:65-68, Obermiozän von Steinheim a. Albuch. (Die tertiären Uludag. Wirbeltiere des Steinheimer Beckens VIII). — Palae- Petronijevic, Z., 1952. Chalicotheriumgrande (Lartet) ontographica, Supplement, 8(8):ll-24, Stuttgart. in den Kohlen des Plevtjabecken (Cerna Gora - K o r o t k e v ic h , O., 1988. Genesis of the Hipparion-fauna Monte-negro). — Recueil des travaux de l'institut de of Eastern Europe [in Russian], — pp. 160, Naukova Geologie Jovan Zujovic, 9:93-100, Belgrade. Dumka, Kiev. P idoplichko, I.G., 1959. New finds of Mammals in the K o st o p o u l o s, D.S., K o u f o s, G.D., S y l v e st r o u , I.A., Neogene of the Ukraine [in Russian]. — Dopovidy S y r id e s , G.E. & T sompachidou , E., this volume. Akademii NaukUkrainskoi RSR, 5:531-534, Kiev. The Late Miocene Mammal Faunas of the Mytilinii R innert, R, 1956. Die Huftiere aus dem Braunkoh­ Basin, Samos Island, Greece: New Collection. 2. lenmiozän der Oberpfalz. — Palaeontographica, A, Lithostratigraphy and Fossiliferous Sites. — Beiträge 107:1-65, Stuttgart. zur Paläontologie, 31:13-26, Wien. Roussiakis, S.J. Sc Theodorou, G.E., 2001. Ancyloth­ K o u f o s, G.D., this volume. The Late Miocene Mammal eriumpentelicum (Gaudry & Lartet, 1856) (Peris­ Faunas of the Mytilinii Basin, Samos Island, Greece: sodactyla, Mammalia) from the classic locality of New Collection. 1. History of the Samos Fossil Mam­ Pikermi (Attica, Greece), stored in the palaeontologi­ mals. — Beiträge zur Paläontologie, 31:1-12, Wien. cal and geological museum of Athens. — Geobios, K o u f o s, G.D., K o st o p o u l o s, D.S. &V l a c h o u , D., this 34:563—584, Lyon. volume. The Late Miocene Mammal Faunas of the Saraç, G., K aya, T. & G eraads, D., 2002. Ancylotherium Mytilinii Basin, Samos Island, Greece: New Collec­ pentelicum (Perissodactyla, Mammalia) from the upper tion. 16. Chronology. — Beiträge zur Paläontologie, Miocene of central and western Turkey. — Geobios, 31:397-408, Wien. 35:241—251, Lyon. K o u f o s, G.D., K o st o p o u l o s, D.S. S c M ercero n , G., Saraç, G. &S en, S., 2005. Chalicotheriidae (Mammalia, this volume. The Late Miocene Mammal Faunas Perissodactyla) from the late Miocene of Akkajdagi, of the Mytilinii Basin, Samos Island, Greece: New Turkey. — [in:] Sen, S. (ed.). Geology, mammals and Collection. 17. Palaeoecology - Palaeobiogeography. environments at Akkaçdagi, late Miocene of Central — Beiträge zur Paläontologie, 31:409-430, Wien. Anatolia. — Geodiversitas, 27(4):591-600, Paris. M a jo r , C.J.F., 1888. Sur un gisement d’ossements fossiles Schaub, S., 1943. Die Vorderextremität von Ancylotherium dans l’île de Samos, contemporain de l’âge de Pikermi. pentelicum G audry & L artet. — Schweizerische — Comptes Rendus des Séances de l’Académie des Paläontologische Abhandlungen, 64:1-36, Basel. Sciences de Paris, 107:1178-1182, Paris. Schlosser, M., 1921. Die Hipparionenfauna von Veles M a jo r , C.J.F., 1894. Le gisement ossifère de Mytilinii et in Mazedonien. — Abhandlungen der Bayerischen catalogue d ossements fossiles recueillis à Mitylinii, île Akademie der Wissenschaften, Mathematisch- de Samos, et déposés au Collège Galliard, à Lausanne. Physikalische Klasse, 29(4):l-55, München. — 51 pp, Lausanne (Georges Bridel & Cie). 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T.M., 2007. Feeding ecology of the Chalicotheriidae silen Säugethier-Überreste von Pikermi. — Abhand­ (Mammalia, Perissodactyla, ). Results lungen der königlichen bayerischen Akademie der from dental micro- and mesowear analyses. — Ver­ Wissenschaften, Mathematische und Physikalische handlungen des Naturwissenschaftlichen Vereins Klasse, 8:109-158, München. Hamburg, Neue Folge, 43:5-31, Hamburg. Wu, W.Y. & C hen, G.F., 1976. A new schizotheriine Sen, S., 1994. Les gisements de mammiferès du Miocène genus from the Neogene of Pingliang, Gansu. — supérieur de Kemiklitepe, Turquie: 5. Rongeurs, Tu- Vertebrata PalAsiatica, 14:194-197, Beijing. bulidentés et Chalicothères. — Bulletin du Muséum Zapfe, H., 196 7. Ancylotheriumpentelicum im Obermiozän National d’Histoire Naturelle, 16(4):97-111, Paris. des Wiener Beckens. — Annalen des Naturhistori­ Symeonidis, N.K., 1973. Chalicotherium goldfussi K aup schen Museums in Wien, 71:401-411, Wien. (Perissodactyla, Mammalia) aus dem Altpliozän von Zapfe, H., 1974. Ein schizotheriiner Chalicotheriidae Pikermi (Griechenland). — Annales Géologiques des aus pannonischen Schottern der Umgebung von Pays Helléniques, 25:301-307, Athens. Krems, Niederösterreich. — Sitzungsberichte der Thenius, E., 1953. Studien über fossile Vertebraten Österreichischen Akademie der Wissenschaften, Griechenlands. III. Das Maxillargebiss von Ancy- Mathematisch-Naturwissenschaftliche Klasse, I, lotherium pentelicum G audry und Lartet. — A n­ 182:289-299, Wien. nales Géologiques des Pays Helléniques, 5:97-106, Zapfe, H., 1976. Die Fauna der miozänen Spaltenfüllung Athens. von Neudorf a.d. March (CSSR), Chalicotheriumgrande Theodorou, G., A thanassiou, A., Roussiakis, S. &Ili- (Blv.). — Sitzungsberichte der Österreichischen Aka­ opoulos, G., 2003. Preliminary remarks on the Late demie der Wissenschaften, Mathematisch-Naturwis­ Miocene herbivores of Kerassia (Northern Euboea, senschaftliche Klasse, 1 , 185:91-112, Wien. Greece). — Deinsea, 10:519-530, Rotterdam. Zapfe, H. 1979. Chalicotherium grande (Blainv.) aus der Tung, Y.S., Huang, W.P. &Qiu, Z.D., 1975. Hipparion miozänen Spaltenfüllung von Neudorf an der March fauna in Anlo, Hohsien, Shansi. — Vertebrata PalA- (Devinska Nova Ves), Tschechoslowakei. — Neue siatica, 13:34-47, Beijing. Denkschriften des Naturhistorischen Museums in W agner, A., 1857. Neue Beiträge zur Kenntnis der fos­ Wien, 2:1-282, Wien. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

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PLATE 1

Ancylotheriumpentelicum (Gaudry & Lartet, 1856), Mytilinii-lB (MTLB), Samos, Greece,

middle Turolian (MN 12).

Fig. 1-2. Right radius and ulna, M TLB-328; 1. cranial and 2. distal view

Fig. 3. Left radius and ulna, M TLB-173; cranial view.

Fig. 4-5. Left femur, M TLB-338; 4. cranial, 5. caudal and 6. medial view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Giaourtsakis, I.X. &K oufos, G.D., Chalicotheriidae. 203

PLATE1 ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

204 Beitr. Paláont., 31, Wien, 2009

PLATE2

Ancylotheriumpentelicum (Gaudry & Lartet, 1856), M ytilinii-lA, 3 (MTLA, MYT), Samos, Greece,

middle Turolian (MN 12).

Fig. 1-3. Right Mt-III, M YT-64; 1. dorsal, 2 . lateral and 3. proximal view.

Fig. 4-6. Right Mt-III, M TLA-277; 4. dorsal, 5. lateral and 6. proximal view.

Fig. 7-8. Ungual phalanx, M TLA-485; 7. dorsal and 8 . plantar view.

Fig. 9. Proximal phalanx, M YT-15; dorsal view. ©Verein zur Förderung der Paläontologie am Institut für Paläontologie, Geozentrum Wien

Giaourtsakis, I.X. &K oufos, G.D., Chalicotheriidae. 205

PLATE 2