Dryopithecins, Darwin, de Bonis, and the European origin of ......791 Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade GEODIVERSITAS •

789GEODIVERSITAS • 2009 • 31 (4) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com

KEY WORDSMammalia,

Primates,Dryopithecus,

Hispanopithecus, Rudapithecus,

Ouranopithecus, hominine origins,

new subtribe.

Begun R. D. 2009. — Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade. Geodiversitas 31 (4) : 789-816.

ABSTRACTDarwin famously opined that the most likely place of origin of the common ancestor of African apes and humans is Africa, given the distribution of its liv-ing descendents. But it is infrequently recalled that immediately afterwards, Darwin, in his typically thorough and cautious style, noted that a fossil ape from Europe, Dryopithecus, may instead represent the ancestors of African apes, which dispersed into Africa from Europe. Louis de Bonis and his collaborators were the fi rst researchers in the modern era to echo Darwin’s suggestion about apes from Europe. Resulting from their spectacular discoveries in Greece over several decades, de Bonis and colleagues have shown convincingly that African ape and human clade members (hominines) lived in Europe at least 9.5 million years ago. Here I review the fossil record of hominoids in Europe as it relates to the origins of the hominines. While I diff er in some details with Louis, we are in complete agreement on the importance of Europe in determining the fate of the African ape and human clade. Th ere is no doubt that Louis de Bonis is a pioneer in advancing our understanding of this fascinating time in our evo-lutionary history.

David R. BEGUNUniversity of Toronto, Department of Anthropology, 19 Russell Street, Toronto, ON, M5S 2S2 (Canada)

[email protected]

Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade

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MOTS CLÉSMammalia,

Primates,Dryopithecus,

Hispanopithecus, Rudapithecus,

Ouranopithecus, origines des hominiens,

sous-tribu nouvelle.

RÉSUMÉLes Dryopithèques, Darwin, de Bonis et l’origine européenne du clade des grands singes d’Afrique et de l’homme.L’avis réputé de Darwin était que l’endroit le plus probable de l’origine de l’ancêtre commun des grands singes africains et de l’homme est l’Afrique, étant donné la distribution des descendants actuels. Mais il est rarement rappelé que immédia-tement après, Darwin, dans son style typiquement minutieux et prudent, indi-quait qu’un grand singe européen, Dryopithecus, représentait peut-être un ancêtre potentiel des grands singes d’Afrique, qui se seraient dispersés de l’Europe vers l’Afrique au cours du Miocène. Louis de Bonis et ses collaborateurs furent parmi les premiers chercheurs de l’ère moderne à adopter cette suggestion de Darwin. Suite à leurs découvertes spectaculaires en Grèce pendant plusieurs décennies, de Bonis et ses collègues ont démontré d’une façon convaincante que des mem-bres du clade des grands singes d’Afrique et de l’homme ont persisté en Europe jusqu’à 9,5 millions d’années au moins. Je résume ici l’histoire des découvertes en Europe concernant l’origine des hominiens. Bien que mes conclusions se dis-tinguent dans les détails de celles de Louis, nous sommes entière ment d’accord sur l’importance de l’Europe dans le destin du clade des grands singes africains et de l’homme (Bonis & Koufos 2004). Il n’y a pas de doute que Louis de Bonis est un pionnier qui a contribué d’une façon exceptionnelle à nos connaissances de cette période fascinante de notre histoire évolutive.

humans to the exclusion of orangutans) was not overwhelming, and the gradistic resemblances among extant great apes are undeniable. I think it is fair to say that Darwin and Huxley were not vindicated until genetic technology was able to es-tablish with little doubt that chimpanzees are more closely related to humans than they are to gorillas, and that the group of African apes and humans as a whole is more closely related to each other than any of them are to orangutans.

During this time, researchers were trying to inte-grate fossil discoveries with our understanding of ape and human evolution. Darwin felt it most likely that African apes and humans evolved in Africa, stating the following: “In each great region of the world the living mammals are closely related to the extinct species of the same region. It is therefore probable that Africa was formerly inhabited by extinct apes closely allied to the gorilla and chimpanzee; and as these two species are now man’s nearest allies, it is somewhat more probable that our early progeni-tors lived on the African continent than elsewhere.” (Darwin 1871: 199).

INTRODUCTION

In 1871, Charles Darwin published the fi rst serious, comprehensive and well-informed analysis of human evolutionary history. In this work, “Th e Descent of Man, and Selection in Relation to Sex”, Darwin compares the morphology of extant great apes to humans, and he concludes that among other things humans and African apes are more closely related to one another than either are to orangutans. Here he was drawing heavily on the work of Th omas Henry Huxley and his most important publication, “Evidence as to Man’s Place in Nature” (Huxley 1863). While today we recognize the phyletic link between African apes and humans, based both on morphological and molecular evidence, it is often forgotten that for much of the 20th century Dar-win and Huxley’s conclusions were rejected, and the great apes were routinely linked together, with humans set apart. In fairness to historically strong advocates of the Pongidae (great apes), such as Adolph Schultz (1936, 1950), the morphological evidence for a hominine clade (African apes and

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However, Darwin knew of a discovery in France of a fossil with close similarities to living great apes. Here is what he had to say about that fossil on the very next line: “But it is useless to speculate on this subject, for an ape nearly as large as a man, namely the Dryopithecus of Lartet, which was closely al-lied to the anthropomorphous Hylobates, existed in Europe during the upper Miocene period; and since so remote a period the earth has certainly undergone many great revolutions, and there has been ample time for migration on the largest scale.” (Darwin 1871: 199).

Lartet himself spoke of the similarities between Dryopithecus Lartet, 1856 and African apes, and these observations were repeated by later research-ers commenting on newer discoveries in France and Germany (Lartet 1856; Gaudry 1890; Branco 1898; Harlé 1898; Schlosser 1901; Abel 1902). However, following impressive discoveries fi rst in Asia and then in Africa, attention shifted away from Europe. “Th e Dryopithecus of Lartet” was eventually lumped in with most other fossil apes, which were distinguished from what were then taken to be the putative ancestors of humans, the ramapi thecines (Simons & Pilbeam 1965). At-tention was turned away from Europe until the early 1970’s, when discoveries from Greece and Hungary, and an emerging understanding of the evolutionary position of the Siwalik fossil apes, were leading to a renewal of interest in Europe (see Begun [2002] for a more detailed history). In 1974, a new species of Dryopithecus, D. macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974), was recognized on the basis of fossil specimens from northern Greece. In the same year Kretzoi (1975) published an account of a genus of fossil ape, Rudapithecus Kretzoi,1969, from Hungary. In both cases, but based on very diff erent lines of evidence, Europe was being repositioned as an area of great importance in interpretations of ape and human evolutionary history.

In the intervening 30 years or so, many new discoveries have been made in both Greece and Hungary, as well as in Spain, and these have shaped the debate on the relevance of Europe in the evolu-tionary history of the African apes and humans. For the remainder of this paper I will review the data

from these three most important locations, along with isolated discoveries from Turkey and Bulgaria, to try to make some sense of the fossil evidence of hominids in the Miocene of Europe (see below for terminology used in this paper).

ABBREVIATIONS InstitutionsAU Ankara University;AUT Aristotle University of Th essaloniki;GMH Geological Museum of Hungary, Budapest;IPS Institut Paleontologic, Sabadell (Institut

Català de Paleontologia);NMB Naturhistorisches Museum Basel.

LocalitiesCO Çorakyerler collection (in AU);RUD Rudabánya primate collection (in GMH);XIR Xirochori collection (in AUT).

EUROPEAN HOMININE TAXONOMY

Before reviewing the fossil record of European hominines it is necessary to explain the taxonomic usage adopted here. With regard to the sample from Greece, I recognize a genus level distinction between Ouranopithecus Bonis & Melentis, 1977 and Graecopithecus von Koenigswald, 1972 (Begun 2002, 2007a; Koufos & Bonis 2005). In Graecopithecus the M2 is close in size to male Ouranopithecus, but the mandible is the size of a female Ouranopithecus. In fact, the breadth of the M2 is greater than the mandibular breadth at that level. Th is suggests an important diff erence in relative tooth size between the two taxa. In addition, the symphysis in Graeco-pithecus is more vertical in orientation, and the M1 is smaller relative to the M2 (Begun 2002). While they are often lumped together by virtue of the fact that both appear to have thickly enameled teeth and both are found in Greece, it is worth noting that the shortest distance between an Ouranopithecus locality (Nikiti) and Pyrgos Vassilissis (Graeco-pithecus) is about 250 km as the crow fl ies today. In the late Miocene the Th ermaikos Gulf and Aegean Sea intervened between the Chalkidiki Peninsula and mainland Greece as it does today, so the land distance between the two localities is even greater. In addition, Nikiti, the youngest Ouranopithecus

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locality, is estimated to be about 9 Ma (Koufos 2003), and Pygros Vassilissis, though diffi cult to date, is likely to be MN 12, or 6-8 Ma, and the ecology of Greece during these diff erent time periods was quite diff erent (Solounias 1981; Bernor 1983; Bonis et al. 1994; Bernor et al. 1996; Mein 1999; Solounias et al. 1999; Steininger 1999; Agustí et al. 2003; Begun & Nargolwalla 2004; Begun 2005; Merceron et al. 2005a; Agustí 2007; Strömberg et al. 2007). Th is taxonomic interpretation is consist-ent with Bonis & Melentis (1984).

A recent review of the taxa usually attributed to Dryopithecus has led to a revised taxonomy (Begun et al. 2008). With the discovery of a number of partial crania, diversity among the dryopithecins has become clearer. Th e large sample from the Vallès Penedès expanded dramatically with the discovery of a partial skeleton from Can Llobateres (Begun & Moyà-Solà 1992; Moyà-Solà et al. 1992; Moyà-Solà & Köhler 1993, 1995, 1996), a second partial skeleton from Hostalets de Pierola (Moyà-Solà et al. 2004), a palate from Can Mata (Moyà-Solà et al. 2009a) and a partial face from Can Mata (Moyà-Solà et al. 2009b). When compared to the postcranial sample and the three partial crania now known from Rudabánya, there are a suffi cient number of diff erences to warrant the recognition of separate genera for each sample (Begun et al. 2008).

Pierolapithecus catalaunicus Moyà-Solà, Köhler, ALba, Casanovas-Villar & Galindo, 2004, from Barranc de Can Vila 1 in Catalonia, Spain, is known from a partial skeleton including much of the face and portions of the postcranial skeleton (Moyà-Solà et al. 2004). Th e locality is dated to MN 7/8, and is therefore older than Can Llobateres and Rudabánya, both of which are MN 9 or early Vallesian (Begun 2002). Barranc de Can Vila 1 is the same age as Dryo-pithecus fontani Lartet, 1856 from St-Gaudens and La Grive (France) and from Can Vila and Can Mata (Spain). Pierolapithecus Moyà-Solà, Köhler, ALba, Casanovas-Villar & Galindo, 2004 is distinct from and more primitive than the Can Llobateres and/or Rudabánya samples in that the lumbar vertebrae have somewhat more anteriorly positioned transverse processes (not known at Rudabánya) and the proximal hand phalanges are shorter, less curved with proximal articular surfaces oriented more dorsally. Th e face is

diffi cult to compare to the Can Llobateres and Ruda-bánya samples because it is distorted. It is unlikely that the face of Pierolapithecus resembled Afropithecus Leakey & Leakey, 1986, as suggested by Moyà-Solà et al. (2004), after the distortion is corrected (Begun & Ward 2005; Begun 2007a). Th e teeth are quite similar to MN 9 dryopithecins but, interest-ingly, the I1 and M3 most closely resembles those attributed to Dryopithecus fontani from La Grive (Begun & Ward 2005) (Fig. 1). In addition, the canines are relatively large, as is the case for the lower canines of Dryopithecus fontani from St-Gaudens. Like MN 7/8 Dryopithecus, Pierolapithecus has partial molar cingula, short premolars and a smaller M1 relative to M2. Th e lower dentition of Dryopithecus from France also has relatively lower premolar talonids, a primitive character, compared to MN 9 Euro pean hominines. Two previously known isolated lower molars also from Can Vila share attributes with French Dryopithecus, including partial buccal cingula and a well-developed tuberculum sextum on the M3 (Fig. 1B) (Begun & Moyà-Solà 1990; Begun 2002). In summary, the samples from Can Vila and France are the same age and have many attributes in common, and they are generally more primitive than MN 9 hominines. I therefore recognize Pierolapithecus as a junior subjective synonym of Dryopithecus (Begun et al. 2008). Th is interpretation is further supported by the recent discovery of a Dryopithecus fontani palate, from the nearby MN 7/8 locality of Can Mata, and a partial face attributed to the new genus Anoiapithecus Moyà-Solà, Alba, Almécija, Casanovas-Vilar, Köh-ler, Esteban-Trivigno, Robles, Galindo & Fortuny, 2009, also from Can Mata, which are both very similar dentally to Pierolapithecus (Moyà-Solà et al. 2009a, b). Th ese new specimens are all attributed to Dryopithecus (Begun et al. in press).

As noted above, MN 9 dryopithecins are derived relative to Dryopithecus as defi ned here. Th e best samples of MN 9 dryopithecins are from Can Llo-bateres and Rudabánya. Here I recognize separate genera for each of these two samples, Hispanopithecus Villalta & Crusafont, 1944 at Can Llobateres and Rudapithecus at Rudabánya, as originally suggested by Kretzoi (1969) and Villalta & Crusafont (1944). Th e best preserved and most informative specimens from Can Llobateres are a face and partial skeleton,

793



AAE F

CD

A

A-D

E, F

B

IPS 18000 and IPS 18800 (Moyà-Solà & Köhler 1993, 1995; Begun 1994; Almécija et al. 2007). Th ese fossils were originally attributed to the same individual. In the cranial specimen the frontal squama of Hispanopithecus is strongly biconcave transversely and anteroposteriorly, while in Rudapi-thecus it is smoothly bi-convex. In Hispanopithecus there is a groove separating the supraorbital ridge from the supraorbital torus. Th e anterior temporal lines in Hispanopithecus are very strongly developed and raised above the cranial surface immediately anterior and medial to it. IPS 18000 is a male, as is RUD 44. Th e latter is larger than IPS 18000 but with less strongly developed temporal ridges. Th e

surface of the zygomatic process of the frontal (the superolateral corner of the orbit) in Hispanopithecus also diff ers from the three Rudapithecus specimens in being more anteriorly oriented. In Hispanopithecus the body of the zygomatic bone is fl at and faces more anteriorly than in Rudapithecus, in which all three specimens have more convex zygoma facing more laterally (Begun 1994). Although highly variable in extant hominoids, the position, size and number of the zygomaticofacial foramina also diff er between the two taxa, with those in Hispanopithecus being relatively large, more numerous, and higher on the zygomatic frontal process (Begun 1994; Moyà-Solà & Köhler 1995).

FIG. 1. — Comparisons between Pierolapithecus Moyà-Solà, Köhler, Alba, Casanovas-Villar & Galindo, 2004 and D. fontani Lartet, 1856: A, Pierolapithecus I1; B, La Grive I1, NMB g. a. 9; C, Pierolapithecus M3, NMB; D, la Grive M3; E, left M3 from Can Vila; F, left M3 from D. fontani. Images of the Pierolapithecus specimens are modifi ed from Moyà-Solà et al. (2004). Scale bars: 1 cm.

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Th e palate of Rudapithecus is narrower than that of Hispanopithecus, and the zygomaticoalveolar crests are thicker anteroposteriorly. Th e mandible of Rudapithecus is generally more robust and oval in cross section. Th e only identifi ably male mandible of Hispanopithecus (IPS 1764) has a triangular shaped cross section between P4-M1. Dentally, the upper incisor crowns are narrow compared to H. laietanus Villalta & Crusafont, 1944, with concave lingual surfaces and single central pillars. Hispanopithecus laietanus I1s have complicated cresting on their lingual surfaces, and H. crusafonti Begun, 1992I1s are extremely narrow with very prominent me-sial, distal and central labial pillars (Begun 1992a, 1994). In Rudapithecus the canines are smaller and more compressed buccolingually and the up-per postcanine teeth tend to be more elongated relative to breadth. Th e upper and lower M3s are

elongated, strongly tapered, and are smaller than the M2s in Rudapithecus. Males of Rudapithecus diff er from males of Hispanopithecus in having large lower premolars, strongly triangular (beaked) P3, large P3 fovea of equal height with buccal crista of equal length, and broad, wrinkled lower molar basins with marginalized cusps (Begun & Kordos 1993; Begun 1994).

Postcranially, Rudapithecus diff ers from Hispano-pithecus in having much less robust manual proximal phalanges with a more proximally oriented proximal articular surface and broader trochlea. Th e metacarpal is also less robust transversely. Th e femoral neck in Rudapithecus is larger supero inferiorly relative to head diameter compared to Hispanopithecus (MacLatchy et al. 2001; Köhler et al. 2002; and pers. obs.). Table 1 lists the most important European hominine taxa and their main diagnostic characters.

TABLE 1. — Principle taxa discussed in this paper and their most important diagnostic craniodental characters.

Dryopithecus Lartet, 1856

Short, vertical premaxilla with evidence of a subnasal step (associated with maxillary over-lap), biconvex premaxilla, modest supraorbital tori and supratoral sulcus, broad interorbital space, robust lateral orbital margins, square orbits, subvertical nasal aperture margins, high zygomaticoalveolar crest, relatively broad canines, short premolars and small M1, low lower premolar talonids, partial molar cingula, thin occlusal enamel, tall crowned postcanines, M3 tuberculum sextum, somewhat robust mandibles, mesio-distally short, tall crowned I1

HispanopithecusVillalta & Crusafont, 1944

Short, vertical premaxilla with evidence of a subnasal step (associated with maxillary overlap), biconvex premaxilla, modest supraorbital tori and supratoral sulcus, strongly developed anterior temporal lines, biconcave vertical frontal squama, reduced subarcuate fossa, ro-bust lateral orbital margins, square orbits, broad interorbital space, ethmoidal frontal sinus, subvertical nasal aperture margins, high zygomaticoalveolar crest, more anteriorly oriented zygoma, thin enamel, mesio-distally short, tall crowned I1, peg-shaped I2

Rudapithecus Kretzoi, 1969

Short, vertical premaxilla with some maxillary overlap, relatively narrow, posteriorly divergent palate, biconcave premaxilla, modest supraorbital tori and supratoral sulcus, vertical frontal squama, elongated, klinorhynch crania, inion high on the skull, reduced subarcuate fossa, fused tympanic and articular temporal, large brain, robust lateral orbital margins, square orbits, broad interorbital space, ethmoidal frontal sinus, subvertical nasal aperture margins, high zygomaticoalveolar crest, convex and more laterally facing zygoma, more compressed canines and elongated postcanines, strongly tapered M3, more peripheral cusp positions, mesio-distally short, tall crowned I1, peg-shaped I2

Ouranopithecus Bonis & Melentis, 1977

Short, vertical premaxilla with some maxillary overlap, modest supraorbital tori and supra-toral sulcus, vertical frontal squama, robust lateral orbital margins, subvertical nasal aperture margins, low zygomaticoalveolar crest, strongly heteromorphic upper incisors, reduced ca-nine size and premolar sectoriality, hyperthick occlusal enamel, tall crowned and megadont postcanines, broad rounded cusps, robust to very robust mandibles

Çorakyerler hominine Very short premaxilla, upper canines nearly aligned with incisors transversely, homomorphic upper premolars, extremely large postcanine, teeth, narrow palate, thick, diamondshaped upper canines cervix reduced in size relative to the molars, thin palatine process, low crowned I1, peg-shaped I2

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Th e classifi cation of European hominines fol-lowed here is as follows:

Subfamily HOMININAE Gray, 1825Tribe DRYOPITHECINI Gregory & Hellman, 1939Subtribe DRYOPITHECINA Gregory & Hellman, 1939

DIAGNOSIS. — Th e Dryopithecina is diagnosed as follows: medium to large bodied hominines (20-45 kg) with thinly enameled teeth, robust and generally mesiodistally short upper incisors, tall crowned lower incisors, compressed canines, large, elongated premolars, M1 close in size to M2, peripheralized molar cusps, seven day cross striation periodicity, short premaxilla, stepped subnasal fossa, short, large caliber incisive canal, larger incisive foramen and fossa, robust palatine process, broad, fl at inferior margin of the nasal aperture, high zygomatic root, large maxil-lary sinus, large ethmoid sinus continuous with a well-developed frontal sinus, thick interorbital space, weakly developed supraorbital tori, prominent anterior temporal lines, elongated brain case, inion slightly above glabella, large brain, fused articular and tympanic temporal, deep temporal fossa, prominent entoglenoid process, robust mandible, large ramus, posteriorly diverging tooth rows, elongated, tapered M3.

Genus Dryopithecus Lartet, 1856 Dryopithecus fontani Lartet, 1856 Dryopithecus catalaunicus Moyà-Solà & Köhler, 2004 Dryopithecus carinthiacus Mottl, 1957Genus Rudapithecus Kretzoi, 1969 Rudapithecus hungaricus Kretzoi, 1969Genus Hispanopithecus Villalta & Crusafont, 1944 Hispanopithecus crusafonti Begun, 1992 Hispanopithecus laietanus Villalta & Crusafont, 1944

Subtribe OURANOPITHECINA new subtribe

DIAGNOSIS. — Th e Ouranopithecina is diagnosed as follows: Large bodied hominines (45+? kg) with thickly enameled teeth, low, rounded cusps, broad, shallow oc-clusal basins, enlarged M3’s, anterior teeth and dental proportions as in Dryopithecina, except small male canine crown dimensions, premaxilla and maxilla as in Dryopithecina except low, thick zygomaticoalveolar crest, periorbital region as in Dryopithecina.

Genus Ouranopithecus Bonis & Melentis, 1977 Ouranopithecus macedoniensis Bonis & Melentis, 1977Genus Graecopithecus von Koenigswald, 1972 Graecopithecus freybergi von Koenigswald,

1972Gen. et sp. nov. (Çorakyerler) (see below)

MACEDONIA, GREECE

As noted above, in 1974 de Bonis and colleagues named a new species of Dryopithecus in Macedonia, from the locality of Ravin de la Pluie. Although from Europe, this fossil is strikingly diff erent from other European fossil apes known at the time, par-ticularly in the robust morphology of the mandible and the large and thickly enameled postcanine teeth (Bonis & Melentis 1976, 1977a, b, 1978, 1980; Begun & Kordos 1997). Th e canines are also unusually small, although the type specimen is a female, and over the years, the relative canine size in this taxon even in males has proven to be a diffi cult issue to resolve (Bonis & Koufos 2001; Kelley 2001). More discoveries of the same taxon at Ravin de la Pluie as well as two other localities in Macedonia, Xirochori and Nikiti, have made clear the diff erences from other European and African Miocene fossil apes from the same time period and before as well.

De Bonis and Melentis attributed the new taxon from Greece to Dryopithecus, but they later re-vised their taxonomic conclusions in light of the similarities they saw with Sivapithecus Pilgrim, 1910, Gigantopithecus von Koenigswald, 1935 and australopithecines (Bonis & Melentis 1977b). Th ese include the large, robust jaws and thickly enameled teeth that distinguish Ouranopithecus from Dryopithecus, but also details of canine and premolar morphology and the structure of the palate (Bonis & Melentis 1977b, 1978; Bonis 1983). Ad-ditional preparation of the palate of Ouranopithecus would lead de Bonis and Melentis to interpret it as a direct link between Miocene apes and australo-pithecines, sharing synapomorphies with the tribe Hominini Grey, 1825, in which they include Homo Linnaeus, 1758 and Australopithecus Dart, 1925 (Bonis 1983; Bonis & Melentis 1984). In eff ect, they are attributing Ouranopithecus to the Hominini,

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in the relative size of the canine, changes in canine-premolar relationships, postcanine megadontia and increases in gnathic robusticity and enamel thick-ness characterize several lineages of fossil hominids (Sivapithecus, Ankarapithecus Ozansoy, 1965, Gripho-pithecus Abel, 1902, Gigantopithecus, Paranthropus Broom, 1939, Paraustralopithecus Arambourg & Coppens, 1968). Th ese features leave an overwhelm-ing impression on facial and dental morphology, though they are all closely interrelated and essen-tially constitute a single trait complex that results from selection for more powerful mastication. As noted, the history of interpretation is quite revealing in this regard. At one time or another, all thickly enameled middle and late Miocene hominoids were grouped into the same genus (Sivapithecus) and linked phyletically to one another and to other similar taxa such as Gigantopithecus (Andrews & Tobien 1977; Bonis & Melentis 1977a, b, 1978; Andrews & Tekkaya 1980; Andrews 1983, 1985). However, when all available characters are consid-ered, Ouranopithecus consistently falls out among the hominines as the sister clade of the African ape and human clade (Begun et al. 1997; Begun 2001, 2002, 2005, 2007a) (Fig. 2B).

Most signifi cantly, Ouranopithecus lacks several characters shared between chimpanzees and early fossil humans such as Australopithecus. Th ese include the structure of the premaxilla, which is more similar to that of Gorilla and Rudapithecus than Australo-pithecus and Pan (Bonis & Melentis 1987; Bonis & Koufos 1993; Begun 2001, 2002; Koufos 2007). Ouranopithecus shares the primitive condition for hominines, a relatively short and vertical premaxilla, a strongly stepped subnasal fossa and a large incisive fossa and canal. Th e nasal border of the premaxilla overlaps minimally with the palatine process of the maxilla. In Pan and Australopithecus the premaxilla is longer and more horizontally oriented, and its nasal border overlaps the maxillary palatine process to a considerably greater degree, which produces a reduced subnasal step, a smaller incisive fossa and a longer, narrower incisive canal (Ward & Kimbel 1983; Begun et al. 1997; Begun 2001, 2002). It could be argued that these premaxillary features also constitute a single character (Schwartz 1997). However, this complex actually cross cuts functional

though they do not say this directly. Bonis (1983) also attributes Dryopithecus and Hispanopithecus to the Dryopithecini Gregory & Hellman, 1939, in which he includes Pan Oken, 1816 and Gorilla Geoff roy-Saint-Hilaire, 1852, which is very close to the phylogeny followed here.

Begun & Kordos (1997) make the case for a slightly diff erent interpretation of Ouranopithecus and Dryopithecus, but one that still places them fi rmly among the hominines (African apes and humans in both Bonis [1983] and Begun [1992b]). Whether the African apes and humans have separate origins in Africa during the early/middle Miocene (Bonis & Melentis 1977a, b; Bonis 1983; Bonis & Koufos 1993) or a common origin in the middle Miocene of Europe (Bonis 1987; Bonis & Koufos 1994; Begun & Kordos 1997; Begun 2000, 2002, 2003, 2005, 2007a; Begun & Nargolwalla 2004), in both scenarios Europe is central to hominine origins as the place in which they evolved and pre-sumably acquired their diagnostic characteristics before returning to Africa.

Louis de Bonis and George Koufos have made a strong case for the phyletic link between Our-anopithecus and Australopithecus (Bonis & Koufos 1993, 1994, 1997, 2001, 2004; Koufos 1993, 1995, 2007; Bonis et al. 1998; Koufos & Bonis 2004). Important characters interpreted as synapo-morphies of an Ouranopithecus-Australopithecus clade include a relatively vertical face, narrow and convex mandibular condyle, distal accessory cusps on the upper and lower M3, rounded P3 lacking a buccal honing facet, relatively small male upper canine more rounded in cross section at the cervix, small deciduous canines, and molarized deciduous premolars (Koufos 2007). Th is hypothesis is rep-resented in Figure 2A.

Begun (2007b) and Begun & Kordos (1997) argue that all of the similarities between Ourano-pithecus and Australopithecus are homoplasies because, 1), this is consistent with the most parsimonious cladogram (Begun et al. 1997; Begun 2001), and 2), they are all related to a single functional com-plex, powerful mastication. Interestingly, Bonis & Melentis (1977a, 1978) suggested that Ourano-pithecus might be closely related to Gigantopithecus based on many of the same characteristics. Reduction

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FIG. 2. — Cladograms depicting alternative hypotheses discussed in the text. Modifi ed from Begun et al. (1997) and Begun (2001, 2002, 2007). Griphopithecus alpani Tekkaya, 1974 and Kenyapithecus kizili Kelley, Andrews & Alpagut, 2008 are thickly enameled middle Miocene hominoids from Paşalar (Turkey). Griphopithecus darwini Abel, 1902 is the type species, from Devínská Nová Ves (Slovakia). Neopithecus brancoi Schlosser, 1901 is an isolated M3 that is most similar to Rudapithecus Kretzoi, 1969 but with insuffi cient anatomy preserved to justify synonomy. Other taxa are discussed in the text. In A, Ouranopithecus is a hominin; in B, it is a dryopithecin.

Proconsul

Proconsul

A

B

?

Griphopithecus alpani

Griphopithecus darwini

Griphopithecus alpani

Griphopithecus darwini

Kenyapithecus kizili

Kenyapithecus kizili

Pongo

Pongo

Hominini

Hominini

Ouranopithecus macedoniensis

Ouranopithecus macedoniensis

New genus from Turkey

New genus from Turkey

Graecopithecus freybergi

Graecopithecus freybergi

Pan

Pan

Gorilla

Gorilla

Dryopithecus fontani

Dryopithecus catalaunicus

Dryopithecus fontani

Dryopithecus catalaunicus

Rudapithecus hungaricus

Rudapithecus hungaricus

Neopithecus brancoi

Neopithecus brancoi

Hispanopithecus crusafontiHispanopithecus laietanus

Hispanopithecus crusafonti

Hispanopithecus laietanus

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In both cases the nomen Rudapithecus hungaricus was used, but without a diagnosis or type specimen designated, and with no taxonomically useful char-acters described. In 1969 a publication appeared with a short diagnosis (Kretzoi 1969). Th us the fi rst available nomen for the sample from Rudabánya in my view is Rudapithecus hungaricus Kretzoi, 1969 (Kordos & Begun 2001). In Kretzoi (1975) a new taxon, Bodvapithecus altipalatus Kretzoi, 1975 is named. Kretzoi interpreted Bodvapithecus Kretzoi, 1975 to be related to Sivapithecus, and Rudapithecus to Ramapithecus (Kretzoi 1975), and this hypothesis was widely accepted (Pilbeam 1979; Szalay & Delson 1979; Wolpoff 1980). Kay & Simons (1983) were the fi rst, to my knowledge, to recognize that the smaller of the two hominid taxa from Rudabánya, Rudapithecus, had close affi nities to Dryopithecus. In subsequent years most researchers have interpreted Rudapithecus and Bodvapithecus to be females and males respectively of the same taxon, Dryopithecus brancoi Schlosser, 1901 (Begun & Kordos 1993). As noted earlier, here this sample is now returned to Rudapithecus.

Th e Rudabánya hominid sample includes three partial crania, two of which preserve signifi cant portions of the brain case. One of these specimens, RUD 200, preserves the connection between the brain case and face, and provides our only glimpse of the architecture of the cranium of a European hominid. Th is specimen is associated with a man-dible, RUD 212, making it the only well preserved skull (cranium and mandible) of a Eurasian hominid (a facial skeleton and mandible is known for Siva-pithecus, but no neurocranium, and a crushed and distorted skull of Oreopithecus is also known).

Th e craniodental sample of Rudapithecus is criti-cal in interpretations of hominine relations. Begun (2005) and Kordos & Begun (2001) list 16 charac-ters of the dentition and cranium shared between Rudapithecus and extant great apes, and 15 ad-ditional characters shared exclusively with extant hominines. In contrast to the characters shared between Ouranopithecus and Australopithecus, these 31 characters come from throughout the cranium and dentition. Th ere is no obvious functional inter-relationship among such hominine characters as a bi-convex premaxilla, supraorbital torus, frontal sinus,

and phylogenetic boundaries. Th e very similar morphology of the Pan and Australopithecus lower face exists despite dramatic diff erences in functional morphology and probably diet. In addition, there is evidence of de-coupling of premaxillary length/orientation and incisive fossa/canal morphology in Ankarapithecus, which shares the former with Sivapithecus and the latter with hominines (Be-gun & Güleç 1998). Ouranopithecus also lacks the spatulate upper lateral incisor of Pan and all Ho-minini, and in this way also shares the primitive morphotype with Gorilla, other dryopithecins and all other anthropoids (Begun et al. 1997). Begun (2002) lists a number of additional characters shared between Pan and Hominini that are not found in Ouranopithecus, and would have to have evolved independently if Ouranopithecus were more closely related to hominins than Pan. Th ese include lower crowned, fl ared molars, less robust lateral orbital pillars, more strongly developed supraorbital tori and sulci, a more horizontal frontal squama, and better developed supraorbital and glabellar pneu-matization. Unlike the characters that are shared by Ouranopithecus and Australopithecus that are all seemingly related to a single functional constraint (heavy mastication), the characters shared by Pan and hominins that are not found in Ouranopithecus are not obviously related to a single functional complex, and cannot easily be explained as a consequence of a single, simple and common adaptive response. For these reasons, I consider Ouranopithecus to be in the outgroup of the extant hominines (Fig. 2B). In the phylogenetic analysis on which Figure 2B is based (Begun et al. 1997; Begun 2001), it is equally parsimonious to place Ouranopithecus as the sister clade to the African apes and humans as it is to link it with dryopithecins as a group that is the sister clade to African apes and humans, as represented in Figure 2B.

RUDABÁNYA, HUNGARY

In 1967 an anonymous newspaper announcement was made describing the discovery of an ape jaw from deposits surrounding the large open pit ore mine at Rudabánya, in northern central Hungary. In the same year the discovery was covered in a popular science magazine article (Tasnádi Kubacska 1967).

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fused articular and tympanic temporal, elongated neurocranium and klinorhynchy. Where known, these characters are for the most part also shared with Ouranopithecus. One signifi cant exception is the position of the root of the zygomatic process of the maxilla, which is low in Ouranopithecus and primitive hominoids such as Proconsul Hopwood, 1933, and high in Rudapithecus and most other hominids (Begun & Kordos 1997). However, this is a good example of the exception that proves the rule. Th e low position of the root of the zygo-matic process in Ouranopithecus is not only shared with Proconsul and Hylobates, but also with robust australo pithecines (Begun & Kordos 1997b; Begun 2007b). In the case of Ouranopithecus and Paran-thropus, the morphology of the zygomatic has been related to elevated levels of stress generated by their greatly enlarged masticatory apparatus (Rak 1983; Begun & Kordos 1997b; Begun 2007b). While low on the alveolar process, the zygomatic bone in robust australopithecines and Ouranopithecus is very diff erent in morphology from the same bone in Proconsul and Hylobates. I would interpret the apparently primitive position of the zygomatic root in Ouranopithecus as a clear homoplasy, given the known phylogeny of extant hominoids, the inferred phylogeny of fossil apes, and the probable functional integration of this character within the heavy mastication morpho-functional complex (Begun 2007b).

Th e postcrania of Rudapithecus also reveal a suite of great ape affi nities. Th e distal end of a humerus and the proximal end of an ulna show the typical suite of great ape characters (large trochlea and capitulum well diff erentiated from each other by a deep zona conoidea, well-developed coranoid, radial and olecranon fossae, large, medial oriented medial epicondyle, strong ulnar trochlear keel, large radially oriented radial notch, robust ulnar shaft [Morbeck 1983; Begun 1992c]). Several carpal bones are known from Rudabánya. Th e scaphoid most closely resembles Pongo Lacépède, 1799, and like Pongo and nearly all non-hominine primates, it is not fused to the os centrale (Kivell & Begun 2009). Th e capitate is also Pongo-like but retains primitive characters found in some cercopithecoids (Kivell & Begun 2009). Th e distal end of a metacarpal closely

resembles those of Hispanopithecus in being strongly constricted dorsally, with large collateral ligament pits. A talus has a typical hominid morphology with a broad, shallow body and an elongated neck, most reminiscent of Pongo. Th e phalanges are typical of suspensory hominoids and are long and strongly curved, with well-developed fl exor sheath ridges and a proximally oriented proximal articular surface (Begun 1993; Deane & Begun 2008).

VALLÈS PENEDÈS, SPAIN

In 1944 Villalta & Crusafont described a new genus of hominoid from the Vallès Penedès of Catalonia, northeastern Spain. Th e type specimen of Hispano-pithecus laietanus comes from La Tarumba, but the vast majority of the sample attributed to this taxon comes from Can Llobateres (Begun & Moyà-Solà 1990; Golpe Posse 1993). A number of other taxa have been recognized based on the Can Llobateres sample (reviewed in Simons & Pilbeam [1965]), but only Hispanopithecus is recognized today.

As noted earlier, a major advance in our under-standing of European hominine phylogeny and adaptation occurred with the discovery and analysis of a craniofacial specimen and partial skeleton of Hispanopithecus from Can Llobateres (Begun & Moyá-Solá 1992; Moyà-Solà et al. 1992; Moyà-Solà & Köhler 1993, 1995, 1996; Begun 1994; Köhler et al. 2002). Details of the diff erences be-tween Hispanopithecus and Rudapithecus are listed above. In general, Hispanopithecus is smaller than Rudapithecus, though they overlap in size. Males of Hispanopithecus are intermediate in size between males and females of Rudapithecus. Th e vertebral column is known only in Hispanopithecus. Th e lumbar vertebrae show the important hominid character of a dorsal position of the transverse proc-ess, a feature that is associated with the short, stiff lower backs and orthogrady of great apes (Ward 1993, 1997, 2007; MacLatchy et al. 2000). Other great ape characters of the postcranium of Hispano-pithecus include a large lunate and robust hamate, the latter with a well-developed hamulus, and large, long and curved manual proximal phalanges with well-developed fl exor sheath ridges. Although the important great ape character of a reduced ulnar styloid process, resulting in the absence of ulnar-

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Th e key is the size and position of the sinus. In dryopithecins, African apes, and humans, the sinus occupies most of the interorbital space from below nasion to glabella, and has a variable pneumatization into the frontal squama. Th is has been interpreted as a true ethmoidal frontal sinus, given the large involvement of the interorbital space and hence the ethmoidal air cells (Begun 1994, 2007a). Siva-pithecus and Pongo lack expansion of the ethmoidal air cells, which invade neither the interorbital region nor the frontal squama. On the other hand, there are many taxa in which the frontal is pneumatized from behind, by the sphenoidal air sinus (Cave & Haines 1940). Th ere are diff ering interpretations of the signifi cance of various paranasal sinus con-fi gurations (Begun 1994; Rossie et al. 2002; Rossie 2005), but researchers agree that the loss of the ethmoidal and frontal sinus expansion in Pongo and Sivapithecus is an important synapomorphy of that clade (Ward & Brown 1986). Hispanopithecus clearly does not share this synapomorphy with Asian great apes (Fig. 3).

Th e frontozygomatic suture in IPS 18000 is said to be in a depressed position relative to the superior orbital margin, which is a tendency seen in Pongo (Moyà-Solà & Köhler 1995; Köhler et al. 2001). In Moyà-Solà & Köhler (1995: 123, fi g. 17) it is clear that there is very substantial overlap between Pan and Pongo, with Hispanopithecus values falling in the range of overlapping values. Furthermore, they use internal biorbital breadth to standardize for body mass, despite the fact that Pongo and Sivapithecus have relatively smaller biorbital breadths given their narrow interorbital regions and their tall, narrow orbits. Figure 3 shows a more likely position of the zygomatic relative to the frontal in IPS 18000. Insets b and c show views of the frontozygomatic suture surfaces. On the zygomatic bone the surface is nearly complete. On the frontal it is preserved but somewhat eroded. Th is corresponds to the description of the bones in Moyà-Solà & Köhler (1995). When the mirror imaged frontal with its frontozygomatic suture surface is matched to the right zygomatic with its suture surface, the posi-tion of the suture itself is clear. It is relatively high on the orbit as in most anthropoids, and lacks the condition found in Sivapithecus and some Pongo.

triquetral contact, is not known for either Hispano-pithecus or Rudapithecus, the absence of a facet for the ulnar styloid on the triquetrum of Dryopithecus catalaunicus strongly suggests that the same was true for the Vallesian taxa.

Moyà-Solà and colleagues have concluded that Hispanopithecus is a member of the Sivapithecus and Pongo clade (Moyà-Solà & Köhler 1993, 1995, 1996; Köhler et al. 2001). Th ey base this conclu-sion on a small number of characters of the face, including the size and position of the zygomatico-facial foramina, the shape and orientation of the zygomatic bone, the position of the frontozygomatic suture, the size of the frontal sinus and the presence of supraorbital costae.

Th e number and size of the zygomaticofacial foramina are highly variable in extant anthropoids (Msuya & Harrison 1996). Pongo does tend to have a higher number of foramina than other great apes, but there is signifi cant overlap. In addition, it is dif-fi cult to say exactly how many foramina are present in IPS 18000 given preservation in the area (Fig. 3). As Figure 3 shows, the position and orientation of the zygomatic bone is unclear as it is not attached to either the maxilla or the frontal bone. Th e small part of the body of the zygomatic bone that is preserved inferolateral to the orbital margin is fl attened, and resembles Pongo in this regard, but so little of the bone is preserved that it is not possible to describe it reliably as overall Pongo-like. In addition, in the case of both the zygomaticofacial foramina and the orientation of the zygomatic bone, three specimens of Rudapithecus lack the Pongo-like morphology that has been suggested for IPS 18000. RUD 44, 77 and 200 all preserve more of the zygomatic bone, and they have neither large and supernummary zygomaticofacial foramina nor a fl at and anteriorly facing zygomatic body.

Inset a in Figure 3 is a view of the partially pre-served frontal sinus of IPS 18000. It is not reduced, as described in Köhler et al. (2001). In fact, it is quite large, and occupies most of the interorbital space and indeed it does invade the space between the endocranial and orbital plates. In dryopithecins this is the typical pattern, which is only found in African apes and humans. It departs most mark-edly from the pattern in Pongo and Sivapithecus.

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Th e orbits are also squared and the interorbital space large, unlike Sivapithecus and Pongo.

Th e presence of supraorbital costae is also said to unite Hispanopithecus with the Sivapithecus/Pongo clade (Moyà-Solà & Köhler 1995). Figure 3 shows the disposition of ridges or costae in the midfrontal region. Supraorbital costae appear on Pongo crania, and in Sivapithecus, and consist of ridges that rim the superior orbital margin and the upper part of the medial orbital margins. Th ey do not meet in the midline, nor do they reach glabella or attach to any eminence at glabella. In contrast, supra orbital tori meet in the midline at glabella, and are usually continuous with an eminence at glabella (Begun 1994). In Figure 3 highlights mark the path of two

tori emanating from a mildly infl ated glabella, both running superolaterally. Superior to the glabella-tori complex is a deep biconcave depression, which is unique to Hispanopithecus (although such a de-pression is suggested by the morphology of Xir-1 [Ouranopithecus], this is not clear due to damage). In addition to the supraorbital tori, there are su-perior orbital margins but they are not thickened and do not project anteriorly from the supraorbital surface as they do in Pongo and Sivapithecus. Th is plus the fact that they do not rim the orbits as described above means that they are unlikely to be homologous to the supraorbital costae of Asian great apes. In the end there are no characters shared uniquely between Hispanopithecus and Asian great

B

A

C

D

FIG. 3. — Views of IPS 18000 from Can Llobateres: A, frontal view of the periorbital region with the right side reconstructed based on a mirror image of the better preserved left side; B, view from below of the interorbital space of IPS 18000 showing the extensive frontal sinus, outlined on the right side; C, lateral view of the frontal fragment showing the frontal’s joint surface of the frontozygomatic suture; D, view from superiorly and medially showing the zygomatic’s joint surface of the frontozygomatic suture. Scale bar: 1 cm.

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size, lower crowned male canines, low crowned I1 with a highly complicated labial surface morpho-logy, extreme upper incisor heteromorphy, incisors nearly aligned with the canines (modifi ed from Güleç et al. [2007]).

Th e Çorakyerler sample shares characters with Ouranopithecus related to a heavily masticated diet, which is often associated with an adaptation to more open habitats than those of living great apes. Indeed, both genera occur in Eurasia at a time of considerable decline in forest cover, drier climates and increased seasonality (Bernor 1983; Bonis et al. 1994; Bernor et al. 1996; Solounias et al. 1999; Agustí et al. 2003; Begun & Nargol-walla 2004; Begun 2005; Merceron et al. 2005a; Koufos et al. 2006; Agustí 2007; Strömberg et al. 2007). As noted earlier, many of these characters occur in Pliocene hominins, but also in other fos-sil hominids with large jaws and thickly enameled teeth. Because the Çorakyerler sample has derived characters shared with African apes and humans not found in Ouranopithecus, it is considered diff erent at the genus level.

Th e Çorakyerler specimens are great apes, with affi nities to European and African taxa and not with Ankarapithecus or South or Southeast Asian taxa (Begun et al. 2003; Begun 2005; Güleç et al. 2007). Th e sample shows its greatest similarities to gorillas, chimpanzees and Ouranopithecus, and is intermediate in age between Ouranopithecus and the earliest known African hominids (Brunet et al. 2002; Vignaud et al. 2002; Güleç et al. 2007; Lebatard et al. 2008). Previous interpretations of Ouranopithecus had suggested this genus as a good candidate for Pliocene hominid ancestry based on derived dentognathic features shared with later Af-rican Pliocene Australopithecus (see above). While the upper premolars of CO 205 and non-robust Australopithecus strongly resemble one another, A. anamensis Leakey, Feibel, McDougall & Walker, 1995 and A. afarensis Johanson, White & Cop-pens, 1978 have more primitive postcanine teeth (White 1977; Ward et al. 2001; Güleç et al. 2007). Furthermore, all late Miocene African hominids (Orrorin Coppens, Senut, Pickford, Gommery, Mein & Cheboi, 2001, Ardi pithecus White, As-faw & Suwa, 1994, and Sahelanthropus Brunet et

apes and a large number of characters shared be-tween dryopithecins and African apes (Begun et al. 1997; Begun 2001).

TURKEY

In 2007 Güleç et al. published a new species of Ourano pithecus, O. turkae, from Çorakyerler, Tur-key. Th e Çorakyerler fossil assemblage is correlated to MN 11, or between 8.7 and 7.4 Ma (Güleç et al. 2007). I had the opportunity to examine and prepare the type specimen, a very large male palate. As noted elsewhere (Begun et al. 2003; Güleç et al. 2007), the specimen resembles Ouranopithecus more closely than any other fossil ape. Th e specimen is considerably larger than the largest Ouranopithecus specimen, and in fact it fi ts comfortably on the mandible of Indopithecus giganteus Pilgrim, 1915 (often referred to as Gigantopithecus bilaspurensis Simons & Chopra, 1969). Th ere are also important diff erences from Ouranopithecus, which justify in my view the re cognition of a new genus.

Th e Çorakyerler maxilla (CO 205) is a large presumed male palate with LI1-M3, RC-M2, pre-serving portions of the right palatine process to the midline, a small fragment of the left palatine proc-ess, most of the left alveolar process, and much of the palatal surface of the right premaxilla. Th ere is some distortion along both tooth rows, but a good estimate of dental arcade shape is possible (Fig. 4). Th e specimen preserves a portion of the right edge of the nasal aperture along the canine root. Th e left side preserves the base of the zygomatic proc-ess and the inferior extension of the canine fossa (Fig. 4). Th e canine and M3 are fully erupted and lightly worn.

Th e new genus from Turkey is distinguished from other hominoids including Ouranopithecus macedoniensis, by the following dentognathic charac-teristics: a short, vertical premaxilla and lower face: narrow palate relative to postcanine size: broad, higher zygomatic root: broad, diamond-shaped canine cross-section at the cervix: homomorphic P3 and P4: oval and symmetrical P3 with pre- and postparacrista of equal length: M3 larger than M2. In addition to the characters noted above, it is further distinguished from Ouranopithecus in the follow-ing characters: larger size, smaller relative canine

803



al., 2002) lack dentognathic specializations associ-ated with heavily masticated diets (Haile-Selassie 2001; Senut et al. 2001; Brunet et al. 2002; Haile-Selassie et al. 2004; Güleç et al. 2007). Evidence from functional morphology and known relations among fossil and living hominids strongly suggests that Ouranopithecus and the Çorakyerler hominine evolved many or all of these dietary adaptations in parallel with later African hominids (Begun & Kordos 1997b; Begun 2007b). On the other hand, the features the Çorakyerler hominine shares with African apes and humans and not Ouranopithecus may indicate a closer relationship to this larger clade (Fig. 2B). A number of characters are shared with Gorilla, and occasionally found in other hominids,

including narrow palates, reduced premolar occlu-sal outline and cusp heteromorphy, peg-shaped I2, short premaxilla and vertically implanted anterior teeth, and a thin maxillary palatine process. Some of these characters, such as the peg-shaped I2, are primitive for the African ape and human clade, as suggested on the basis of previous comparisons to Dryopithecus and Ouranopithecus (Begun 1994, 2001, 2002; Begun & Kordos 1997).

Th e presence of a fossil great ape in a relatively open habitat in the Turolian of Turkey will require a rethinking of ideas of Eurasian hominoid extinctions, previously attributed to environmental changes at this time (Bonis & Koufos 1999; Bonis et al. 1999; Solounias et al. 1999; Eronen & Rook 2004; Rook &

A

B

C D

FIG. 4. — Views of the Çorakyerler hominine: A, palatal; B, lateral; C, medial; D, frontal. Scale bar: 1cm.

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resemble living chimpanzees in molar occlusal morpho logy, enamel thickness, gross and microwear, and shearing quotients (Begun 1994; Ungar 1996; Begun & Kordos 1997b; Kay & Ungar 1997; Smith et al. 2003). Th is has been widely interpreted as an adaptation to a soft fruit diet. However, there are some diff erences between thinly enameled dryo-pithecins and chimpanzees. Th e incisors of the fossil taxa tend to be narrow compared to chimpanzees, and relatively tall crowned. It has been suggested that this may be a response to the need to process foods with tough protective coverings (Begun & Kordos 1997). Th e narrow mesiodistal length of the teeth may have served to generate higher more focused occlusal forces, and the crown height may be a response to wear. Strongly developed lingual marginal ridges and labial pillars may also have served to resist labiolingual bending and torsion. Th is interpretation is supported by a recent analysis of incisor crown curvatures that also links thinly enameled dryopithecins to hard object feeders (Deane 2007). Th e combination of a soft fruit frugivore signal from molar occlusal morphology, microwear, shearing quotients and enamel thickness and a hard object feeding signal from the incisors is unique and may indicate that hard object pro-cessing was part of a fall back feeding strategy that allowed dryopithecins to exploit a wider range of more diffi cult to process foods in times of resource shortage. Th e mandibular corpora of these thinly enameled dryopithecins are also somewhat more robust than in chimpanzees, which is consistent with this interpretation. Rudapithecus and Hispanopithecus localities are interpreted as closed forests but with a greater degree of seasonality than is typical for tropical or many subtropical forests today (Andrews et al. 1997; Agustí et al. 1999, 2003; Bernor et al. 2004; Rook et al. 2004; Agustí 2007; Merceron et al. 2007b). Th is suggests periods of resource shortage that would have made an eff ective fall back feeding strategy of paramount importance for survival. It is possible that the ability to exploit more embed-ded foods also represents a pre-adaptation that led eventually to a more extreme and committed hard object feeding strategy in Ouranopithecus.

As noted above, Ouranopithecus had a robust masticatory apparatus with all the classic indica-

Bernor 2004; Agustí 2007). Th e fauna associated with the Çorakyerler hominoids indicates a relatively open habitat (Güleç et al. 2007). Th e large, thickly enameled, low-cusped postcanine teeth are likely associated with a heavily masticated diet. However, the canine/pre molar complex in the Çorakyerler hominine, unlike Sahelanthropus, Orrorin or Ardip-ithecus, retains functional honing, at the same time that the male canines are reduced compared to the large, sharp, and prominently projecting canines of most other late Miocene Eurasian and extant great apes. Th e Çorakyerler hominine is the youngest and largest of all Turkish Miocene hominoids and dif-fers from all modern and fossil great apes in what is known of its anatomy and environment. Th e antici-pated recovery of additional cranial and postcranial remains at Çorakyerler will clarify the adaptations and phylogenetic status of this new genus.

BULGARIA

Spassov & Geraads (2008) describe a hominid P4 from a Turolian locality in the Chirpan district of southern Bulgaria. Th ey estimate the age of this fossil assemblage at about 7 Ma, which makes it most likely somewhat younger that the Çorakyerler homi nine. Spassov & Geraads (2008) indicate that the new Bulgarian specimen resembles Ouranopithecus. Th ey do not speculate on the ecological context of the new hominid from Bulgaria, though the presence of the proboscidean Anancus Aymard, 1855 and the spiral horned antelope Protragelaphus Dames, 1883 suggests perhaps a relatively open woodland environment (Cerling et al. 1999; Merceron et al. 2005b; Spassov & Geraads 2008), which would be broadly consistent with the paleoecology of Ourano pithecus localities (Bonis et al. 1999; Mer-ceron et al. 2007a).

ECOLOGICAL DIVERSITY IN EUROPEAN MIOCENE HOMININES

Th e large number of European Miocene hominine taxa bear witness to a broad range of dietary and positional behavior adaptations. In terms of diet there are two major patterns. Th e older dryopithecins, Dryopithecus, Hispanopithecus and Rudapithecus

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tors of a hard object diet. Th ese include hyperthick enamel, large postcanine teeth with low, rounded cusps, robust mandibles and powerfully developed attachment sites for the muscles of mastication (Bonis & Koufos 1993, 1994, 1997; Begun & Kordos 1997b; Begun 2007b). Th is is supported by dental microwear analyses as well (Ungar & Kay 1995; Ungar 1996; Merceron et al. 2005c). Ouranopithecus also appears to have some reduction in the size of the canines, at least in the maxilla, which leaves more room in the jaw for enlarged postcanine teeth. Th e anterior teeth are often very strongly worn, suggesting intensive anterior dental processing, which is often associated with embedded foods. Th e paleoecology of Ouranopithecus locali-ties is widely regarded as more open than earlier dryopithecins (Bonis et al. 1999), and this is also consistent with a feeding strategy emphasizing embedded foods, with perhaps a greater percentage coming from terrestrial sources.

Th ere is an interesting functional parallel between the dryopithecins and crown hominines (African apes and fossils and living humans). Both clades are very diverse and speciose when all fossil taxa are considered. And both clades run the spectrum from soft fruit frugivores (e.g., Rudapithecus, Pan) to extreme hard object feeders (Ouranopithecus, Paranthropus). Th is could be revealing of an emer-gent property of cranial biology in hominines. Hominine cranial form and possibly other adapta-tions such as brain size channel a radiation of taxa that tended to fi ll the same niches independently, fi rst in Europe in the middle to late Miocene, and later in Africa in the late Miocene and Pliocene (Begun & Kordos 1997).

Diversity in positional behavior in dryopithecins is less well documented, mainly because postcrania are not described for Ouranopithecus. Hispano-pithecus and Rudapithecus are both interpreted as highly arboreal (Begun 1988, 1992b, c, 1994, 2007b; Moyà-Solà & Köhler 1996; Begun & Ko-rdos 1997; Almécija et al. 2007). Interpretations diff er however on the precise nature of the arboreal locomotion in the two genera. Hispanopithecus is thought by some to have been a strong climber with well-developed characters related to orthogrady and suspension while retaining features indicative

of palmigrady (Almécija et al. 2007). Like Rudap-ithecus, Hispanopithecus has long, strongly curved phalanges with strongly developed fi brous fl exor sheath ridges, all of which is associated in living primates with a specialized adaptation to suspensory positional behavior (Susman 1979; Begun 1993; Richmond & Whalen 2001; Deane & Begun 2008). However, the metacarpal heads in Hispanopithecus are strongly constricted dorsally, with deep collat-eral ligament pits expanded dorsally. Th e proximal articular surfaces of the proximal phalanges are said to be oriented more dorsally than in modern hominoids. Th ese characters and the presence of short metacarpals are interpreted to indicate that the hand postures of Hispanopithecus included frequent hyper extension at the metacarpo-phalangeal joint, which is a feature of palmigrade primates (Almécija et al. 2007). However, palmigrade monkeys do not have metacarpal heads as dorsally constricted as Hispanopithecus. Hominoids, in contrast, lack these joint characteristics and, given the great develop-ment of their fl exor musculature, they have limited mobility in extension (Tuttle 1969). Th e metacarpal heads also lack the typical palmar grooves, or fl ut-ing, of the joint surface proximally that are found in most palmigrade monkeys that are related to the presence of sesamoid bones (Lewis 1977; Almécija et al. 2007). While the proximal articular surfaces of the proximal phalanges have some dorsal expan-sion, they are large, deep and rounded, and extend palmarly between the two large palmar tubercles, again unlike the condition in palmigrade mon-keys in which they tend to be broad, shallow and in a more dorsal position. Th e proximal end of the proximal phalanges are clearly tilted palmarly and do not really face dorsally to any signifi cant extent. Th e shortness of the metacarpals relative to the phalanges is also not clear cut. Comparisons are limited to the fourth digital ray, in which the metacarpal is in two pieces. Although the authors indicate that the length can be easily reconstructed, the result is a fourth metacarpal that is signifi cantly shorter than the second metacarpal, much more so than in extant anthropoids, except hylobatids, which otherwise have very slender metacarpals. It is also very unusual to see a large second metacar-pal and small second proximal phalanx compared

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strong suspensory signal (Morbeck 1983; Begun 1988, 1992c, 1993, 1994; Begun & Kordos 1997; Deane & Begun 2008). Preliminary analysis of the femora, pelvis and hand bones is consistent with this interpretation. In most cases the great ape that Rudapithecus most closely approaches is Pongo.

Dryopithecus (including Pierolapithecus) is rep-resented by an exquisite skeleton including well-preserved ribs, vertebrae and hand bones (Moyà-Solà et al. 2004). Th e ribs point to a broad, shallow hominoid-like thorax and the carpals are great ape like in the absence of contact between the triquetrum and ulna. Th e vertebrae are intermedi-ate in the position of the transverse processes and the phalanges are shorter and less curved than in Hispano pithecus (Moyà-Solà et al. 2004). Moyà-Solà et al. (2004) interpret this mixture of characters to refl ect a unique combination of behaviors, including powerful climbing and orthogrady and palmigrade hand posture, but not suspension. Begun & Ward (2005) suggest that the vertebral and phalangeal characters of Dryopithecus catalaunicus do not pre-clude suspension, and the strongly developed sec-ondary shaft characters of the phalanges are most consistent with suspension, despite the somewhat reduced length compared to Hispanopithecus. It is noteworthy that many Old and New World monkeys practice suspensory positional behaviors, though perhaps not as adeptly as hominoids, and in some cases with the aid of a prehensile tail, despite lack-ing the hominoid-like characters of Dryopithecus. Either way, Dryopithecus is the oldest hominid with well-preserved and unambiguous evidence of a great ape body plan (Moyà-Solà et al. 2004).

AFRICAN LATE MIOCENE HOMININES AND THE PALEOBIOGEOGRAPHYOF HOMININE ORIGINS

As noted earlier, there is agreement among many researchers that the dryopithecins are hominines, even if there is some disagreement on their precise relationships to crown hominines. Th ere are, how-ever, hominines in the African late Miocene, and some suggest that these evolved not from European hominines but in situ in Africa. Th e European taxa

to the fourth digital ray. In hylobatids the fourth ray proximal phalanx is shorter and smaller in ar-ticular dimensions than the second. If the fourth meta carpal were really as short as is indicated in Almécija et al. (2007), one would expect a similar pattern with the proximal phalanges.

While the metacarpal robusticity and phalangeal relative size are uncertain in Hispanopithecus, the morphology of the metacarpal heads and the proxi-mal articular surfaces of the proximal phalanges, which are neither monkey-like nor ape-like, may suggest a unique feature of the positional repertoire of Hispanopithecus. Frequent palmigrady would sug-gest the need to stabilize the metacarpo-phalangeal joint in hyperextension, which should lead to dorsal expansion of the distal metacarpal articular surface and collateral ligament pits that are more palmarly displaced. Th e opposite is the case in Hispanopithecus. Large dorsal collateral ligament pits would cause the collateral ligaments to become taut and bring the joint into maximum close packed position in a relatively wide range of fl exion, as expected in a suspensory animal. Th e joint would be relatively loose in extension. Instead of palmigrady, the un-usual morphology of the metacarpo-phalangeal joint may simply refl ect the need to stabilize the joint in a wide range of fl exed postures.

Th e other especially informative aspects of the postcrania of Hispanopithecus are the femur and lumbar vertebrae. Th e best preserved vertebrae show the typical great ape character of a dorsally displaced transverse process, which has been related to the presence of a short, stiff lower back (Moyà-Solà & Köhler 1996; Ward 2007). Th e well-preserved femur has features of the hip joint related to suspension and high hip mobility (Köhler et al. 2002). In sum, the postcranial skeleton of Hispanopithecus preserves much evidence of a highly arboreal, climbing, sus-pensory hominoid. Possible retention of relatively frequent palmigrady has been suggested but the evidence in my view is inconclusive.

Rudapithecus from Rudabánya is known from a number of phalanges, hand and foot bones, the distal end of the humerus, proximal ends of a radius and ulna, two partial femora and a partial pelvis. Th e phalanges, humeral, radial and ulnar specimens are very similar to extant great apes with a clear

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are either dismissed as non-hominine or consid-ered to have dispersed into Europe in the middle Miocene from an unknown middle Miocene Af-rican ancestor (Rook & Bernor 2004; Pickford & Senut 2005; Kunimatsu et al. 2007; Suwa et al. 2007). Th e evidence of the hominine affi nities of the dryopithecins is actually quite strong, both from the analysis of Ouranopithecus and the Dryopithecus-Hispanopithecus-Rudapithecus group (see above and the numerous references cited). Th e real question is not whether European late Miocene hominids are hominines, but, do the European hominines represent one or more side branches of hominine evolution, having dispersed into Europe one or more times from unknown ancestors, or is the much larger record of Miocene hominines in Europe the true source of later hominines that dispersed into Africa in the late Miocene? Th ere is strong evidence for the latter hypothesis in data from the morphology, geochronology and paleobiogeography of European hominines and other land mammals (Begun 2001, 2005, 2007a; Begun & Nargolwalla 2004; Nargol-walla 2009). Th e morphological evidence is reviewed above. Hominines occur in Europe more than 2 million years before they appear in Africa. Th ey are more widespread and diverse than African hominines. Many land mammals are known to have expanded their ranges between Africa and Europe in the late Miocene, during the temporal interval between the fi rst appearance of Vallesian European hominines and the fi rst appearance of African hominines, including many taxa with preferences for forests (Fortelius et al. 1996; Leakey et al. 1996; Dawson 1999; Ginsburg 1999; Heissig 1999; Made 1999; Solounias et al. 1999; Agustí et al. 2001; Winkler 2002; Lehmann et al. 2006; Agustí 2007; Likius et al. 2007; Nargolwalla 2009). Th ese dispersals or range expansions are widely believed to be related to changing climatic conditions in Europe, including increases in seasonality and a decline in forest cover (Bernor et al. 1979; Bernor 1983; Quade et al. 1989; Fortelius et al. 1996; Leakey et al. 1996; Agustí et al. 1999, 2001, 2003; Bonis & Koufos 1999; Cer-ling 1999; Cerling et al. 1999; Magyar et al. 1999; Solounias et al. 1999; Fortelius & Hokkanen 2001; Koufos 2003; Eronen & Rook 2004; Fortelius et al. 2006; Agustí 2007). Th e claim that the ecology

of the most likely dispersal route between Europe and Africa, the eastern Mediterranean, was already too inhospitable for apes (Rook & Bernor 2004), is misleading for several reasons. Firstly, the drying that is typical of the broadly defi ned “Pikermian biome” begins to show an impact on environments after 10 Ma, and it was not until 8-9 Ma that the eff ects become pronounced (Solounias et al. 1999; Agustí et al. 2003; Eronen & Rook 2004; Fortelius et al. 2006; Agustí 2007). Th ere is no paleoecologi-cal evidence to suggest that the eastern Mediter-ranean was inhospitable to forest living primates before 10 Ma. Secondly, many recent analyses question the overall “savanna-like” quality of the eastern Mediterranean in the late Miocene, and several stress the persistence of forest even after 10 Ma, despite the overall drying trend. Akgün et al. (2007) identify mixed mesophytic and swamp forest assemblages in western Anatolia in the late Miocene. Bruch et al. (2006) found evidence for a warm and humid climate overall in the Pannonian basin and well into the Balkans in many fossil plant localities until 9 Ma, with no indication of a Medi-terranean climate. Pickford et al. (2006) identify a humid climate in the southern Mediterranean be-tween 10-11 Ma. Fortelius et al. (2006) produced palaeoprecipitation maps for the late Miocene, through to about 9.5 Ma, which show very little regional diff erentiation between central Europe and the eastern Mediterranean. Ivanov et al. (2002) and Griffi n (2002) describe fl uctuating cycles of warm/humid and cooler/dryer conditions in the northern Balkans in the late Miocene. Finally, the ape that dispersed from Europe to Africa is undiscovered at present, and therefore we have no knowledge of its ecological preferences, but if it was like many living primates it was probably capable of surviv-ing in a diversity of environments. Th us, there is no reason a priori to believe that hominines could not have taken part in the general dispersal of land mammals between Europe and Africa during the late Miocene, and in fact Darwin speculated 150 years ago that this may well have taken place.

Th ere is a good chance that African hominines evolved from a European ancestor. What is the nature of this ancestor and the dynamics of early homi-nine evolution in Africa? While the broad picture

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microCT analysis revealing several characters of the enamodentine junction (EDJ) that support this al-location. Pickford et al. (2009) recently described as hominoid a very poorly preserved mandibular specimen from Niger with an estimated age of 5-11 Ma. Th is specimen, along with other isolated teeth from Kenya in the same temporal range (Pickford & Senut 2005), are too poorly preserved to be assigned to any known hominoid clade with any confi dence.

Chororapithecus Suwa, Kono, Katoh, Asfaw & Beyene, 2007, represented by a handful of isolated teeth from 10-10.5 Ma deposits in Ethiopia, is said to possibly be an early member of the Gorilla clade (Suwa et al. 2007). Th e overall morphology of the molars resembles many thickly enameled late Miocene hominid molars, but the EDJ, while mostly fl at like most hominids with thick enamel, has a subtle cresting pattern that Suwa et al. (2007) interpret to be homologous to a much better devel-oped crest on gorilla molars. Again, the connection to Gorilla is possible, though much more data are needed to make a strong case for this relationship. Chororapithecus is at least 2 Ma younger than Dryo-pithecus, and does not therefore directly challenge the hypothesis that hominines fi rst appear in Europe. However, it does present an interesting potential scenario for the evolution of thinly enameled teeth from a thickly enameled ancestor. Rook & Bernor (2004) consider it unlikely that Ouranopithecus could be ancestral to Aridpithecus, the latter hav-ing thin molar enamel, although morphologically the teeth are broadly similar to one another and to australopithecines generally. Th e transformation from a Chororapithecus dental morphology to that of the extant gorilla is much more dramatic, and is a cautionary tale about what it is wise to consider likely or unlikely a priori in evolutionary history.

CONCLUSIONS

Th is review stresses the phyletic unity and ecologi-cal diversity of European middle and late Miocene hominines, and explicitly recognizes the pioneer-ing eff orts of Louis de Bonis in providing strong evidence for decades to support the Afro-European

of paleoecology in the eastern Mediterranean may favor the view that a more open woodland taxon such as Ouranopithecus or a relative is more likely to have expanded its range from Europe to Africa than a forest form such as Rudapithecus, there were opportunities for expansion by forest dwelling taxa as well (Nargolwalla & Begun 2009). Th e small but growing record of hominines from Africa does not help resolve this question. Nakalipithecus Kunimatsu et al., 2007, a hominine candidate from 9.8 Ma deposits in Kenya, is similar in many features to Ouranopithecus (Kunimatsu et al. 2007). Th e main specimen, a mandible, preserves the molars but they are very strongly worn, with indications of partial buccal cingula, unlike Ouranopithecus and more like the Proconsul-like molars from 10.5 Ma deposits at Ngorora (Hill & Ward 1988). However, an upper female canine is very similar to Ouranopithecus, es-pecially in the morphology of the cervix and lingual aspect (Kunimatsu et al. 2007). Kunimatsu et al. (2007) suggest that Nakalipithecus may be ances-tral to Ouranopithecus, given its slightly older age and more primitive morphology. Th is may be the case, although more fossils are needed to eff ectively distinguish Nakalipithecus from the late surviving proconsulids from Ngorora. Even if Nakalipithecus is broadly ancestral to Ouranopithecus, they are so close in known age (9.8 vs. 9.6 Ma) that they should be considered essentially contemporaneous, and either one could have originated fi rst. In addition, range expansions of late Miocene hominines are likely to have been complex and multidirectional, with expansions and contractions contributing to complex patterns of speciation and extinction. Whatever the relationship is between Ourano-pithecus and Nakalipithecus, the oldest hominine is still Dryopithecus, from localities in Europe that are more than 2 Ma older.

Samburupithecus Ishida & Pickford, 1997 has also been proposed as an early hominine based on its overall robust morphology and thickly enameled molars (Ishida & Pickford 1997). However, Begun (2007a) notes that Samburupithecus retains many primitive characters with Proconsul that are lack-ing in hominids, concluding that it is most likely to be a late surviving member of the Proconsuloi-dea. Olejniczak et al. (2009) recently conducted a

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Miocene hominine connection. Diversity in Euro-pean Miocene hominines parallels that of Pliocene to recent hominines, with a range of ecological preferences from suspensory, highly arboreal soft fruit frugivores living in closed forests, to hard ob-ject feeders, possibly much more terrestrial, living in more open settings.

Th e conclusion that hominines originate and experience their initial adaptive radiation in Europe is signifi cant in setting the ecological and temporal context for the evolution of our subfamily. Homi-nines and pongines are likely to have evolved from early middle Miocene thickly enameled taxa, such as Griphopithecus from Europe and Western Asia or a close relative from later localities in Africa (Nachola pithecus Ishida, Kunimatsu, Nakatsukasa & Nakano, 1999, Equatorius Ward, Brown, Hill, Kel-ley & Downs, 1999, Kenyapithecus Leakey, 1962). While it may seem unnecessarily complicated to suggest that multiple dispersals and/or range exten-sions occurred between Africa and Eurasia during the evolution of the hominids before the hom-inines disperse from Europe to Africa a fi nal time in the late Miocene (Begun 2002, 2005, 2007a; Andrews & Kelley 2007), this is the scenario pro-posed for several taxa including aardvarks, several carnivores, antelopes, hippos and probably at least one lineage of proboscidean (Leakey et al. 1996; Ginsburg 1999; Heißig 1999a, b; Made 1999; Boisserie et al. 2003; Werdelin 2003; Begun & Nargolwalla 2004; Lehmann et al. 2006; Likius et al. 2007; Koufos & Bonis 2008).

Th e major changes experienced by both the hominines and the pongines in Eurasia include diversifi cation of positional behavior and the de-velopment of the extant hominid suspensory body plan, specialization of dietary adaptations, increases in overall body and brain size, and an overall slow-ing of life history (Begun & Kordos 2004; Kelley 2004; Russon & Begun 2004). Th e one aspect of hominid biology that unites all of these charac-teristics is behavioral fl exibility and the ability to cope with diverse and changing ecological condi-tions (Potts 2004; Russon & Begun 2004). Th ese adaptations fi rst appear in Eurasia in response to Eurasian ecological conditions, and they are likely to have permitted hominids to survive and adapt

to changing conditions in the late Miocene, and disperse into Southeast Asia and Africa. Specifi -cally, the adaptations that developed in European hominines persist in extant hominines, and set the stage for the development of further changes in positional behavior, diet and eventually brain size that characterize the evolutionary history of the hominins.

Acknowledgements I am grateful to Stéphane Peigné and Gildas Mer-ceron for asking me to contribute to this volume honoring the career of Louis de Bonis. Th anks as well to Gildas Merceron, George Koufos and an anonymous reviewer who provided many useful comments. I thank the many curators who have al-lowed me to study the fossils in their care, especially Louis, who hosted me during my fi rst visit to examine European Miocene fossils, in 1984. His generosity and patience with an upstart graduate student will always be fondly remembered. I wish him much continued success in all his endeavours.

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