Buronius manfredschmidi-A new small hominid from the early late Miocene of Hammerschmiede (Bavaria, Germany).

The known diversity of European middle and late Miocene hominids has increased significantly during the last decades. Most of these great apes were frugivores in the broadest sense, ranging from soft fruit frugivores most like chimpanzees to hard/tough object feeders like orangutans, varying in size from larger than siamangs (over 17 kg) to larger than most chimpanzees (~60-70 kg). In contrast to the frequent sympatry of hominoids in the early-to-middle Miocene of Africa, in no European Miocene locality more than one hominid taxon has been identified. Here we describe the first case of hominid sympatry in Europe from the 11.62 Ma old Hammerschmiede HAM 5 level, best known from its excellent record of Danuvius guggenmosi. The new fossils are consistent in size with larger pliopithecoids but differ morphologically from any pliopithecoid and from Danuvius. They are also distinguished from early and middle Miocene apes, share affinities with late Miocene apes, and represent a small hitherto unknown late Miocene ape Buronius manfredschmidi. With an estimated body mass of about 10 kg it represents the smallest known hominid taxon. The relative enamel thickness of Buronius is thin and contrasts with Danuvius, whose enamel is twice as thick. The differences between Buronius and Danuvius in tooth and patellar morphology, enamel thickness and body mass are indicative of differing adaptations in each, permitting resource partitioning, in which Buronius was a more folivorous climber.

Comment on "Origin of human bipedalism as an adaptation for locomotion on flexible branches".

Thorpe et al. (Reports, 1 June 2007, p. 1328) concluded that human bipedalism evolved from a type of bipedal posture they observed in extant orangutans with seemingly human-like extended knees. However, humans share knuckle-walking characters with African apes that are absent in orangutans. These are most parsimoniously explained by positing a knuckle-walking precursor to human bipedalism.

Dental development and life history in Anapithecus hernyaki.

The sample of Anapithecus from Rudabánya, Hungary, is remarkable in preserving a large number of immature individuals. We used perikymata counts, measurements of root length and cuspal enamel thickness, and observations of the sequence of tooth germs that cross match specific developmental stages in Anapithecus to construct the first composite picture and time scale for dental development in a pliopithecoid (Catarrhini, Primates). We conclude that the age of eruption of M1 in Anapithecus was similar to various macaque species (approximately 1.45 months), but that M2 and M3 emergence were close to 2.2 and 3.2 years, respectively (both earlier than expected for similarly sized cercopithecoids). There may have been little difference in individual tooth formation times between cercopithecoids and Anapithecus, but the degree of molar overlap during M1, M2, and M3 crown development, which is extreme in Anapithecus, is fundamentally different. Overall dental development in Anapithecus was very rapid. Old World monkeys appear derived in lacking significant molar overlap, and hominoids may be derived in having longer tooth formation times, both resulting in longer overall dental development times. This is consistent with the general conclusion that the Pliopithecoidea is an outgroup to the Cercopithecoidea and the Hominoidea. On the other hand, rapid dental formation in Anapithecus may be an apomorphy indicative of an unusually rapid life history or unique pressures related to diet and maturation. Folivory and/or predation pressure may be responsible for generating selection to more rapidly erupt permanent teeth and possibly attain adult body masses in Anapithecus. Whatever the case, Anapithecus, with an M3 emergence of approximately 3.2 years, is dramatically faster than any extant catarrhine of similar body mass. This represents yet another unusual attribute of this poorly known fossil catarrhine.

The oldest Eurasian hominoid.

Engelswies is an early Miocene vertebrate locality in southern Germany with a rich assemblage of terrestrial mammals, invertebrates and fossil plants. It is dated to 16.5-17.0 Ma based on magnetostratigraphy, biostratigraphy and lithostratigraphy, and includes among the faunal remains a hominoid upper molar fragment, the oldest hominoid so far identified from Europe. The evidence from Engelswies suggests that hominoids arrived in Eurasia about 17 Ma, roughly contemporaneously with pliopithecoids and Deinotherium, and before the last marine transgression to isolate Eurasia from Africa. Thick enamel and low dentine penetrance may have been key adaptations that contributed to the success of hominoids of dentally modern aspect in western Eurasia and ultimately to their ability to spread to eastern Eurasia and Africa in the middle and late Miocene.

Primates from Rudabánya: allocation of specimens to individuals, sex and age categories.

Fossil primates have been known from the late Miocene locality of Rudabánya since 1965. Numerous campaigns of collecting, sampling and excavation have been carried out since that time by several teams of researchers, but the sample of primates has never been fully catalogued and published. Here we provide a comprehensive list of all primate specimens from Rudabánya with provenience data and allocation to individuals. At the main locality of R II 16 individuals are attributed to Anapithecus and nine to Dryopithecus, based on dental remains. Anapithecus comes mostly from a layer of gray to black marl and Dryopithecus is found mostly in a less consolidated overlying black mud. However, both taxa are found in both layers. Anapithecus is represented by larger proportions of juveniles and females, and Dryopithecus by more adult and subadult males. Both species are represented primarily by dental remains, but those of Dryopithecus are more commonly associated with mandibles and maxillae, while in Anapithecus most individuals are represented by associated dentitions. Dryopithecus is better represented by postcrania other than phalanges. Anapithecus age and sex frequency distributions are more typical of those of carnivore and chimpanzee prey assemblages than Dryopithecus.

Origin of human bipedalism: The knuckle-walking hypothesis revisited.

Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

Restoration of the type and palate of Ankarapithecus meteai: taxonomic and phylogenetic implications.

A small number of large hominoid specimens are known from the late Miocene of Turkey. (This paper focuses on the two specimens known until 1996. New fossils described by Alpagut et al. ([1996] Nature 382:349-351) are briefly discussed in this paper as well.) Among these, a fragmentary mandible is the type specimen of Ankarapithecus meteai. Another specimen, a partial face, is from the same taxon. Based on the morphology of this face, Ankarapithecus meteai was synonymized with Sivapithecus (Andrews and Tekkaya [1980], Paleontology 23:85-95). The facial specimen was recently restored by the authors, exposing much anatomy that had previously been obscured. The new anatomical details reveal many important differences from Sivapithecus and justify a revision of the nomen Ankarapithecus. Ankarapithecus was a large hominid (great apes and humans) in the clade that also includes Sivapithecus and Pongo. Sivapithecus and Pongo share derived characters not found in Ankarapithecus, which is thus the sister clade to the Sivapithecus-Pongo clade. While the results of this analysis support the sister relationship of Sivapithecus and Pongo, there is some uncertainty regarding paleobiogeographic and taxonomic relationships to the large hominids from the Siwaliks. An Ankarapithecus-like taxon may have been ancestral to Sivapithecus, or an early Siwalik (Chinji formation) taxon, which predates both Sivapithecus sensu stricto and Ankarapithecus, may be ancestral to both.

A new reconstruction of RUD 77, a partial cranium of Dryopithecus brancoi from Rudabánya, Hungary.

A newly reconstructed cranium (RUD 77) of the Miocene fossil hominoid Dryopithecus, formerly Rudapithecus (Kretzoi [1969] Symp. Biol. Hung. 9:3-11; Begun and Kordos [1993] J. Hum. Evol. 25:271-286) is presented here. This specimen, from the late Miocene locality of Rudabánya, in northeastern Hungary, consists of portions of the neurocranium, face, and postcanine dentition. Newly recovered portions of the parietal, occipital, temporal, zygomatic, and premaxillary bones, which are described here for the first time, in association with previously described portions of this specimen (Kordos [1987] Ann. Hist. Nat. Mus. Natl. Hung. 79:77-88) make RUD 77 among the most complete and well preserved neurocrania of any Miocene hominoid. Detailed anatomical descriptions and measurements are provided here, along with comparisons to other relatively complete Miocene hominoid cranial remains, and to living hominoids. While a more complete phylogenetic analysis is in preparation based on the sample as a whole, it is suggested here that RUD 77 provides some additional evidence in support of a previous hypothesis that Dryopithecus is more closely related to the African apes and humans than is Sivapithecus.

Miocene fossil hominids and the chimp-human clade.

Miocene hominoids from Europe are among the earliest members of the great ape and human clade (the Hominidae). One of these forms, represented by well-preserved cranial remains from Rudabánya, Hungary, sheds new light on the question of the evolutionary relations among living hominids. This new evidence supports the view that humans have a specific evolutionary relation with chimpanzees, to the exclusion of all other apes.

Phyletic diversity and locomotion in primitive European hominids.

A major contribution of previous analyses of Miocene hominoid postcrania is the recognition of a great ape grade of locomotor morphology in the late Miocene. However, in the absence of a consideration of the taxonomic and phylogenetic implications of the specimens concerned, the importance of this conclusion remains unappreciated. This paper presents a revised view of the positional implications of late Miocene hominid fossils and considers some of the taxonomic and phyletic implications of these specimens. The taxonomic status of a number of large catarrhine specimens from Europe (attributed to Dryopithecus, Sivapithecus, Austriacopithecus, Paidopithex, Rudapithecus) is discussed. The functional and phyletic significance of this material reveals a complex pattern of behavioral and phyletic diversity among large-bodied catarrhines in Europe and suggests that this diversity evolved in situ from circum-Mediterranean middle Miocene ancestors.

A large pliopithecine molar from Germany and some notes on the Pliopithecinae.

A lower molar of a large-bodied pliopithecine from the West German locality of Salmendingen is described and attributed to the taxon Anapithecus cf. hernyaki. This identification is used with other evidence to support the hypothesis that a femur from the nearby locality of Eppelsheim should be attributed to the Pliopithecinae. These newly identified pliopithecine fossils provide evidence of primate diversity and morphological uniqueness in the Late Miocene of Europe.