Notes on Ramapithecus, the earliest known hominid, and Dryopithecus

Recent paleontological analysis of the Higher Primate subfamily Dryopithecinae shows that fossils in this group can be referred to two genera, Ramapithecus and Dryopithecus. Ramapithecus is known from India and East Africa in Late Miocene or Early Pliocene time (about 14 m. years ago). The remains of Ramapithecus resemble closely the equivalent parts of the later Hominidae and contrast with those of the Pongidae. It is concluded that Ramapithecus is the earliest known hominid, some 5 or 6 times older than the oldest Pleistocene hominids. Dryopithecus is a pongid and contains as subgenera (Dryopithecus), (Proconsul), and (Sivapithecus). Probably part of (Proconsul) is ancestral to the chimpanzee and part to the gorilla, while part of (Sivapithecus) is ancestral to the orang-utan.     

Problems in the interpretation of Ramapithecus: with special reference to anterior tooth reduction

The dental proportions of Ramapithecus specimen FT 1271-2 (from Fort Ternan, Kenya) have been compared with undoubted fossil pongids from the Miocene of East Africa. Compared to its Miocene pongid contemporaries, Ramapithecus exhibits distinct anterior tooth reduction both in its incisor and canine dimensions. This distinction is most clearly seen in comparisons of Ramapithecus with pongids of similar cheek tooth size, i.e., Dryopithecus africanus and Pan paniscus. The differences in dental proportions between the phylogenetic lines of D. africanus to Pan and Ramapithecus to Homo are discussed in terms of various dietary hypotheses. The similarities in dental proportions of Gorilla and Ramapithecus illustrate their non-frugivorous dietary preferences, but have little or no value as far as the taxonomic assessment of Ramapithecus is concerned.     

Hominoids, hominids and the problem of the lower boundary of the Anthropogene

This paper distinguishes two phases within the Anthropogene, each being characterized by specific features of hominization: the earlier phase the Eoanthropogene, is characterized by the presence of bipedal, small-brained hominoids. These include the australopithecine and habiline fossils, all of which are classified in a separate family, the Australopithecidae. In the second phase, the Euanthropogene, the family Hominidae is present, with its two species, Homo erectus and H. sapiens. The two families, Australopithecidae and Hominidae, are united into a superfamily, the Hominoidea; whilst the pongids are placed in a separate family (Pongidae) and superfamily (Pongoidea). The bearing of the work of the Soviet palaeo-neurologist, the late V. I. Kochetkova on the status of the australopithecines, is reviewed in some detail.     

The nature of the transition in the dental mechanism from pongids to hominids

Many recent discoveries of Ramapithecus, and of probably ground-living dryopithecines, Dryopithecus (subgenus Sivapithecus), clarify the nature of the transition of the dental mechanism from that of pongids to the hominid stage with reduced canines and flattened cheek teeth with thick enamel.Faunal correlation with potassium/argon dated sites indicates that Sivapithecus and Ramapithecus appeared in the Old World about the same time, approximately 13 million years ago. The thickened molar enamel of these hominoids suggests a terrestrial adaptation in both groups, probably resulting from climatic changes. This adaptation was not necessarily a unique event in the ancestry of the two genera, for species of the two seem to have been different sizes when the change was made.New Ramapithecus finds come from Pyrgos, near Athens, from Çandir in Anatolia, and from Rudabánya, Hungary. At the latter site various specimens preserve all upper and lower teeth in place, while the Çandir and Pyrgos mandibles give important new information about symphyseal structure and orientation, as well as about arcade arrangement. The Rudabánya finds confirm, as do the others, marked facial foreshortening, relatively orthal incisors, anteriorly abbreviated mandible and canine reduction in Ramapithecus. The dental mechanics of Ramapithecus suggested from earlier described finds recovered in the Siwalik deposits of India and Pakistan, as well as at Fort Ternan, Kenya are clarified by the finds from Athens, Anatolia, and Hungary. Like Australopithecus, Ramapithecus mandibles have well-developed double transverse, shallow but transversely thick horizontal rami and anteriorly shifted, vertically oriented, deep ascending branches. These addes resemblances increase the probability that Ramapithecus is in or near the ancestry of Australopithecus and other hominids.     

“Australopithecus afarensis”: A composite species

In response to a critique byFerguson (1989),Leonard (1991) reiterates most of his original arguments for supporting “Australopithecus afarensis”Johanson, White, andCoppens, 1978 as a single species. He disregards the principle of morphological equivalence by comparing the dental metrics and morphology of a hominid with those of species of the Pongidae, which do not correspond with the degree of variation in hominids, instead of with those of species of the Hominidae. He fails to refute clear evidence that the range of variation of dental metrics and morphology in “A. afarensis” exceeds that seen in species of the Hominidae. On the basis of extreme variation, “A. afarensis” is, therefore, interpreted as representing a composite species.     

Molekulare und., adaptive Evolution, Kladistik und Stammesgeschichte1: Ergänzungen zu einer Arbeitshypothese

Molecular and “adaptive” evolution, cladistic and phylogeny. Supplementations to a working hypothesis The methods of molecular biology and morpholo for the taxonomical assessment have given in several cases very different results. The following examples are discussed: Ailuropoda melanoleuca as an ursoid member of the Carnivora (Ursidae or Ailuropodidae?), Homo saiens and his position to the African apes (Pan and Gorilla) (Pongidae or Hominidae?), Theroipithecus gelada and its affinities to the baboons (Papionini or Theropithecini ?) and Oreopithecus bambolii from the late Miocene of Italy and its taxonomical position within the catarrhine primates. For the three extant species not only the “two level evolution” (molecular and morphological-adaptative evolution) is evident, but also the working hypothesis, that the “adaptative” evolution is more rapidly as the molecular evolution, as the author is suggesting since 1969. The results of molecular methods (immunoloy, amino acid sequences, electrophoresis, DNA-DNA hybridization) are more important for the phylogeny than for the taxonomy. For the cladistic method it is clear, that a cladogram is not a phylogenetic tree, as suggested Peter Ax (1984).     

Karyotypic fission theory and the evolution of old world monkeys and apes

The karyotypes of living catarrhines are correlated with the current concepts of their fossil record and systematic classification. A phylogeny, beginning at the base of the Oligocene, for those animals and their chromosome numbers is presented. Todd? (1970) theory of karyotypic fissioning is applied to this case — three fissioning events are hypothesized. A late Eocene event (the primary catarrhine fissioning) is hypothesized to underlie the diversification of the infraorder Catarrhini into its extant families, the second fissioning underlies the radiation of the Pongidae/Hominidae in the Miocene and the third accounts for the high chromosome numbers (54–72) and the Neogene (Miocene-Pliocene-Pleistocene) radiation of members of the genus Cercopithecus. Published catarrhine chromosome data, including that for “marked” chromosomes (those with a large achromatic region that is the site for ribosomal RNA genes) are tabulated and analysed. The ancestral X chromosome is always retained in the unfissioned metacentric state. The Pongidae/Hominidae have 15 pairs of mediocentric chromosomes that survived the second fissioning whereas the other chromosomes (besides the X) are thought to be fission-derived acrocentrics. Both the detailed karyology and the trend from low to high numbers is best interpreted to support Todd? concept of adaptive radiations correlated with karyotypic fissioning in ancestral populations.     

Indopithecus giganteus distinct fromSivapithecus indicus

The very large Eurasian Miocene apeIndopithecus giganteus is distinct from contemporaniousSivapithecus (non-Dryopithecus)indicus. The probabilities that length and width for the only specimen ofI. giganteus could be sampled from populations similar or identical to those ofS. indicus are less than six chances in 100,000 for both parameters.     

Evidence for an HLA-C-like locus in the orangutan Pongo pygmaeus

 HLA-B and C are related class I genes which are believed to have arisen by duplication of a common ancestor. Previous study showed the presence of orthologues for both HLA-B and C in African apes but only for HLA-B in Asian apes. These observations suggested that the primate C locus evolved subsequent to the divergence of the Pongidae and Hominidae. From an analysis of orangutan Tengku two HLA-C-like alleles (Popy C*0101 and Popy C*0201) were defined as well as three HLA-B-like (Popy-B) alleles. By contrast, no Popy-C alleles were obtained from orangutan Hati, although three Popy-B alleles were defined. Thus an HLA-C-like locus exists in the orangutan (as well as a duplicated B locus), implying that the primate C locus evolved prior to the divergence of the Pongidae and Hominidae and is at least 12–13 million years old. Uncertain is whether all orangutan MHC haplotypes contain a C locus, as the failure to find C alleles in some individuals could be due to a mispairing of HLA-C-specific primers with certain Popy-C alleles. These results raise the possibilities that other primate species have a C locus and that the regulation of natural killer cells by C allotypes evolved earlier in primate evolution than has been thought. Received: 18 January 1999 / Revised: 23 March 1999     

Sexual dimorphisms in dental dimensions of higher primates

Dental dimensions and distributions of dental dimensions of males and females were compared for great apes (Pan, Gorilla, and Pongo, and humans (Homo). The results were examined and discussed with reference to fossil primates Sivapithecus and Ramapithecus. The analyses focused on patterns of sexual dimorphism, both with regard to mean dimensions and the distribution of those dimensions. Sex differences in mean canine dimensions were large and significant for Gorilla and Pongo, significant but smaller for Pan, and small but occasionally significant for Homo. The dispersions of measures were greater for males than for females in Gorilla and Pan but did not differ significantly for Pongo or Homo. Examination of the noncanine teeth revealed complex sex differences. In the anterior teeth, sex differences in mean dimensions were generally apparent for Gorilla and Pongo, less so for Pan, and least of all in Homo. The patterns of dispersion of measures of anterior teeth differed markedly from those of the canines. Pan exhibited the same pattern for anterior and canine teeth. Gorilla showed the opposite pattern. Pongo and Homo showed similar dispersions for males and females in many cases. Sex differences in posterior teeth followed the pattern of the canines for Gorilla and were absent for Pan. Pongo exhibited mean differences in dimensions across sex, but dispersions were similar. The pattern for Homo was most like that of Pongo, but with fewer significant differences. The genera differed with regard to the number of significant differences in means or dispersions along the tooth row. It is clear that the patterns of dimorphism differ qualitatively across all extant genera of great apes and humans. It appears that the pattern for Homo most closely resembles that of Ramapithecus, whereas Pongo most closely resembles Sivapithecus. The patterns for Gorilla and Pan appear to be unlike either of the fossil forms. It is suggested that the qualitatively distinct patterns of dental sexual dimorphism indicate substantial flexibility during recent primate evolution and that the degree of structural flexibility demonstrated provides a basis for appreciating potential for plasticity of gender differences in behavioral, social, and cultural systems.     

Palatine fenestrae,the orangutan and hominoid evolution

Although most mammals develop relatively large double anterior palatine fenestrae that patently communicate with the nasal cavity, four extant primates—Homo sapiens, Pongo, Pan andGorilla—do not. While these four have closed-down these foramenal structures,Homo sapiens andPongo are unique in forming a single foramen palatally. Among fossil taxa,Homo, Australopithecus, Sivapithecus (=Ramapithecus) andRudapithecus also develop a single foramen palatally. Dryopithecines, the presumed fossil apes, preserve the two patent fenestrae. In light of dental features that are considered diagnostically “hominid,” which are also found in the orangutan, it is suggested that this “ape,” rather thanPan, is phylogenetically closer toHomo.     

Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals

Sivapithecus is a Miocene great ape from South Asia that is orangutan-like cranially but is distinctive postcranially. Work by others shows that the humerus resembles large terrestrial cercopithecoids proximally and suspensory hominoids distally, but most functional interpretations nevertheless situate Sivapithecus in an arboreal setting. We present a new quantitative analysis of the Sivapithecus capitate and hamate. Though the functional morphology of both bones suggests some degree of arboreality, the overall morphology is most similar to knuckle-walking African apes. Other features of the Sivapithecus humerus and hind limb are also functionally consistent with knuckle-walking, and we suggest that this locomotor behavior is a valid alternative functional interpretation of the postcranial morphology. We speculate that knuckle-walking in Sivapithecus would have evolved independently from African apes, perhaps for similar ecological reasons. The discovery of a possible pongine knuckle-walker challenges the hypotheses that (1) knuckle-walking evolved only once in hominoids and (2) knuckle-walking is too highly specialized to be the positional behavior from which human bipedalism evolved. The possibility of knuckle-walking in Sivapithecus may help to explain not only the curious combination of characters that typify the postcranium but also the unique postcranial morphology of extant Pongo. Furthermore, it may clarify the distribution of fossil pongines across many ecological zones in Eurasia in the Miocene and Pleistocene, as well as, independently, the spread of African apes across a diversity of environments in equatorial Africa.     

The environmental background of hominid emergence and the appearance of the genusHomo

The human biologist usually considers ecology of recent humanity. This essay explores the question of whether the human biologist specialising in the ecology of living peoples has anything to learn from the palaeo-anthropologist, studying ancient hominids andtheir adaptive mechanisms over a deep time dimension. Since the Hominidae are under discussion, the definition of the hominids is reviewed. Historically, three phases are recognised. A rethinking of the classification of the hominoids has become necessary for the old and classical systematics, which divided this superfamily into the Hominidae and the Pongidae, is now outmoded. Since no consensus on such a re-classification has yet been reached, the author adheres to the classical system for the time being. The Hominidae emerged between about 8 and 5 million years ago. At that time, Africa was subject to major cooling and aridification and considerable changes in the flora and fauna were occurring. Wet forests were retreating, savanna was spreading and the animals of Africa were undergoing many changes, partly by faunal interchange with Asia following the drying up of the Mediterranean, and partly by autochthonous evolution among the pre-existing species of the continent. The Hominidae could well have emerged from the striking environmental modifications of this late Miocene phase. Critical changes occurred in hominid evolution between 3 and 2 million years before the present. The pre-existing speciesAustralopithecus africanus acquired the form of a postulated derivedA. africanus; the hominid lineage underwent cladogenetic splitting into robust and hyper-robust australopithecines and the genusHomo; Homo babilis appeared; stone tools are first found in the archaeological record; spoken language seems to have been acquired. These sensational events, within the space of one million years, took place against the background of conspicuous changes in the climate and physical geography of Africa, the flora and non-hominid fauna. Mankind became increasingly dependent upon stone culture. Hence a new element was added to the range of modes of adjustment, an element which must have greatly increased the ecological flexibility of the hominids. From the end of the Pliocene era onwards, culture should be seen as a constituent of man's environment and, at the same time, a highly advantageous component of human adaptational processes. In later and recent mankind, it may be difficult to extricate the respective roles of biological, social and physical factors, on the one hand, and cultural aspects on the other, as mechanisms and facilitators of adaptation to diverse econiches.     

Isotopic reconstructions of habitat change surrounding the extinction of Sivapithecus,a Miocene hominoid,in the Siwalik Group of Pakistan

This research presents an isotopic study of a wide range of mammalian taxa from the Miocene Siwalik Group of Pakistan, with a focus on two time intervals—9.3–9.2 Ma, when the hominoid Sivapithecus was present, and 8.1–8.0 Ma, shortly after Sivapithecus became extinct. The purpose of this investigation is to reconstruct the vegetation mosaic at both time levels in order to determine what Sivapithecus habitat was like and what changes in habitat and climate may have taken place by the time Sivapithecus went extinct. Both carbon and oxygen stable isotopes were sampled from inorganic carbonate in tooth enamel. Results indicate a vegetation mosaic of both closed and open habitat at both time intervals, but a decrease in forest accompanied by an increase in open habitat (including C₄ grasses) during the younger interval. Individuals from many species cluster with each other with respect to both carbon and oxygen isotope values, indicating a spectrum of feeding adaptations exploiting different parts of the habitat. Isotopic values for Sivapithecus suggest that it fed in the forest upper canopy. Taxa feeding in the most closed, wet habitat become extinct by 8.1 Ma. Furthermore, higher δ¹⁸O values over time suggest a change in climate with a reduction in annual rainfall, perhaps accompanied by changes in precipitation sources or rainfall regime as well. These results suggest that forests became fragmented over time. While still present in the younger level, much of the forest was replaced by open habitat, including patches of C₄ grass. Forest loss and fragmentation is a likely cause of the extinction of Sivapithecus.     

A late divergence hypothesis

The possibility of a Middle-Late Miocene separation of the human lineage from the lineages leading to the extant great apes, based on paleontological and phenetic evidence, is presented. Middle Miocene Sivapithecus, rather then Early Miocene Dryopithecus, is supported as a last common ancestor of Pongo, Pan, Gorilla, and Homo. Estimates for the branching of the lineages are a maximum of 15 m.y.a. for the Pongo lineage and a range from 14-6 m.y.a. for the Pan, Gorilla, and Ausralopithecus/Homo lineages. Weaknesses of the late divergence hypothesis are discussed.     

Evolution of the second orangutan: phylogeny and biogeography of hominid origins

Aim To resolve the phylogeny of humans and their fossil relatives (collectively, hominids), orangutans (Pongo) and various Miocene great apes and to present a biogeographical model for their differentiation in space and time. Location Africa, northern Mediterranean, Asia. Methods Maximum parsimony analysis was used to assess phylogenetic relationships among living large‐bodied hominoids (= humans, chimpanzees, bonobos, gorillas, orangutans), and various related African, Asian and European ape fossils. Biogeographical characteristics were analysed for vicariant replacement, main massings and nodes. A geomorphological correlation was identified for a clade we refer to as the ‘dental hominoids’, and this correlation was used to reconstruct their historical geography. Results Our analyses support the following hypotheses: (1) the living large‐bodied hominoids represent a monophyletic group comprising two sister clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas (collectively, the African apes); and (2) the human–orangutan clade (dental hominoids) includes fossil hominids (Homo, australopiths, Orrorin) and the Miocene‐age apes Hispanopithecus, Ouranopithecus, Ankarapithecus, Sivapithecus, Lufengpithecus, Khoratpithecus and Gigantopithecus (also Plio‐Pleistocene of eastern Asia). We also demonstrate that the distributions of living and fossil genera are largely vicariant, with nodes of geographical overlap or proximity between Gigantopithecus and Sivapithecus in Central Asia, and between Pongo, Gigantopithecus, Lufengpithecus and Khoratpithecus in East Asia. The main massing is represented by five genera and eight species in East Asia. The dental hominoid track is spatially correlated with the East African Rift System (EARS) and the Tethys Orogenic Collage (TOC). Main conclusions Humans and orangutans share a common ancestor that excludes the extant African apes. Molecular analyses are compromised by phenetic procedures such as alignment and are probably based on primitive retentions. We infer that the human–orangutan common ancestor had established a widespread distribution by at least 13 Ma. Vicariant differentiation resulted in the ancestors of hominids in East Africa and various primarily Miocene apes distributed between Spain and Southeast Asia (and possibly also parts of East Africa). The geographical disjunction between early hominids and Asian Pongo is attributed to local extinctions between Europe and Central Asia. The EARS and TOC correlations suggest that these geomorphological features mediated establishment of the ancestral range.     

First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès‐Penedès Basin,Catalonia, NE Spain): Taxonomic and phylogenetic implications

A well‐preserved 11.8‐million‐years‐old lower face attributed to the seminal taxon Dryopithecus fontani (Primates, Hominidae) from the Catalan site ACM/C3‐Ae of the Hostalets de Pierola area (Vallès‐Penedès Basin, Catalonia, NE Spain) is described. The new data indicate that D. fontani is distinct at the genus level from Late Miocene European taxa previously attributed to Dryopithecus, which are here reassigned to Hispanopithecus. The new facial specimen also suggests that D. fontani and the Middle Miocene Pierolapithecus catalaunicus are not synonymous. Anatomical and morphometric analyses further indicate that the new specimen shows a combination of lower facial features—hitherto unknown in Miocene hominoids—that resembles the facial pattern of Gorilla, thus providing the first nondental evidence of gorilla‐like lower facial morphology in the fossil record. Considering the current evidence, the gorilla‐like facial pattern of D. fontani is inferred to be derived relative to previously known stem hominids, and might indicate that this taxon is either an early member of the Homininae or, alternatively, a stem hominid convergent with the lower facial pattern of Gorilla. The biogeographic implications of both alternatives are discussed. This new finding in the Hostalets de Pierola section reinforces the importance of this area for understanding the elusive question of the Middle Miocene origin and early radiation of great apes. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Hominoid dental variability and species number at the late Miocene site of Lufeng,China

The large hominoid sample from the late Miocene site of Lufeng, China, has been variously claimed to contain either one or two species, but very few metric data in support of either position have been published. We calculate coefficients of variation for the dental remains both for the two presumed species and for the pooled sample as a whole using the summary statistics published by Wu & Oxnard (Wu & Oxnard: American Journal of Primatology 5:303–344, 1983a, Nature 306:258–260, 1983b). These are compared to the same measures of single-sex and combined-sex samples of extant hominoids. We also present metric characterizations of male and female canines of extant great apes, with which we evaluate the gender composition of the Lufeng canine sample. In a two-species alternative, the two presumed species have measures of variability and canine representation that are more compatible with single-sex samples representing males and females, respectively. The pooled dental sample has measures of variability within the ranges of single species of extant great apes. We conclude there is a single large hominoid species represented at Lufeng that is highly sexually dimorphic. The phylogenetic relationships of this species are briefly considered. It is generally primitive in craniodental morhpology and is unlikely to be a member of the Sivapithecus-Pongo clade.     

Morphometric analysis of hominoid lower molars from Yuanmou of Yunnan Province, China

Shape analyses of cross-sectional mandibular molar morphology, using Euclidean Distance Matrix Analysis, were performed on 79 late Miocene hominoid lower molars from Yuanmou of Yunnan Province, China. These molars were compared to samples of chimpanzee, gorilla, orangutan,Lufengpithecus lufengensis, Sivapithecus, Australopithecus afarensis, and human mandibular molars. Our results indicate that the cross-sectional shape of Yuanmou hominoid lower molars is more similar to the great apes that to humans. There are few differences between the Yuanmou,L. lufengensis, andSivapithecus molars in cross-sectional morphology, demonstrating strong affinities between these three late Miocene hominoids. All three of the fossil samples show strong similarities to orangutans. From this, we conclude that these late Miocene hominoids are more closely related to orangutants than to either the African great apes or humans.     

Secnidazole vs. paromomycin: Comparative antiprotozoan treatment in captive primates

The antiprotozoan activity of secnidazole was studied in Cercocebus t. torquatus, Cercopithecus campbelli, Erythrocebus patas (Cercopithecidae), and Gorilla gorilla (Pongidae) compared with that of paromomycin in Cercocebus t. lunulatus (Cercopithecidae), E. patas, and G. gorilla (Pongidae) by coprological analysis. The antiprotozoan activity of both drugs depended on the parasite species and the host species. The drugs acted in a similar way on Entamoeba coli parasitising C. t. torquatus, and E. patas. This activity was different from that observed on I. buestchlii from the same host species. Nevertheless, E. coli parasitising cercopithecids and pongids responded to drugs differently.