Parallel evolution in the hominoid trunk and forelimb

The evolutionary history of the living hominoids has remained elusive despite years of exploration and the discovery of numerous Miocene fossil ape species. Part of the difficulty can be attributed to the changing nature of our views about the course of hominoid evolution. In the 1950s and 1960s, individual Miocene taxa were commonly viewed as the direct ancestors of specific living ape species, suggesting an early divergence of the modern lineages.^1–5 However, in most cases, the Miocene forms were essentially “dental apes,” resembling extant species in dental and a few cranial features, but possessing more primitive postcranial features that suggested arboreal quadrupedalism rather than suspensory habits. With the introduction of molecular methods of phylogenetic reconstruction and the increasing use of cladistic analysis, it has become apparent that the radiation leading to the modern hominoids was somewhat more recent than had been believed, and that most of the Miocene hominoid species had little to do with the evolutionary history of the living apes. © 1998 Wiley-Liss, Inc.     

Nucleotide sequences of immunoglobulin-epsilon pseudogenes in man and apes and their phylogenetic relationships

To understand the phylogenetic relationships between hominoids, the nucleotide sequences of immunoglobulin-epsilon processed pseudogenes from chimpanzee, gorilla and orangutan were determined. The basic structures of these processed pseudogenes agreed with their human counterpart. Although the degrees of nucleotide differences between man and the African apes had no statistical significance, all the analytical data examined supported the theory that chimpanzee is the closest relative of man. This result was consistent with that deduced by our recent qualitative study. Studies on the nucleotide sequences of globin genes have suggested that the molecular clock runs more slowly in hominoids than in non-hominoid primates. According to the present data, however, further retardation of the evolutionary rate was not observed in the human lineage. Assuming that orangutan diverged 14 million years ago and that the evolutionary rate between the orangutan lineage and the lineage leading to the other three species is constant, the divergence dates of chimpanzee and gorilla were estimated to be 4.9(+/- 0.9) and 5.9(+/- 0.9) million years ago, respectively.     

Enamel thickness,microstructure and development in Afropithecus turkanensis

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.     

Patterns of covariation in the hominoid craniofacial skeleton: implications for paleoanthropological models

Living species are often used as analogues for fossil ones. When this is done, the implicit assumption is made that hominids and living hominoids vary in the same way. This paper addresses the validity of this assumption by comparing patterns of facial variation among humans and African apes. In particular, it addresses three major questions that underlie approaches to reconstructing hominid relationships. First, is phenotypic variation similar between closely related species? Second, if it is dissimilar, why? Third, is it feasible to use analogue species for modeling purposes? Measurements are obtained from 542 crania of adult apes and humans. Care is taken to choose homologous data, and account for differences in population size and structure. Variance/covariance and correlation matrices among the species are compared using common principal component (CPC) analysis, random skewers methods and matrix correlations. Morphological distances (D(2)) are calculated between population means, and between randomized pairs of individuals within each population, to evaluate intraspecific variation. Morphological distances are also calculated between randomized pairs of individuals using the variation patterns of analogue populations, in order to evaluate the efficacy of such substitutions. Results show that while the hominoids share a similar pattern of facial variation overall, the patterns do diverge. This difference generally corresponds to the phylogenetic relationships among these species, suggesting that patterns of variation may have diverged through time in the large bodied hominoids. Because interpretation of relationships in the fossil record is confounded by a lack of understanding of how variation changes through time, exploration of such patterns of divergence can provide important clues to understanding human evolution. Additionally, neglecting to account for this divergence when using living analogues as variation "yardsticks" can give rise to interpretations of the fossil record that are more speciose than is warranted.     

Apes and Tricksters: The Evolution and Diversification of Humans’ Closest Relatives

The evolutionary history of humans comprises an important but small branch on the larger tree of ape evolution. Today’s hominoids—gibbons, orangutans, gorillas, chimpanzees, and humans—are a meager representation of the ape diversity that characterized the Old World from 23–5 million years ago. In this paper, I briefly review this evolutionary history focusing on features important for understanding modern ape and human origins. As the full complexity of ape evolution is beyond this review, I characterize major geographic, temporal, and phylogenetic groups using a few flagship taxa. Improving our knowledge of hominoid evolution both complicates and clarifies studies of human origins. On one hand, features thought to be unique to the human lineage find parallels in some fossil ape species, reducing their usefulness for identifying fossil humans. On the other hand, the Miocene record of fossil apes provides an important source for generating hypotheses about the ancestral human condition; this is particularly true given the dearth of fossils representing our closest living relatives: chimpanzees and gorillas.     

Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees

To study the genomic divergences among hominoids and to estimate the effective population size of the common ancestor of humans and chimpanzees, we selected 53 autosomal intergenic nonrepetitive DNA segments from the human genome and sequenced them in a human, a chimpanzee, a gorilla, and an orangutan. The average sequence divergence was only 1.24% +/- 0.07% for the human-chimpanzee pair, 1.62% +/- 0.08% for the human-gorilla pair, and 1.63% +/- 0.08% for the chimpanzee-gorilla pair. These estimates, which were confirmed by additional data from GenBank, are substantially lower than previous ones, which included repetitive sequences and might have been based on less-accurate sequence data. The average sequence divergences between orangutans and humans, chimpanzees, and gorillas were 3.08% +/- 0.11%, 3.12% +/- 0.11%, and 3.09% +/- 0.11%, respectively, which also are substantially lower than previous estimates. The sequence divergences in other regions between hominoids were estimated from extensive data in GenBank and the literature, and Alus showed the highest divergence, followed in order by Y-linked noncoding regions, pseudogenes, autosomal intergenic regions, X-linked noncoding regions, synonymous sites, introns, and nonsynonymous sites. The neighbor-joining tree derived from the concatenated sequence of the 53 segments--24,234 bp in length--supports the Homo-Pan clade with a 100% bootstrap value. However, when each segment is analyzed separately, 22 of the 53 segments (approximately 42%) give a tree that is incongruent with the species tree, suggesting a large effective population size (N(e)) of the common ancestor of Homo and Pan. Indeed, a parsimony analysis of the 53 segments and 37 protein-coding genes leads to an estimate of N(e) = 52,000 to 96,000. As this estimate is 5 to 9 times larger than the long-term effective population size of humans (approximately 10,000) estimated from various genetic polymorphism data, the human lineage apparently had experienced a large reduction in effective population size after its separation from the chimpanzee lineage. Our analysis assumes a molecular clock, which is in fact supported by the sequence data used. Taking the orangutan speciation date as 12 to 16 million years ago, we obtain an estimate of 4.6 to 6.2 million years for the Homo-Pan divergence and an estimate of 6.2 to 8.4 million years for the gorilla speciation date, suggesting that the gorilla lineage branched off 1.6 to 2.2 million years earlier than did the human-chimpanzee divergence.     

Major features in the evolution of early hominoid locomotion

Evolution of hominoid locomotion is a traditional topic in primate evolution. Views have changed during the last decade because a number of crucial differences between early and advanced hominoid morphologies have been demonstrated. Increasing evidence on primate behaviour and ecology show that any direct analogies between living and fossil hominoids must be made extremely carefully. The necessity of synthesizing data on primate behaviour, locomotion, morphology and ecology and simultaneously defining the framework in which the data should be interpreted are explained. Results of our studies of ontogeny of locomotor and behavioural patterns (LBP) are presented that could help identify the main features of early hominoid locomotor patterns (LP) and the mechanisms of their changes. The early hominoid LP was different from those of pronograde monkeys and specialized antipronograde living apes. Some similar features could be expected between early hominoid LP and the LP of ceboid monkeys. Analogous mechanisms of change of LBP exist in all groups of living higher primates. Crucial early mechanisms of change are the ontogenetic shifts in LBP connected with ethoecological changes. Analysis of fossil evidence has shown that Miocene hominoids differ morphologically from any group of living primates. Certain features present in Miocene hominoids could be found in Atelinae and living Asian apes but they are limited to some functional regions of the postcrania only. Consequently the early hominoid general LP can not be strictly analogous either to that of any monkey group or to the LP of apes. We suppose that certain pronograde adaptations, such as climbing, bipedality, limited suspensory activity and sitting constituted the main part of their LP.     

Evolution of the hominoid vertebral column: The long and the short of it

The postcranial axial skeleton exhibits considerable morphological and functional diversity among living primates. Particularly striking are the derived features in hominoids that distinguish them from most other primates and mammals. In contrast to the primitive catarrhine morphotype, which presumably possessed an external (protruding) tail and emphasized more pronograde trunk posture, all living hominoids are characterized by the absence of an external tail and adaptations to orthograde trunk posture. Moreover, modern humans evolved unique vertebral features that satisfy the demands of balancing an upright torso over the hind limbs during habitual terrestrial bipedalism. Our ability to identify the evolutionary timing and understand the functional and phylogenetic significance of these fundamental changes in postcranial axial skeletal anatomy in the hominoid fossil record is key to reconstructing ancestral hominoid patterns and retracing the evolutionary pathways that led to living apes and modern humans. Here, we provide an overview of what is known about evolution of the hominoid vertebral column, focusing on the currently available anatomical evidence of three major transitions: tail loss and adaptations to orthograde posture and bipedal locomotion.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

Hominid and gelada baboon evolution: Agreement between molecular and fossil time scales

The time of origin of the hominid lineage has long been debated. Macromolecular studies have consistently shown genetic distances between living humans and African apes to be quite small. The molecular clock hypothesis proposes that the time of separation of these lineages is relatively recent (in the range of 4–8 million years ago) and not 15 million years or more ago as usually suggested. Three independent molecular comparisons yield a mean estimate of 4.6 million years for the hominid-African pongid divergence. The relationship of Theropithecusand Papiois a parallel case within Primates of two taxa which are quite similar at the molecular level, but which are usually thought to have separated relatively long ago. The two cases of seeming discordance between different lines of evidence are analogous. Each involves a speciation event which eventually resulted in one substantially derived lineage and one or more relatively unchanged lineages. In each case, claims of the antiquity of the divergence event extend to at least twice the age of the first certain appearance of the more derived lineage in the fossil record. Finally, in each case, the molecular clock model suggests a range of possible divergence times that overlaps with the first appearances of undoubted hominids and Theropithecusin the fossil record. This test involving paleontological evidence supports the molecular clock hypothesis.     

Molar crown formation in the Late Miocene Asian hominoids, Sivapithecus parvada and Sivapithecus indicus

During the past decade, studies of enamel development have provided a broad temporal and geographic perspective on evolutionary developmental biology in Miocene hominoids. Here we report some of the first data for molar crown development in one hominoid genus, Sivapithecus. The data are compared to a range of extant and extinct hominoids. Crown formation times (CFTs), daily rates of enamel secretion (DSR), Retzius line number and periodicity, and relative enamel thickness (RET) were calculated in a mandibular first molar of Sivapithecus parvada and a maxillary first molar of Sivapithecus indicus from the Siwalik sequence of Pakistan. A CFT of 2.40 years for the protoconid of S. parvada and 2.25 years for the protocone of S. indicus lie within the range of first molar (M1) formation times for the majority of Miocene hominoids (1.96-2.40 years, excluding Proconsul heseloni), and are similar to an M(1) from Gorilla (2.31 years) and M(1)s from Pan (2.22-2.39 years). This is unlike the longer CFTs in modern humans, which appear to be linked with their extended growth period. In contrast to extant great apes and humans, daily rates of enamel secretion are rapid in the Sivapithecus M1s during the early stages of growth, which seems to be a common pattern for most Miocene apes. The rapid accumulation of cuspal enamel in the Sivapithecus molars produced thicker enamel than either Pan or Gorilla in a comparable period of time. Future studies on larger samples of living and fossil hominoids are needed to clarify trends in crown development, which may be better understood in the context of life history strategies coupled with good data on body mass and brain size.     

Divergence time estimates for the early history of animal phyla and the origin of plants, animals and fungi

In the past, molecular clocks have been used to estimate divergence times among animal phyla, but those time estimates have varied widely (1200-670 million years ago, Ma). In order to obtain time estimates that are more robust, we have analysed a larger number of genes for divergences among three well-represented animal phyla, and among plants, animals and fungi. The time estimate for the chordate-arthropod divergence, using 50 genes, is 993 +/- 46 Ma. Nematodes were found to have diverged from the lineage leading to arthropods and chordates at 1177 +/- 79 Ma. Phylogenetic analyses also show that a basal position of nematodes has strong support (p > 99%) and is not the result of rate biases. The three-way split (relationships unresolved) of plants, animals and fungi was estimated at 1576 +/- 88 Ma. By inference, the basal animal phyla (Porifera, Cnidaria, Ctenophora) diverged between about 1200-1500 Ma. This suggests that at least six animal phyla originated deep in the Precambrian, more than 400 million years earlier than their first appearance in the fossil record.     

Genetic aspects in hominid evolution

Genomic comparison between apes and humans have made important contributions to our understanding of human evolution. The modern period of karyological comparisons between humans and other primates began about forty years ago and has been marked by a series of technical revolutions. In the 1960s pioneering genetic and chromosomal comparisons of human and great apes suggested, as had Darwin a century before, that our closest relative were the African apes. Early immunological analyses placed human/apes divergence at about five million year ago. Acceptance of man’s late divergence from the African apes was delayed by the scarcity of paleontological evidence coupled with a fallacious Asiatic origin hypothesis of the hominoids. Chromosome banding techniques in the seventies and high resolution methods in the eighties allowed a detailed comparison of the chromosomes between closely related primates and reinforced the hypothesis of an African origin for humans. It was clearly shown that humans were more closely related to African apes than to the orang-utan. The last decade has seen a vigorous integration of molecular and cytogenetic. This powerful combination promises to be quite fruitful because chromosomes can be compared directly at the DNA level. Fluorescentin situ hybridisation (FISH), chromosome painting, is a colourful technique for establishing chromosomal homology between species. Results obtained by FISH over the last ten years have resolved the cytogenetic problem of the homology between humans, apes, hylobates and Old World monkeys and defined the chromosomal syntenies and major translocations involved in the genome evolution of higher primates.     

Reexamination of the African hominoid trichotomy with additional sequences from the primate beta-globin gene cluster.

Additional DNA sequence information from a range of primates, including 13.7 kb from pygmy chimpanzee (Pan paniscus), was added to data sets of beta-globin gene cluster sequence alignments that span the gamma 1, gamma 2, and psi eta loci and their flanking and intergenic regions. This enlarged body of data was used to address the issue of whether the ancestral separations of gorilla, chimpanzee, and human lineages resulted from only one trichotomous branching or from two dichotomous branching events. The degree of divergence, corrected for superimposed substitutions, seen in the beta-globin gene cluster between human alleles is about a third to a half that observed between two species of chimpanzee and about a fourth that between human and chimpanzee. The divergence either between chimpanzee and gorilla or between human and gorilla is slightly greater than that between human and chimpanzee, suggesting that the ancestral separations resulted from two closely spaced dichotomous branchings. Maximum parsimony analysis further strengthened the evidence that humans and chimpanzees share the longest common ancestry. Support for this human-chimpanzee clade is statistically significant at P = 0.002 over a human-gorilla clade or a chimpanzee-gorilla clade. An analysis of expected and observed homoplasy revealed that the number of sequence changes uniquely shared by human and chimpanzee lineages is too large to be attributed to homoplasy. Molecular clock calculations that accommodated lineage variations in rates of molecular evolution yielded hominoid branching times that ranged from 17-19 million years ago (MYA) for the separation of gibbon from the other hominoids to 5-7 MYA for the separation of chimpanzees from humans. Based on the relatively late dates and mounting corroborative evidence from unlinked nuclear genes and mitochondrial DNA for the close sister grouping of humans and chimpanzees, a cladistic classification would place all apes and humans in the same family. Within this family, gibbons would be placed in one subfamily and all other extant hominoids in another subfamily. The later subfamily would be divided into a tribe for orangutans and another tribe for gorillas, chimpanzees, and humans. Finally, gorillas would be placed in one subtribe with chimpanzees and humans in another, although this last division is not as strongly supported as the other divisions.     

Cautious climbing and folivory: a model of hominoid differentation

Despite the large and growing number of Miocene fossil catarrhine taxa, suitable common ancestors of great apes and humans have yet to be agreed upon. Considering a) the conservative and primitive nature of the hominoid molar cusp pattern, and b) the variability of secondary dental features, it is difficult to discern whether a hominoid dentition is primitive, secondarily simplified to the primitive condition or too far derived to be ancestral to any of the living forms. Nonetheless, the inability to recognize a common ancestor is primarly due to the absence of a model of hominoid differentiation that provides a basis for its recognition. Vertical climbing as the limiting component of cautious climbing, explains all of the locomotor anatomy shared by living hominoids. Comparison of the shared derived characters of hominoids to those of forms which have converged on hominoidsi.e colobines, atelines, lorisines, paleopropithecines and sloths suggest that early hominoids were probably folivores. In arboreal forms there is a strong link between a large body size, folivory and cautious climbing. Comparison of craniodental characters of committed folivores to committed frugivores from among each of the compared groups with the exception of lorisines, indicates that many of the distinguishing craniodental characters of humans and great apes are adaptations to folivory. Many of these characters, however, are also present in Jolly's seed eating complex. As such folivory may be the heritage factor which Jolly hypothesized to account for differential reduction of canines in fossilTheropithecus and hominids.     

Anatomy of the hominoid wrist joint: Its evolutionary and functional implications

Observations on the behavior of living hominoids show generic differences in the use and posture of the wrist joint. Both orang-utans and hylobatids usually use the wrist in suspensory behaviors. However, orang-utans emphasize markedly adducted and flexed wrist postures, while hylobatids emphasize violent forearm and wrist rotation. African apes, especially the gorilla, use the wrist more frequently than other hominoids for terrestrial quadrupedal weight-bearing. Humans use the wrist less frequently for supportive purposes than do other hominoids. These behavioral differences correspond to structural specializations in the proximal carpal joint of each of the hominoid genera. Although each of the hominoid genera has apparently modified its proximal carpal joint best to serve its characteristic behaviors, all hominoids share a unique proximal carpal joint that permits approximately 160ℴ of forearm rotation. The hylobatid proximal carpal joint is specialized in exhibiting a marked development of those structures limiting forearm rotation, but it is in most respects the least derived— that is, closest to the nonhominoid anthropoids. Chimpanzees show a proximal carpal joint that is more generalized than those of the other great apes but more derived than that of hylobatids. The human and gorilla proximal wrist joints, on the other hand, show marked modifications for weight-bearing in terrestrial behaviors. Orang-utans have the most derived proximal carpal joint, which in many respects parallels that of the slow-climbing nonhominoid primates. The comparative anatomy and structural specializations of the wrist joint support (a) an early divergence of hylobatids from the common hominoid stock, (b) a common ancestry for gorillas and humans separate from the other hominoids, and (c) a long independent evolutionary period for orang-utans since their divergence from the common hominoid stock, or one that was marked by strong selection pressures for wrist specializations. Unfortunately, the generalized condition of the chimpanzee’s wrist joint and the very derived condition of the orang-utan wrist provide uncertain evidence as to which of the two was first to diverge from the common hominoid stock. Identification of hominoid wrist specializations as reflecting real phylogenetic relationships or parallelisms depends on how well the phytogeny inferred from wrist morphology accords with those arrived at from the study of other systems.     

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.     

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.     

Age at first molar emergence in Lufengpithecus lufengensis and its implications for life-history evolution

The late Miocene hominoid Lufengpithecus from Yunnan Province, China, is crucial for understanding hominoid evolution in Asia. Given that age at first permanent molar emergence is a key life-history trait in primates, the present study determined the age at death of the Lufengpithecus lufengensis juvenile PA868, which was in the process of erupting its first molar. Using a perikymata periodicity of 7-11 days, along with estimation of cusp formation time and the postnatal delay of crown mineralization, perikymata counts obtained from the permanent central incisor and canine germs indicate that the age at death of PA868 was 2.4-4.5 years based on the central incisor germ, and 2.5-4.7 years based on the canine germ. The age at the first molar emergence was actually slightly younger (by about 0.3 years), as demonstrated by tiny wear facets on this tooth, which indicate that gingival emergence had occurred sometime before death. The average age at first molar emergence of Lufengpithecus lufengensis PA868 is estimated to be 3.2-3.3 years, with a range of 2.1-4.4 years. In comparison to extant primates and other fossil hominoids, the life history of Lufengpithecus lufengensis is similar to that of extant great apes and the Miocene hominoids Afropithecus turkanensis and Sivapithecus parvada, as well as Plio-Pleistocene Australopithecus, and different from monkeys, gibbons, and modern humans.     

A New Malaria Agent in African Hominids

Plasmodium falciparum is the major human malaria agent responsible for 200 to 300 million infections and one to three million deaths annually, mainly among African infants. The origin and evolution of this pathogen within the human lineage is still unresolved. A single species, P. reichenowi, which infects chimpanzees, is known to be a close sister lineage of P. falciparum. Here we report the discovery of a new Plasmodium species infecting Hominids. This new species has been isolated in two chimpanzees (Pan troglodytes) kept as pets by villagers in Gabon (Africa). Analysis of its complete mitochondrial genome (5529 nucleotides including Cyt b, Cox I and Cox III genes) reveals an older divergence of this lineage from the clade that includes P. falciparum and P. reichenowi (∼21±9 Myrs ago using Bayesian methods and considering that the divergence between P. falciparum and P. reichenowi occurred 4 to 7 million years ago as generally considered in the literature). This time frame would be congruent with the radiation of hominoids, suggesting that this Plasmodium lineage might have been present in early hominoids and that they may both have experienced a simultaneous diversification. Investigation of the nuclear genome of this new species will further the understanding of the genetic adaptations of P. falciparum to humans. The risk of transfer and emergence of this new species in humans must be now seriously considered given that it was found in two chimpanzees living in contact with humans and its close relatedness to the most virulent agent of malaria.