Hominoid Primates Research Articles

Dental metric comparisons of Morotopithecus and Afropithecus: Implications for the validity of the genus Morotopithecus

Morotopithecus bishopi and Afropithecus turkanensis are two large-bodied hominoid primates from early Miocene deposits of eastern Africa. Researchers have used both cranial and postcranial characters to distinguish these two species. Unfortunately, of the fossil material attributed to each, only the face, palate, and upper dentition are preserved well enough in both species for direct comparisons. There are currently no known directly comparable postcranial elements. In this study, we reevaluated dental characters argued to distinguish the type specimens of Morotopithecus from Afropithecus: relative size of the upper premolars and M 3. Exact randomization methods were used to address two questions. First, is it possible to find the degree of dental-size difference observed between Morotopithecus (UMP 62-11) and Afropithecus (KNM-WK 16999) within extant African hominoids? Second, what is the probability of observing the levels of difference found between the fossils among pairs of extant individuals? Metric differences in relative premolar and M 3 size were calculated between all possible pairs within the extant sample and the observed difference of the fossil pair was then compared to the resulting distribution of extant pairs. The observed size differences for all comparisons in the fossil teeth were well within the variation observed in the extant African hominoid samples ( p > 0.05). In light of these results and other currently available cranial evidence, we suggest that the type specimens of Morotopithecus and Afropithecus are not different enough to support taxonomic distinction.

The relationship between uric acid and its oxidative product allantoin: a potential indicator for the evaluation of oxidative stress in birds

Uric acid is the main nitrogenous waste product in birds but it is also known to be a potent antioxidant. Hominoid primates and birds lack the enzyme urate oxidase, which oxidizes uric acid to allantoin. Consequently, the presence of allantoin in their plasma results from non-enzymatic oxidation. In humans, the allantoin to uric acid ratio in plasma increases during oxidative stress, thus this ratio has been suggested to be an in vivo marker for oxidative stress in humans. We measured the concentrations of uric acid and allantoin in the plasma and ureteral urine of white-crowned sparrows (Zonotrichia leucophrys gambelii) at rest, immediately after 30 min of exercise in a hop/hover wheel, and after 1 h of recovery. The plasma allantoin concentration and the allantoin to uric acid ratio did not increase during exercise but we found a positive relationship between the concentrations of uric acid and allantoin in the plasma and in the ureteral urine in the three activity phases. In the plasma, the slope of the regression describing the above positive relationships was significantly higher immediately after activity. We suggest that the slope indicates the rate of uric acid oxidation and that during activity this rate increases as a result of higher production of free radicals. The present study demonstrates that allantoin is present in the plasma and in the ureteral urine of white-crowned sparrows and therefore might be useful as an indicator of oxidative stress in birds.

Molecular evolution of the HERV-E family in primates

More than 50 copies of HERV-E family elements have been estimated to exist in the human genome. Here we examined the recent evolutionary history of the HERV-E family by a PCR approach using genomic DNA from hominoid primates and a human monochromosomal panel. From the HERV-E family, 25 and 68 env fragments, were identified and analyzed from hominoid primates and human chromosomes 1, 2, 3, 4, 6, 7, 8, 10, 11, 12, 13, 14, 15, 16, 17, 19, X, and Y, respectively. They showed 76.7-99.6% sequence similarity to that of HERV-E (accession no. M10976). Phylogenetic analysis of HERV-E env family distinctively divided into two groups (groups I and II) that each contained three subgroups. Divergence times of the two groups were estimated as 10.7 MYr for group I and 41.3 MYr for group II using an average evolutionary rate of 0.3% per MYr. These data are consistent with that of PCR analysis, which showed a band of the HERV-E family in the genomes of the hominoids, Old World monkeys, and New World monkeys. Therefore, the HERV-E family may have integrated into the primate genome after prosimians and New World monkeys diverged. Then they proliferated extensively in the genome of humans and great apes during primate evolution.

Morphological evidence of marine adaptations in human kidneys

Amongst primates, kidneys normally exhibiting lobulated, multipyramidal, medullas is a unique attribute of the human species. Although, kidneys naturally multipyramidal in their medullary morphology are rare in terrestrial mammals, kidneys with lobulated medullas do occur in: elephants, bears, rhinoceroses, bison, cattle, pigs, and the okapi. However, kidneys characterized with multipyramidal medullas are common in aquatic mammals and are nearly universal in marine mammals. To avoid the deleterious effects of saline water dehydration, marine mammals have adaptively thickened the medullas of their kidneys--which enhances their ability to concentrate excretory salts in the urine. However, the lobulation of the kidney's medullary region in marine mammals appears to be an adaptation to expand the surface area between the medulla and the enveloping outer cortex in order to increase the volume of marine dietary induced hypertonic plasma that can be immediately processed for the excretion of excess salts and nitrogenous waste. A phylogenetic review of freshwater aquatic mammals suggest that most, if not all, nonmarine aquatic mammals inherited the medullary pyramids of their kidneys from ancestors who originally inhabited, or frequented, marine environments. So this suggest that most, if not all, aquatic mammals exhibiting kidneys with lobulated medullas are either marine adapted--or are descended from marine antecedents. Additionally, a phylogenetic review of nonhuman terrestrial mammals possessing kidneys with multipyramidal medullas suggest that bears, elephants and possibly rhinoceroses, also, inherited their lobulated medullas from semiaquatic marine ancestors. The fact that several terrestrial mammalian species of semiaquatic marine ancestry exhibit kidneys with multipyramidal medullas, may suggest that humans could have, also, inherited the lobulated medullas of their kidneys from coastal marine ancestors. And a specialized marine diet in ancient human ancestry could, also, explain the reactivation and enumeration of corporeal eccrine sweat glands and the copious secretion of salt tears. The substantial loss of genetic variation in humans relative to other hominoid primates, combined with the apparent isolation of early Pliocene human ancestors from particular retroviruses that infected all other African primate species, may suggest that such a semiaquatic marine phase, during the emergence of Homo, may have occurred on an island off the coast of Africa during the early Pliocene.

Barth syndrome: TAZ gene mutations, mRNAs, and evolution

Barth syndrome (MIM 302060) is an X-linked condition that includes dilated cardiomyopathy, neutropenia, failure to thrive, abnormal mitochondria, and 3-methylglutaconic aciduria. The mutated gene, TAZ, first described in 1996, appeared to produce a large set of alternatively spliced mRNAs with initiations of transcription upstream of exons 1 and 3. Since then, disease-causing mutations have been found in all exons including, most recently, a missense mutation in the controversial exon 5. Because of the initially described second initiation of transcription in intron 2, with in-frame initiation of translation in exon 3, we hypothesized that subjects with mutations in exons 1 and 2 would produce more normal "short product" that might attenuate their phenotype. Moreover, it was of interest to determine which splice variants were potentially functional as exon 5 is not present in yeast and rodents, and the variant lacking this exon is the most abundant. Using RT-PCR, we characterized TAZ mRNAs in cultured lymphocytes from nine subjects with Barth syndrome and two healthy controls. The TAZ genes and mRNAs of primates were also included. We found the following: (1) there is only one site for initiation of transcription, and the normal alternatively spliced assortment is limited to full-length, delta5, delta7, delta5delta7; (2) there are two alternative splice sites within introns 1 and 2 that could potentially produce an in-frame product; (3) exon 5 evolved into "exonhood" in the primate lineage after the split between Old World monkeys and hominoid primates; and (4) our results suggest that only two functional protein variants exist in lymphocytes: delta5 and full-length. Although exon 5 does not appear to be required for TAZ function in yeast and monkeys, its evolution to a highly conserved spliced exon in hominoid primates and the recent finding of an exon 5 mutation in a patient with Barth syndrome suggest that the full-length variant is important to TAZ function.

Quantification of dentine shape in anthropoid primates

The external shape and thickness of the enamel component of primate molars have been employed extensively in phylogenetic studies of primate relationships. The dentine component of the molar crown also has been suggested to be indicative of phylogenetic relationships, but few studies have quantified dentine morphology in order to evaluate this possibility. To explore the utility of dentine shape as an indicator of phylogenetic affinity, a two-dimensional geometric morphometric analysis (EDMA-II) was performed using nine homologous landmarks on a sample of sectioned maxillary molars of extant ceboid, cercopithecoid, and hominoid primates. Results indicate that dentine shape (the configuration of the enamel-dentine junction, or EDJ) can distinguish taxa at every taxonomic level examined, including superfamilies, subfamilies, and closely related genera and species. This supports the idea that dentine morphology may be useful for phylogenetic studies. It is further suggested that the morphology of the EDJ may be more conservative than enamel morphology, and perhaps better-suited to phylogenetic studies. Among the samples studied, cercopithecoid primates have a unique dentine shape, and it is suggested that the development of bilophodont molars may be related to the distinctive EDJ configuration in cercopithecoids.

Periodic explosive expansion of human retroelements associated with the evolution of the hominoid primate.

Five retroelement families, L1 and L2 (long interspersed nuclear element, LINE), Alu and MIR (short interspersed nuclear element, SINE), and LTR (long terminal repeat), comprise almost half of the human genome. This genome-wide analysis on the time-scaled expansion of retroelements sheds light on the chronologically synchronous amplification peaks of each retroelement family in variable heights across human chromosomes. Especially, L1s and LTRs in the highest density on sex chromosomes Xq and Y, respectively, disclose peak activities that are obscured in autosomes. The periods of young L1, Alu, LTR, and old L1 peak activities calibrated based on sequence divergence coincide with the divergence of the three major hominoid divergence as well as early eutherian radiation while the amplification peaks of old MIR and L2 account for the marsupial-placental split. Overall, the peaks of autonomous LINE (young and old L1s and L2s) peaks and non-autonomous SINE (Alus and MIRs) have alternated repeatedly for 150 million years. In addition, a single burst of LTR parallels the Cretaceous-Tertiary (K-T) boundary, an exceptional global event. These findings suggest that the periodic explosive expansions of LINEs and SINEs and an exceptional burst of LTR comprise the genome dynamics underlying the macroevolution of the hominoid primate lineage.

Evolution of the Hominoid Semenogelin Genes, the Major Proteins of Ejaculated Semen

The hominoid primates (apes and humans) exhibit remarkable diversity in their social and sexual behavioral systems. This is reflected in many ways in their anatomy and physiology. For example, the testes and seminal vesicles are relatively large in species with high sperm competition like the chimpanzee and small in species with low or no sperm competition like the gorilla. Additionally, the chimpanzee is the only hominoid primate known to produce a firm copulatory plug, which presumably functions in sperm competition by blocking insemination of subsequent males. Here we examine the molecular evolution of the semenogelin genes (SEMG1 and SEMG2), which code for the predominant structural proteins in human semen. High molecular weight complexes of these proteins are responsible for the viscous gelatinous consistency of human semen; their rodent homologs are responsible for the formation of a firm copulatory plug. Chimpanzees have an expanded SEMG1 gene caused by duplications of tandem repeats, each encoding 60 amino acids, resulting in a protein nearly twice as long as that of humans. In contrast, at both SEMG1 and SEMG2 we observed several gorilla haplotypes that contain at least one premature stop codon. We suggest that these structural changes in the semenogelin proteins that have arisen since the human-chimpanzee-gorilla split may be responsible for the physiological differences between these species ejaculated semen that correlate with their sociosexual behavior.

BooK Reviews

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Les nouveaux restes du squelette d’ Ugandapithecus major (Miocène inférieur de Napak, Ouganda)

New postcranial remains of Ugandapithecus major (Early Miocene, Uganda). During the 1998–2001 field seasons of the Uganda Palaeontology Expedition to Karamoja, several postcranial specimens of Miocene hominoid primates, including a proximal femur lacking the head, and an isolated femoral head were found that belong to a large bodied hominoid. The former specimen was found at Napak I and the latter at Napak IX. These remains increase the skeletal sample of Ugandapithecus major recently described at the site (Senut et al., 2000). The new specimens provide interesting information about the postcranial skeleton of the large ape species discovered by Bishop in the 1960’s and described by Rafferty and co-authors (Rafferty et al., 1995), and Gommery and co-authors (1998). The features observed on the various pieces suggest that Ugandapithecus major was a large-bodied tree climber. Comparisons with the Moroto postcranial elements reveals that Morotopithecus is a mixture of two taxa, one of which is Ugandapithecus major, the other Afropithecus turkanensis (Pickford, in press).

Phylogeny of the Neogene hominoid primates of Eurasia: v.2: Hominoid evolution and climatic change in Europe

Phylogeny of the Neogene hominoid primates of Eurasia: v.2: Hominoid evolution and climatic change in Europe

Isolation and phytogeny of SINE‐R retroposons derived from human endogenous retrovirus HERV‐K family in schizophrenia

SINE‐R retroposons have been derived from human endogenous retrovirus HERV‐K family and found to be hominoid specific. Both SINE‐R retroposons and HERV‐K family are potentially capable of affecting the expression of closely located genes. Using the genomic DNA from patients with schizophrenia, we identified 26 SINE‐R retroposons and analyzed them with the sequences derived from the hominoid primates. The SINE‐R retroposons from schizophrenia showed 89.7–96.6% sequence similarities with the sequence of the schizo‐cDNA clone that derived from postmortem tissue from the frontal cortex of an individual suffering from schizophrenia. Phylo‐genetic analysis using the neighbor‐joining method revealed that the new SINE‐R retroposons in schizophrenia have proliferated independently during hominid evolution. Such retroposons have great relevance to genomic change connected to human diseases. The data suggest that new SINE‐R retroposons identified in schizophrenia deserve further investigation as potential leads on t...

人類進化と新生代後期の環境変化 ヒト科の出現 中新世におけるヒト上科の展開

The human lineage is a branch of the huge evolutionary tree of the Hominoidea, which today includes ourselves, chimpanzees, gorillas, orangutans, and gibbons. All extant hominoids, except for the single world-wide species Homo sapiens, are now endangered and restricted in their distribution to tropical/subtropical forests of Africa and Southeast Asia. However, hominoids were once a more flourishing group millions of years ago. In the Miocene, there were various kinds of hominoid primates, and their distribution was much wider than today, from the southern tip of Africa through Arabia, to Europe in the west and through India to China in the east. It is from this great diversity of Miocene hominoids that the earliest human ancestors emerged, probably during the Late Miocene. The first part of this article provides readers with general information about extant hominoids. The second part is devoted to Miocene fossil hominoids with emphasis on African fossils. Although the main subject of this article is fossil hominoids, some recent findings of early hominids are also mentioned to at the end.

Human and ape molecular clocks and constraints on paleontological hypotheses.

Although the relationships of the living hominoid primates (humans and apes) are well known, the relationships of the fossil species, times of divergence of both living and fossil species, and the biogeographic history of hominoids are not well established. Divergence times of living species, estimated from molecular clocks, have the potential to constrain hypotheses of the relationships of fossil species. In this study, new DNA sequences from nine protein-coding nuclear genes in great apes are added to existing datasets to increase the precision of molecular time estimates bearing on the evolutionary history of apes and humans. The divergence of Old World monkeys and hominoids at the Oligocene-Miocene boundary (approximately 23 million years ago) provides the best primate calibration point and yields a time and 95% confidence interval of 5.4 +/- 1.1 million years ago (36 nuclear genes) for the human-chimpanzee divergence. Older splitting events are estimated as 6.4 +/- 1.5 million years ago (gorilla, 31 genes), 11.3 +/- 1.3 million years ago (orangutan, 33 genes), and 14.9 +/- 2.0 million years ago (gibbon, 27 genes). Based on these molecular constraints, we find that several proposed phylogenies of fossil hominoid taxa are unlikely to be correct.

Biomechanical scaling of the hominoid mandibular symphysis.

Experimental investigation of mandibular bone strain in cercopithecine primates has established that the mandible is bent in the transverse plane during the power stroke of mastication. Additional comparative work also supports the assumption that the morphology of the mandibular symphysis is functionally linked to the biomechanics of lateral transverse bending, or "wishboning" of the mandibular corpus. There are currently no experimental data to verify that lateral transverse bending constitutes an important loading regime among hominoid primates. There are, however, allometric models from cercopithecoid primates that allow prediction of scaling patterns in hominoid mandibular dimensions that would be consistent with a mechanical environment that includes wishboning as a significant component. This study uses computed tomography (CT) scans to visualize cortical bone distribution in the anterior corpus of a sample of four genera of extant hominoids. From the cortical bone contours, area properties of the mandibular symphysis are calculated, and these variables are subjected to an allometric analysis to detect whether scaling of jaw dimensions are consistent with a wishboning loading regime. Scaling of the hominoid symphysis recalls patterns observed in cercopithecoid monkeys, which lends indirect support for the hypothesis that wishboning is an integral part of the masticatory loading environment in living apes. Inclination of the symphysis, rather than changes in cross-sectional shape or development of the superior transverse torus, represents a morphological solution for minimizing the potentially harmful effects of wishboning in the jaws of these primates.

Intraspecies heterogeneity of immunoglobulin alpha-chain constant region genes in rhesus macaques.

Immunoglobulin A (IgA) is the major antibody class present in external secretions and is also an important component of serum immunoglobulins. On mucosal surfaces, IgA represents a first line of defence by neutralizing invading pathogens. The number of IgA constant-region genes (C alpha) present in different mammalian species is variable. Immunoglobulin C alpha genes differ mainly in the sequences located in the hinge region. IgA molecules, whose hinge regions are remarkably similar to those of the respective human molecules, are present in hominoid primates. In this report, we show that two alleles of a single immunoglobulin C alpha are present in rhesus macaques (Macaca mulatta). In addition, we show that intraspecies immunoglobulin C alpha allelic polymorphism is very high in this non-human primate species. Specifically, five different hinge regions, some of which are proline-rich, were identified from a total of eight rhesus macaque immunoglobulin C alpha-chains. The five hinge regions were different from those present in hominoid primates, both in length and in sequence. These results represent the first example of high levels of intraspecies immunoglobulin constant-region variability and suggest that IgAs of variable structure and function may be present in rhesus macaques. As rhesus macaques are widely used as animal models for the development of vaccines for acquired immune deficiency syndrome (AIDS), the possible presence of structurally and functionally variable IgA molecules in different animals should be taken into account when designing experimental strategies to induce mucosal antibody responses to human immunodeficiency virus (HIV).

Étude des hétérochronies par superposition procruste : application aux crânes de primates Hominoidea

The procrustes superimposition method is well adapted to heterochronic studies in the field of evolutionary biology. 1) The procrustes method gives a precise and mathematical definition of two of the three heterochronic variables: size and shape. 2) It allows us to describe complex anatomical structures and thus to analyse the whole structure and not just to proceed trait by trait. 3) The approach is statistical and the different hypotheses and results may be statistically tested. 4) When applied to heterochronies the method allows us to test if there is a common shape change related to allometry. In the present study of three species of Hominoid primates, the procrustes superimposition reveals that various heterochronic processes are simultaneously present. Size–age–shape dissociations between species, already present in the first ontogenetic stage, are amplified with growth until adult stage. As compared with that of the chimpanzee, the growth of the gorilla skull is accelerated in terms of size–shape covariation and size alone. The growth of the human skull is neotenic as compared with that of the apes.

Evolution of the X-linked Zinc Finger Gene and the Y-linked Zinc Finger Gene in Primates

We have sequenced the partial exon of the zinc finger genes (ZFX and ZFY) in 5 hominoids, 2 Old World monkeys, 1 New World monkey, and 1 prosimian. Among these primate species, the percentage similarities of the nucleotide sequence of the ZFX gene were 96-100% and 91.2-99.7% for the ZFY gene. Of 397 sites in the ZFX and ZFY gene sequences, 20 for ZFX gene and 42 for ZFY gene were found to be variable. Substitution causes 1 amino acid change in ZFX, and 5 in ZFY, among 132 amino acids. The numbers of synonymous substitutions per site (Ks) between human and the chimpanzee, gorilla and orangutan for ZFY gene were 0.026, 0.033, and 0.085, respectively. In contrast, the Ks value between human and hominoid primates for the ZFX gene was 0.008 for each comparison. Comparison of the ZFX and ZFY genes revealed that the synonymous substitution levels were higher in hominoids than in other primates. The rates of synonymous substitution per site per year were higher in the ZFY exon than in the SRY exon, and higher in the ZFY exon than in the ZFY intron, in hominoid primates.

Evolution of the X-linked zinc finger gene and the Y-linked zinc finger gene in primates.

We have sequenced the partial exon of the zinc finger genes (ZFX and ZFY) in 5 hominoids, 2 Old World monkeys, 1 New World monkey, and 1 prosimian. Among these primate species, the percentage similarities of the nucleotide sequence of the ZFX gene were 96-100% and 91.2-99.7% for the ZFY gene. Of 397 sites in the ZFX and ZFY gene sequences, 20 for ZFX gene and 42 for ZFY gene were found to be variable. Substitution causes 1 amino acid change in ZFX, and 5 in ZFY, among 132 amino acids. The numbers of synonymous substitutions per site (Ks) between human and the chimpanzee, gorilla and orangutan for ZFY gene were 0.026, 0.033, and 0.085, respectively. In contrast, the Ks value between human and hominoid primates for the ZFX gene was 0.008 for each comparison. Comparison of the ZFX and ZFY genes revealed that the synonymous substitution levels were higher in hominoids than in other primates. The rates of synonymous substitution per site per year were higher in the ZFY exon than in the SRY exon, and higher in the ZFY exon than in the ZFY intron, in hominoid primates.

The Evolution of neogene terrestrial ecosystems in Europe

1. Introduction J. Agusti, L. Rook and P. Andrews Part I. Palaeogeography of the Circum-Mediterranean Region: 2. Mediterranean and Paratethys palaeogeography during Oligocene and Miocene Fred Rogl 3. Pliocene tephra correlations between East African hominid localities, the Gulf of Aden and the Arabian Sea Peter B. deMenocal and Francis H. Brown 4. Climatic perspectives for neogene environmental reconstructions Eileen M. O'Brien and Charles R. Peters Part II. Miocene Mammalian Successions: 5. A critical re-evaluation of the Miocene mammals units in western Europe: dispersal events and the problems of correlation J. Agusti, L. Cabrera and M. Garces 6. Large mammals from the Vallesian of Spain Jorge Morales, Manuel Nieto, Meike Kholer and Salvador Moya-Sola 7. Trends in rodent assemblages from the Aragonian (Early-Middle Miocene) of the Calatayud-Daroca basin, Aragon, Spain R. Daams, A. J. van der Meulen, P. Pelaez-Campomanes and M. A. Alvarez-Sierra 8. The Late Miocene small mammal succession from France, with emphasis on the Rhone Valley localities Pierre Mein 9. Late Miocene mammals from central Europe Jens Lorenz Franzen and Gerharrd Storch 10. An overview on the Italian Miocene land mammal faunas Lorenzo Rook, Laura Abbazzi and Burkhart Engesser 11. The Miocene large mammal succession in Greece Louis de Bonis and George D. Koufos 12. Chronology and mammal faunas of the Miocene Sinap formation, Turkey Juha Pekka Lunkka, Mikael Fortelius, John Kappelman and Sevket Sen 13. The Late Miocene small mammal succession in Ukraine Valentin A. Nesin and Vadim A. Topachevsky Part III. Palaeoenvironments: Non-Mammalian Evidence: 14. Marine invertebrate (chiefly foraminiferal) evidence for the palaeogeography of the Oligocene-Miocene of western Eurasia, and consequences for terrestrial vertebrate migration Robert Wynn Jones 15. Palaeoclimatic implications of the energy hypothesis from Neogene corals of the Mediterranean region Brian R. Rosen 16. Contribution to the knowledge of Neogene climatic changes in western and central Europe by means of non-marine molluscs Daniela Esu 17. Sedimentary facies analysis in palaeoclimatic reconstructions. Examples from the Upper Miocene-Pliocene successions of south-central Tuscany (Italy) Marco Benvenuti, Mauro Papini and Giovanni Testa 18. Neogene vegetation changes in western European and west circum-Mediterranean areas Jean-Pierre Suc, Severine Fauquette, Mostefa Bessegik, Adele Bertini, Zhuo Zheng, Georges Clauzon, Danica Suballyova, Fiomena Diniz, Pierre Quezel, Najat Feddi, Martine Clet, Ezzedine Bessais, Naima Bachiri Taoufiq, Henriette Meon and Nathalie Combourieu-Nebout Part IV. Palaeoenvironments: Mammalian Evidence: 19. Shrews (Mammalia, Insectivora, Soricidae) as palaeoclimatic indicators in the European Neogene Jelle W.F. Reumer 20. Mammal turnover and global climate change in the Late Micoene terrestrial record of the Valles- Penedes basin (NE Spain) Jorge Agusti , Lluis Cabrera, Miguel Garces and Manel Llenas 21. Palaeoenvironments of Late Miocene primate localites in Macedonia, Greece L. de Bonis, G. Bouvrain and G. D. Koufos 22. The paleoecology of the Pikermian biome and the savanna myth Nikos Solounias, J. Michael Plavcan, Jay Quade and Lawrence Witmer 23. Vicariance biogeography and paleoecology of Eurasian Miocine hominoid primates Peter Andrews and Raymond L. Bernor 24. Conclusion J. Agusti, L. Rook and P. Andrews.