A whole new world of ancestors: Eocene anthropoideans from Africa

The last decade has witnessed enormous gains in our knowledge of early anthro-poidean primates,

1 “Anthropoidean” refers to members of the suborder Anthropoidea, whch contaings New and Old world monkeys, apes, and humans. These primates are also often called “simians,” “simiiforms” or “anthropoids;” the latter term is potentially confusing because it has often been used to refer only to the great apes.

the oldest known relatives of monkeys, apes and humans. Recent fossil finds in Egypt, Algeria, Tunisia, and Oman, along with the associated geological research at these sites, have radically changed our models of anthro-poidean origins and differentiation. Instead of first appearing as robust-jawed herbivorous primates in the earliest Oligocene, it is now apparent that there was radiation of small-bodied, fruit-and-insect-eating anthropoideans during the Eocene. These early forms included at least two oligopithecines (squirrel-monkey-sized primates with a catarrhine dental formula) and two early “parapithecid monkeys” (three-premolared primates with lumpy, bunodont dentitions). In addition, several smaller species from Algeria and Egypt, ranging in size from pygmy marmosets to tamarins, are not definitely assignable to previously known families. Alongside the early anthropoideans, there are also at least four recently identified prosimian families. The continental Eocene of Africa—for years, little more than a blank on the paieontologi-cal map—now comprises an increasingly productive field source of new data that is important in deciphering phyletic and adaptive aspects of the prosimian-anthropoid transition.     

Foot bones from Omo: implications for hominid evolution

We reanalyze a hominid talus and calcaneus from Omo dating to 2.2 mya and 2.36 mya, respectively. Although both specimens occur at different localities and times, both tarsals articulate well together, suggesting a single taxon on the basis of size and function. We attribute these foot bones to early Homo on the basis of their morphology. The more modern-like tarsal morphology of these Omo foot bones makes them very similar to a talus from Koobi Fora (KNM-ER 813), a specimen attributed to Homo rudolfensis or Homo erectus. Although the Omo tarsals are a million years younger than the oldest known foot bones from Hadar, both localities demonstrate anatomical differences representing two distinct morphological patterns. Although all known hominid tarsals demonstrate clear bipedal features, the tarsal features noted below suggest that biomechanical changes did occur over time, and that certain features are associated with different hominid lineages (especially the robust australopithecines).     

Birth of 'human-specific' genes during primate evolution

Humans and other Anthropoids share very similar chromosome structure and genomic sequence as seen in the 98.5% homology at the DNA level between us and Great Apes. However, anatomical and behavioral traits distinguish Homo sapiens from his closest relatives. I review here several recent studies that address the issue by using different approaches: large-scale sequence comparison (first release) between human and chimpanzee, characterization of recent segmental duplications in the human genome and analysis of exemplary gene families. As a major breakthrough in the field, the heretical concept of human-specific genes has recently received some supporting data. In addition, specific chromosomal regions have been mapped that display all the features of gene nurseries and could have played a major role in gene innovation and speciation during primate evolution. A model is proposed that integrates all known molecular mechanisms that can create new genes in the human lineage.     

Introduction: Current studies on primate socioecology and evolution

Primate socioecology is being dramatically transformed as new studies are undertaken in Madagascar and the New World. Simultaneously, the incorporation of primate studies in a broader framework of theory in behavioral ecology promises to more intelligently inform anthropology's traditional concerns with primate and hominid evolution.     

Understanding primate brain evolution

We present a detailed reanalysis of the comparative brain data for primates, and develop a model using path analysis that seeks to present the coevolution of primate brain (neocortex) and sociality within a broader ecological and life-history framework. We show that body size, basal metabolic rate and life history act as constraints on brain evolution and through this influence the coevolution of neocortex size and group size. However, they do not determine either of these variables, which appear to be locked in a tight coevolutionary system. We show that, within primates, this relationship is specific to the neocortex. Nonetheless, there are important constraints on brain evolution; we use path analysis to show that, in order to evolve a large neocortex, a species must first evolve a large brain to support that neocortex and this in turn requires adjustments in diet (to provide the energy needed) and life history (to allow sufficient time both for brain growth and for 'software' programming). We review a wider literature demonstrating a tight coevolutionary relationship between brain size and sociality in a range of mammalian taxa, but emphasize that the social brain hypothesis is not about the relationship between brain/neocortex size and group size per se; rather, it is about social complexity and we adduce evidence to support this. Finally, we consider the wider issue of how mammalian (and primate) brains evolve in order to localize the social effects.     

Non-marine Teredolites from the Middle Eocene of southern England

Middle Eocene, non-marine sediments from southern England contain examples of Teredolites borings in two contrasting palaeoenvironmental settings, viz.: (A) as in situ borings in an allochthonous lignite in an abandoned river channel and (B) as bored logs in cross-bedded fluvial sandstones of probable point-bar origin. The lignite is 0.30 m thick, of which the upper 0.20 m is intensely bored. Rounded pebbles of ?charcoal at its base also show small borings. A log in a fluvial sandstone shows densely-packed. radial club-shaped borings. now filled with sandstone. The wood substrate has subsequently been oxidized away. To our knowledge, this is the first detailed account of Teredolites from an ancient freshwater setting.     

Extractive foraging and the evolution of primate intelligence

One of the two major theories regarding the evolution of intelligence in primates is that feeding strategies determine mental development. Evidence for this theory is reviewed and related to extractive foraging, which is the act of locating and/or processing embedded foods such as underground roots and insects or hard-shelled nuts and fruits. It is shown that, although only cebus monkeys and chimpanzees in the wild use tools in extractive foraging, many other species of mammals (including primates) and birds are capable of extracting embedded foods without tools. Extractive foraging by primates is compared to extractive foraging by other mammals and birds to assess whether: 1) extractive foraging involves cognition, and 2) extractive foraging by primates is unique in a way that may mean it played a role in the development of intelligence among primates. This comparison reveals that some acts of extractive foraging by nonprimates are equally sophisticated as those of primates. It is suggested that extractive foraging played no significant role in the evolution of primate intelligence. Hypotheses for testing precise differences in extractive foraging ability across taxa are offered, and the roles of olfactory cues, manual dexterity, and strength in extractive foraging are evaluated. In conclusion, the hominization process is briefly reviewed in relation to foraging behavior. A ?package? of traits that, in combination, is unique to hominids is discussed: tool-aided extractive foraging, division of labor by sex with food exchange, and feeding of juveniles.     

Studies of primate protein variation and evolution: Microelectrophoretic detection

Genetic variation at 16 protein and enzyme loci in Cercopithecus aethiops and several other primate species has been surveyed, using cellulose acetate microelectrophoresis. Resolution of several standard variant proteins is comparable to that achieved on starch gel or polyacrylamide gel. Although both intraspecific and interspecific variation was observed for some loci, the data generally support the concept that extracellular proteins are more likely to be polymorphic within a species, while intracellular proteins generally vary between species, if at all. These methodologies are particularly appropriate for screening multiple-locus variation in large numbers of samples; their relevance to studies of molecular evolution and evaluation of theories of kin selection is discussed.This research was supported in part by California State Agricultural Experiment Station Funds to the University of California, Berkeley, and the Wenner Gren Foundation.     

Built to last: The structure,function, and evolution of primate dental enamel

The teeth of every primate, living and extinct, are covered by a hard, durable layer of enamel. This is not unique: Almost all mammals have enamel-covered teeth. In addition, all of the variations in enamel structure that occur in primates are also found in other groups of mammals. Nevertheless, the very complexity of enamel and the variation we see in it on the teeth of living and fossil primates raise questions about its evolutionary significance. Is the complex structure of primate enamel adaptive? What, if anything, does enamel structure tell us about primate phylogeny? To answer these questions, we need to look more closely at the characteristics of prismatic enamel in primates and at the distribution of those characteristics, both in relation to our knowledge of primate dental function and feeding ecology and from a phylogenetic perspective.     

Ramapithecus and Sivapithecus from China: Some implications for higher primate evolution

The pattern of overall dental dimensions in over 900 teeth of ramapithecines from Lufeng in China is examined using frequency distribution histograms and fitted normal curves, and compared with data for extant hominoids. A prior study has demonstrated unequivocally that at least two groups of animals must have existed at Lufeng [Wu and Oxnard, 1983; Oxnard, 1983a]. The present investigation confirms this finding in more detail. In addition it shows that one fossil group possesses smaller teeth with a lesser degree of sexual dimorphism and approximately equal numbers of adult males and females, and the other possesses larger teeth with a rather larger degree of sexual dimorphism and a female-male ratio that may have approximated from as low as 2:1 to as high as 4:1. Comparisons of patterns of difference along the tooth row demonstrate that both these forms differ from modern apes in their sexual dimorphism, the smaller form being more like humans than the larger, which is more like apes, especially orangutans. Comparisons of the areas of the canine teeth with each of the other functional segments of the tooth row again show that the smaller form is basically similar to modern humans and that the larger resembles extant great apes. Comparisons of other functional dental areas seem to relate to dietary and masticatory functions. Thus the cutting areas are large relative to the chewing areas in omnivorous humans, whereas in the essentially vegetarian great apes this ratio is smaller. The smaller fossil resembles the human condition and may have been somewhat omnivorous; the larger one more resembles the apes and may have been somewhat more vegetarian. However, these comparisons also show that the way in which the larger form resembles the apes is associated with special development of the canines, which is different from that in any modern ape. Comparisons show that the canines in the larger form project far beyond the normal line of tooth crowns. Finally, comparisons show that canine sexual dimorphism in height is marked in the larger form. Neither of these last two features is true of the smaller fossil. These findings have implications for our understanding of the evolution of early pongids and hominids, and for the evolution of primate sexual dimorphisms and dental mechanisms.     

Hindlimb adaptations in Ourayia and Chipetaia, relatively large-bodied omomyine primates from the Middle Eocene of Utah

North American omomyids represent a tremendous Eocene radiation of primates exhibiting a wide range of body sizes and dietary patterns. Despite this adaptive diversity, relatively little is known of the postcranial specializations of the group. Here we describe hindlimb and foot bones of Ourayia uintensis and Chipetaia lamporea that were recovered from the Uinta B member (early Uintan Land Mammal Age), Uinta Formation, Utah. These specimens provide insights into the evolution of postcranial adaptations across different body sizes and dietary guilds within the Eocene primate radiation. Body mass estimates based on talar measurements indicate that Ourayia uintensis and Chipetaia lamporea weighed about 1,500-2,000 g and 500-700 g, respectively. Skeletal elements recovered for Ourayia include the talus, navicular, entocuneiform, first metatarsal, and proximal tibia; bones of Chipetaia include the talus, navicular, entocuneiform, and proximal femur. Both genera had opposable grasping big toes, as indicated by the saddle-shaped joint between the entocuneiform and first metatarsal. Both taxa were arboreal leapers, as indicated by a consistent assemblage of characters in all represented bones, most notably the somewhat elongated naviculars, the high and distinct trochlear crests of the talus, the posteriorly oriented tibial plateau (Ourayia), and the cylindrical head of the femur (Chipetaia). The closest resemblances to Ourayia and Chipetaia are found among the Bridger omomyines, Omomys and Hemiacodon. The results of our comparisons suggest that the later, larger, more herbivorous omomyines from Utah retained a skeletal structure characteristic of earlier, smaller North American omomyids.     

The Fayum primate forest: Did it exist?

The primates have the reputation of being essentially arboreal, forest-adapted animals. Yet there are many genera and species that inhabit an extremely wide array of non-forest habitats. Nevertheless, palaeoprimatologists often tend to depict fossil primate habitats as being more arboreal and more forest-like than is justified by the facts. It is worthwhile, therefore, to reconsider some current interpretations. In this paper, evidence of the Fayum Oligocene primate deposits are reviewed and discussed. The following conclusions emerge:(1) The large number of primate species indicates that the Fayum ecosystem was an optimum or near-optimum habitat for primates. (2) The lithological characteristics point to a sahélien type of climate. (3) The calcified and silicified root systems, having diameters up to 4 cm, suggest a sahélien type of shrub, bushland and/or small-tree vegetation. (4) The large fossilized logs cannot have grown on the spot and apparently represent driftwood from a more humid climatic belt in the south, as is indicated by damage resulting from fluvial transportation and by palaeobotanical data. (5) There may have been some minor patches or strips of medium-height forests and/or wood-lands in the Fayum delta, but there is no evidence of these.Thus the tall forest in which the earliest known African primates are currently supposed to have lived probably never existed. Grounds for this conclusion were presented by Unger 121 years ago, by Beadnell 75 years ago and by Kräusel 41 years ago, but sank into oblivion. The classic image of the primates as arboreal specialists seems to have interfered with seeing the facts. However, more extensive verification of the evidence by means of palaeobotanical research is still required. The fossil material to do so is readily available.     

Understanding hind limb weight support in chimpanzees with implications for the evolution of primate locomotion

Most quadrupedal mammals support a larger amount of body weight on their forelimbs compared with their hind limbs during locomotion, whereas most primates support more of their body weight on their hind limbs. Increased hind limb weight support is generally interpreted as an adaptation that reduces stress on primates' highly mobile forelimb joints. Thus, increased hind limb weight support was likely vital for the evolution of primate arboreality. Despite its evolutionary importance, the mechanism used by primates to achieve this important kinetic pattern remains unclear. Here, we examine weight support patterns in a sample of chimpanzees (Pan troglodytes) to test the hypothesis that limb position, combined with whole body center of mass position (COM), explains increased hind limb weight support in this taxon. Chimpanzees have a COM midway between their shoulders and hips and walk with a relatively protracted hind limb and a relatively vertical forelimb, averaged over a step. Thus, the limb kinematics of chimpanzees brings their feet closer to the COM than their hands, generating greater hind limb weight support. Comparative data suggest that these same factors likely explain weight support patterns for a broader sample of primates. It remains unclear whether primates use these limb kinematics to increase hind limb weight support, or whether they are byproducts of other gait characteristics. The latter hypothesis raises the intriguing possibility that primate weight support patterns actually evolved as byproducts of other traits, or spandrels, rather than as adaptations to increase forelimb mobility. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Leopard predation and primate evolution

Although predation is an important driving force of natural selection its effects on primate evolution are still not well understood, mainly because little is known about the hunting behaviour of the primates' various predators. Here, we present data on the hunting behaviour of the leopard (Panthera pardus), a major primate predator in the Tai; forest of Ivory Coast and elsewhere. Radio-tracking data showed that forest leopards primarily hunt for monkeys on the ground during the day. Faecal analyses confirmed that primates accounted for a large proportion of the leopards' diet and revealed in detail the predation pressure exerted on the eight different monkey and one chimpanzee species. We related the species-specific predation rates to various morphological, behavioural and demographic traits that are usually considered adaptations to predation (body size, group size, group composition, reproductive behaviour, and use of forest strata). Leopard predation was most reliably associated with density, suggesting that leopards hunt primates according to abundance. Contrary to predictions, leopard predation rates were not negatively, but positively, related to body size, group size and the number of males per group, suggesting that predation by leopards did not drive the evolution of these traits in the predicted way. We discuss these findings in light of some recent experimental data and suggest that the principal effect of leopard predation has been on primates' cognitive evolution.     

Genetics and the evolution of primate enamel thickness: a baboon model

The thickness of mammalian tooth enamel plays a prominent role in paleontology because it correlates with diet, and thicker enamel protects against tooth breakage and wear. Hominid evolutionary studies have stressed the importance of this character for over 30 years, from the identification of "Ramapithecus" as an early Miocene hominid, to the recent discovery that the earliest hominids display molar enamel intermediate in thickness between extant chimpanzees and Australopithecus. Enamel thickness remains largely unexplored for nonhominoid primate fossils, though there is significant variation across modern species. Despite the importance of enamel thickness variation to primate evolution, the mechanisms underlying variation in this trait have not yet been elucidated. We report here on the first quantitative genetic analysis of primate enamel thickness, an analysis based on 506 pedigreed baboons from a captive breeding colony. Computed tomography analysis of 44 Papio mandibular molars shows a zone of sufficiently uniform enamel thickness on the lateral surface of the protoconid. With this knowledge, we developed a caliper metric measurement protocol for use on baboon molars worn to within this zone, enabling the collection of a data set large enough for genetic analyses. Quantitative genetic analyses show that a significant portion of the phenotypic variance in enamel thickness is due to the additive effects of genes and is independent of sex and tooth size. Our models predict that enamel thickness could rapidly track dietary adaptive shifts through geological time, thus increasing the potential for homoplasy in this character. These results have implications for analyses of hominoid enamel thickness variation, and provide a foundation from which to explore the evolution of this phenotype in the papionin fossil record.     

Brief communication: New primate remains from the Miocene of Namibia,Southern Africa

Miocene primates from southern Africa are extremely rare. For this reason we wish to place on record several interesting new fossil primate specimens recently recovered from the Miocene sites of Berg Aukas and Harasib in the Otavi Mountain region of northern Namibia. The new finds consist of a virtually complete atlas vertebra from Berg Aukas attributable to the hominoid Otavipithecus namibiensis and two teeth and four postcranial fragments from Harasib referrable to Cercopithecoidea. The atlas vertebra exhibits anatomical characteristics intermediate between those of modern cercopithecoids and hominoids which may be indicative of a transition from pronograde to orthograde postures. The cercopithecoid remains show that the earliest Old World monkeys known from southern Africa were small, approximately the size of vervet monkeys. These new specimens are important because they provide the first evidence relating to possible positional behaviors of Otavipithecus and the earliest fossil record of cercopithecoids from southern Africa. © 1996 Wiley-Liss, Inc.     

Origin and evolution of primate social organisation: a reconstruction

The evolution and origin of primate social organisation has attracted the attention of many researchers, and a solitary pattern, believed to be present in most nocturnal prosimians, has been generally considered as the most primitive system. Nocturnal prosimians are in fact mostly seen alone during their nightly activities and therefore termed 'solitary foragers', but that does not mean that they are not social. Moreover, designating their social organisation as 'solitary', implies that their way of life is uniform in all species. It has, however, emerged over the last decades that all of them exhibit not only some kind of social network but also that those networks differ among species. There is a need to classify these social networks in the same manner as with group-living (gregarious) animals if we wish to link up the different forms of primate social organisation with ecological, morphological or phylogenetic variables. In this review, we establish a basic classification based on spatial relations and sociality in order to describe and cope properly with the social organisation patterns of the different species of nocturnal prosimians and other mammals that do not forage in cohesive groups. In attempting to trace the ancestral pattern of primate social organisation, the Malagasy mouse and dwarf lemurs and the Afro-Asian bushbabies and lorises are of special interest because they are thought to approach the ancestral conditions most closely. These species have generally been believed to exhibit a dispersed harem system as their pattern of social organisation ('dispersed' means that individuals forage solitarily but exhibit a social network). Therefore, the ancestral pattern of primate social organisation was inferred to be a dispersed harem. In fact, new field data on cheirogaleids combined with a review of patterns of social organisation in strepsirhines (lemurs, bushbabies and lorises) revealed that they exhibit either dispersed multi-male systems or dispersed monogamy rather than a dispersed harem system. Therefore, the concept of a dispersed harem system as the ancestral condition of primate social organisation can no longer be supported. In combination with data on social organisation patterns in 'primitive' placentals and marsupials, and in monotremes, it is in fact most probable that promiscuity is the ancestral pattern for mammalian social organisation. Subsequently, a dispersed multi-male system derived from promiscuity should be regarded as the ancestral condition for primates. We further suggest that the gregarious patterns of social organisation in Aotus and Avahi, and the dispersed form in Tarsius evolved from the gregarious patterns of diurnal primates rather than from the dispersed nocturnal type. It is consequently proposed that, in addition to Aotus and Tarsius, Avahi is also secondarily nocturnal.