New primates from the late Eocene of Thailand: a contribution to primate diversity in the Paleogene of Asia

We report the discovery of two new primates from the late Eocene Krabi Basin, southern Thailand. One isolated upper molar displays morphological features (protocone united with hypocone by the prehypocrista, postprotocrista extended distobuccally) suggesting possible phylogenetic relationships with Amphipithecidae, while an isolated lower molar is tentatively referred to as a new tarsiiform, mainly on the basis of its paraconid and entoconid morphology. Although very scarce, these remains expand the record of Paleogene primates in Southeast Asia, and testify to their successful radiation in that area.     

The phylogenetic significance of strepsirhinism in Paleogene primates

Two lines of evidence reviewed here argue against the recent proposal that strepsirhinism is an apomorphous feature in primates, shared only by adapiforms and lemuriforms. These are (1) the presence of strepsirhinism in several extant taxa of nonprimate mammals, including Tupaiidae, Tenrecidae, Erinaceus,and Didelphis,and (2) the inferred presence of strepsirhinism not only in adapiforms, but also in all plesiadapiforms and omomyids for which the relevant anatomical regions are known. Therefore, strepsirhinism cannot be invoked as an adaptive innovation underlying the initial strep-sirhine/haplorhine dichotomy. Likewise, the apparent retention of strepsirhinism in omomyids suggests either that the haplorhine oronasal configuration of extant tarsiids and anthropoids was acquired independently or that tarsiids and anthropoids form a clade to the exclusion of omomyids.     

Estimating the body size of eocene primates: A comparison of results from dental and postcranial variables

Estimating body weights for fossil primates is an important step in reconstructing aspects of their behavior and ecology. To date, the body size of Eocene euprimates—the Adapidae and Omomyidae—has been estimated only from molar area. Studies on other primates and mammals demonstrate that body weights estimated from teeth are not always concordant with those estimated from postcranial variables. We derive estimates for Eocene primates based on tarsal bone variables to compare with previously published values derived from dental measures. Stepsirhine-wide, family-level, and subfamily-level models are developed and compared. We also compare the accuracy and precision of dental- and tarsal-based regression models for predicting weight in extant species. Tarsal bone and dental area measures prove to be equally robust in predicting body weight; however, highly disparate estimates are often obtained from different variables. Equations based on lower-level taxonomic groups perform better than more widely based models. However, all equations considered yield fairly large errors, which can affect interpretations of paleoecology. The choice of the more robust prediction is not straightforward.     

Significance of primate petrosal from Middle Eocene fissure-fillings at Shanghuang, Jiangsu Province, People's Republic of China

An isolated petrosal bone belonging to a diminutive primate is reported from Middle Eocene fissure-fills near Shanghuang (southern Jiangsu Province, People's Republic of China), the type locality of several newly described primates (Eosimias sinensis, a basal anthropoid; Adapoides troglodytes, a basal adapinan; Tarsius eocaenus, a congener of extant tarsiers; and Macrotarsius macrorhysis, the first Asian representative of an otherwise exclusively North American genus). Because of its fragmentary condition and unique combination of characters, the Shanghuang petrosal cannot be assigned unambiguously to any of the Shanghuang primate taxa known from dental remains. However, the possibility that the petrosal represents either an adapid or a tarsiid can be dismissed because it lacks defining basicranial apomorphines of these groups. By contrast, the element does present arterial features consistent with its being haplorhine. Deciding between the likeliest candidates for its allocation—Omomyidae and Eosimiidae—is difficult, in part because it is not known what (or even whether) basicranial characters can be used to distinguish these clades. If the Shanghuang petrosal is that of an cosimiid, as both direct and indirect evidence appears to indicate, the following implications emerge: (1) as long suspected on other grounds, anthropoids share a closer evolutionary history with Omomyidae (and Tarsiiformes) than they do with Adapidae (and Strepsirhini); (2) the specialised basicranial anatomy of extant anthropoids and their immediate cladistic relatives is derived from a primitive precursor whose otic morphology was like that of omomyids in most known respects; (3) the evolution of the defining dental and basicranial apomorphies of extant Anthropoidea has been distinctly mosaic in pattern.     

Functional anatomy of the tarsier foot

Tarsiers possess a very odd musculoskeletal foot anatomy that goes beyond their acknowledged specialized leaping adaptations. Tarsius has evolved a fundamentally different method of bone rotation to achieve an inverted foot position during grasping and has developed an unusual muscular system for holding onto vertical supports. Although galagos and tarsiers possess elongated foot bones as adaptations for leaping, galagos utilize many more types of movements, have specialized osteological surface for inversion, and have a more common type of muscle development in the foot and leg than tarsiers possess. Likewise, the Omomyidae, the ancestral lineage of Tarsius, exhibit a lack of morphological similarity with Tarsius in the known foot joints.     

The distal tibia of primates with special reference to the omomyidae

The morphology of the distal tibia and its joint surfaces is described in the late Eocene European Necrolemur,the middle Eocene North American Hemiacodon,and an omomyid species from the lower part of the Bridger Formation of North America. Necrolemur,like Tarsius,exhibits tibiofibular fusion, although to a less advanced degree. The Bridger omomyids, however, show no evidence of fusion but are similar to galagos in the conformation of this joint. The distal tibia of euprimates is distinguished by several derived features. These correlate with derived features of the astragalus and are functionally related to the abduction of the foot that accompanies dorsiflexion in primates. Tarsius,omomyids, and anthropoids share a suite of features which distinguish them from strepsirhines; these maybe haplorhine synapomorphies, but the polarity of these features is difficult to determine. If they are synapomorphies, abduction accompanying dorsiflexion and movement at the inferior tibiofibular joint were restricted in ancestral haplorhines. In living primates such restriction is associated with small body size and saltatorial locomotion.     

Evolution of eye size and shape in primates

Strepsirrhine and haplorhine primates exhibit highly derived features of the visual system that distinguish them from most other mammals. Comparative data link the evolution of these visual specializations to the sequential acquisition of nocturnal visual predation in the primate stem lineage and diurnal visual predation in the anthropoid stem lineage. However, it is unclear to what extent these shifts in primate visual ecology were accompanied by changes in eye size and shape. Here we investigate the evolution of primate eye morphology using a comparative study of a large sample of mammalian eyes. Our analysis shows that primates differ from other mammals in having large eyes relative to body size and that anthropoids exhibit unusually small corneas relative to eye size and body size. The large eyes of basal primates probably evolved to improve visual acuity while maintaining high sensitivity in a nocturnal context. The reduced corneal sizes of anthropoids reflect reductions in the size of the dioptric apparatus as a means of increasing posterior nodal distance to improve visual acuity. These data support the conclusion that the origin of anthropoids was associated with a change in eye shape to improve visual acuity in the context of a diurnal predatory habitus.     

Canine size and shape in male anthropoid primates

While studies of canine dimorphism in primates are common, only a few have examined canine tooth size independently within each sex. Recently, Greenfield and Washburn (Am. J. Phys. Anthropol. 84:17–34, 1991) proposed that there are two types of male canines which reflect different allometric scaling patterns of canine crown height against canine occlusal dimensions. They also suggest that proportional canine shape, measured as canine crown height (or projection) relative to the occlusal dimensions, provides an estimate of the canine's function as a weapon, though they provide no test of this hypothesis. This analysis critically examines the claim that there are two types of male canines among anthropoids. It then tests the hypothesis that relative male canine size (measured against body weight) and proportional canine shape are related to estimates of intermale competition, diet, and substrate (used as a surrogate measure of predation pressure). While there is strong taxonomic variation in canine size and shape among male anthropoids, no evidence is found for two discrete canine types. Rather, within families and subfamilies, canine dimensions scale isometrically against body weight and against each other, with a continuum of canine shapes among different taxa. While variation in male canine size is associated with intermale competition and substrate, even when taxonomic variation is controlled, variation in proportional canine shape is not. Neither canine size nor shape are generally associated with variation in diet. © 1993 Wiley-Liss, Inc.     

New postcranial elements for the earliest Eocene fossil primate Teilhardina belgica

Teilhardina belgica is one of the most primitive fossil primates known to date and the earliest haplorhine with associated postcranials, making it relevant to a reconstruction of the ancestral primate morphotype. Here we describe newly discovered postcranial elements of T. belgica. It is a small primate with an estimated body mass between 30 and 60 g, similar to the size of a mouse lemur. Its hindlimb anatomy suggests frequent and forceful leaping with excellent foot mobility and grasping capabilities. It can now be established that this taxon exhibits critical primate postcranial synapomorphies such as a grasping hallux, a tall knee, and nailed digits. This anatomical pattern and behavioral profile is similar to what has been inferred before for other omomyids and adapiforms. The most unusual feature of T. belgica is its elongated middle phalanges (most likely manual phalanges), suggesting that this early primate had very long fingers similar to those of living tarsiers.     

Intrasexual competition and canine dimorphism in anthropoid primates.

A number of factors, including sexual selection, body weight, body-weight dimorphism, predation, diet, and phylogenetic inertia have been proposed as influences on the evolution of canine dimorphism in anthropoid primates. Although these factors are not mutually exclusive, opinions vary as to which is the most important. The role of sexual selection has been questioned because mating system, which should reflect its strength, poorly predicts variation in canine dimorphism, particularly among polygynous species. Kay et al. (1988) demonstrate that a more refined estimate of intermale competition explains a large proportion of the variation in canine dimorphism in platyrrhine primates. We expand their analysis, developing a more generalized measure of intermale competition based on the frequency and intensity of male-male agonism. We examine the relative influences of predation (inferred by substrate use), female body weight, body-weight dimorphism, diet, and sexual selection on the evolution of anthropoid canine dimorphism. Intermale competition is very strongly associated with canine dimorphism. Predation also has a marked effect on canine dimorphism, in that savanna-dwelling species consistently show greater canine dimorphism than other species, all other factors being held equal. Body-weight dimorphism is also strongly associated with canine dimorphism, though apparently through a common selective basis, rather than through allometric effects. Body weight seems to play only a minor, indirect role in the evolution of canine dimorphism. Diet plays no role. Likewise, we find little evidence that phylogenetic inertia is a constraint on the evolution of canine dimorphism.     

Allometric scaling in the dentition of primates and prediction of body weight from tooth size in fossils

Tooth size varies exponentially with body weight in primates. Logarithmic transformation of tooth crown area and body weight yields a linear model of slope 0.67 as an isometric (geometric) baseline for study of dental allometry. This model is compared with that predicted by metabolic scaling (slope = 0.75). Tarsius and other insectivores have larger teeth for their body size than generalized primates do, and they are not included in this analysis. Among generalized primates, tooth size is highly correlated with body size. Correlations of upper and lower cheek teeth with body size range from 0.90–0.97, depending on tooth position. Central cheek teeth (P and M) have allometric coefficients ranging from 0.57–0.65, falling well below geometric scaling. Anterior and posterior cheek teeth scale at or above metabolic scaling. Considered individually or as a group, upper cheek teeth scale allometrically with lower coefficients than corresponding lower cheek teeth; the reverse is true for incisors. The sum of crown areas for all upper cheek teeth scales significantly below geometric scaling, while the sum of crown areas for all lower cheek teeth approximates geometric scaling. Tooth size can be used to predict the body weight of generalized fossil primates. This is illustrated for Aegyptopithecus and other Eocene, Oligocene, and Miocene primates. Regressions based on tooth size in generalized primates yield reasonable estimates of body weight, but much remains to be learned about tooth size and body size scaling in more restricted systematic groups and dietary guilds.     

Patterns of dental emergence in early anthropoid primates from the Fayum Depression,Egypt

Abstract

Paleontological field work in the Fayum Depression of Egypt has produced a remarkable diversity of fossil anthropoids, and this, combined with advances in genetic analyses of living anthropoids, has led to establishment of a temporal and phylogenetic framework for anthropoids that is achieving some degree of consensus. Less well understood are the evolutionary mechanisms and selective factors behind the origin and early diversification of anthropoids. One area that has remained under explored is investigation into the life history patterns of early anthropoids, a major omission given that understanding patterns of growth and development is essential for interpreting the paleobiology of fossil species. Here we detail dental emergence sequences for five species in four families of early anthropoid primates from the Fayum, and use these data to test Schultz’s Rule concerning the timing of emergence of molars versus premolars in mammals. Two important results are generated: (1) only one species had a dental eruption sequence identical to that observed among crown catarrhine primates; and (2) in all cases, the permanent canine was the last post-incisor dental element to fully erupt, a finding that may be significant for interpreting early anthropoid behavioral strategies.     

Variation in the shape of the palate in adapis (Eocene,Adapiformes) compared with living primates

This study is a quantitative analysis of the variation of palate shape inAdapis, an Eocene fossil primate. The observed coefficients of variation of palate dimensions inAdapis are statistically greater than in most living primate species. The histogram of the index (palate width/palate length) emphasizes the great heterogeneity in the fossil species. Two allometric scalings of palate dimensions (length and width) on condylobasal length were performed. Plots of perpendicular distances from the two regression lines show that living species are always more concentrated around the middle of the diagrams than isAdapis. The space occupied byAdapis is as great as, or even greater, than those occupied by living genera. Hence, several species are certainly presenent in the fossil sample, and the varieties distinguished by Stehlin in 1912, are in fact valid species names.     

New Uintan primates from Texas and their implications for North American patterns of species richness during the Eocene

New omomyid fossils from the Purple Bench locality of the Devil's Graveyard Formation, middle Eocene (Uintan) of southwest Texas, are described. One specimen represents a new genus and species, herein named Diablomomys dalquesti. This new species is allocated to the tribe Omomyini, sister taxon to Omomys and Chumashius. A second specimen represents a range extension of the Utah species Mytonius hopsoni to the Trans-Pecos region of Texas. Previously, only one omomyid species (Omomys carteri) had been documented from Purple Bench and other late Uintan localities in the Devil's Graveyard Formation. These new omomyid fossils are of particular significance because Purple Bench is stratigraphically intermediate between the older late Bridgerian/early Uintan localities and the younger Duchesnean localities of Trans-Pecos Texas. With a more southerly location in the continental United States, the Devil's Graveyard Formation amplifies our understanding of patterns of North American primate richness at a time when the higher-latitude sites of the western interior were undergoing significant climatic cooling and increases in seasonality with commensurate faunal reorganization. Although the Uintan (approximately 46.5-40Ma) was a time in which anaptomorphine richness decreased dramatically, the results of this analysis suggest that Uintan omomyine richness is higher than was previously appreciated, particularly at lower latitudes.     

Intrasexual competition and body weight dimorphism in anthropoid primates

Body weight dimorphism in anthropoid primates has been thought to be a consequence of sexual selection resulting from male-male competition for access to mates. However, while monogamous anthropoids show low degrees of weight dimorphism, as predicted by the sexual selection hypothesis, polygynous anthropoids show high variation in weight dimorphism that is not associated with measures of mating system or sex ratio. This observation has led many to debate the role of other factors such as dietary constraints, predation pressure, substrate constraints, allometric effects, and phylogeny in the evolution of anthropoid weight dimorphism. Here, we re-evaluate variation in adult body weight dimorphism in anthropoids, testing the sexual selection hypothesis using categorical estimates of the degree of male-male intrasexual competition (“competition levels”). We also test the hypotheses that interspecific variation in body weight dimorphism is associated with female body weight and categorical estimates of diet, substrate use, and phylogeny. Weight dimorphism is strongly associated with competition levels, corroborating the sexual selection hypothesis. Weight dimorphism is positively correlated with increasing female body weight, but evidence suggests that the correlation reflects an interaction between overall size and behavior. Arboreal species are, on average, less dimorphic than terrestrial species, while more frugivorous species tend to be more dimorphic than folivorous or insectivorous species. Several alternative hypotheses can explain these latter results. Weight dimorphism is correlated with taxonomy, but so too are competition levels. We suggest that most taxonomic correlations of weight dimorphism represent “phylogenetic niche conservatism”; however, colobines show consistently low degrees of weight dimorphism for reasons that are not clear. Am J Phys Anthropol 103:37–68, 1997. © 1997 Wiley-Liss, Inc.     

Hemiacodon engardae, a new species of omomyid primate from the earliest Uintan Turtle Bluff Member of the Bridger Formation, southwestern Wyoming, USA

In this paper, we describe a new species of Hemiacodon known only from University of Colorado Museum Loc. 92189 (Donna's Locality) in the Turtle Bluff Member of the Bridger Formation, Green River Basin, southwestern Wyoming. Donna's locality has yielded a diverse mostly small-bodied mammalian assemblage of Bridgerian and first appearance Uintan mammalian taxa, as well as range-through taxa. Together with H. engardae sp. nov., the faunal assemblage from Donna's Locality and more recently discovered localities in the same stratigraphic interval provides the first conclusive paleontological evidence of an earliest Uintan age (Ui1A biochron) for the Turtle Bluff Member of the Bridger Formation.The new species is represented by a sample of 11 specimens consisting of well-preserved upper and lower premolars and lower molars. H. engardae is distinct from H. gracilis on the basis of overall larger size as well as a combination of features of the premolars and molars related to a greater development of shearing crests. This suggests that H. engardae may have incorporated more foliage into its diet than the Bridgerian species, H. gracilis.     

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.     

Paleobiology of the oligopithecines,the earliest known anthropoid primates

Anthropoid primates of the subfamily Oligopithecinae are late Eocene in age, and have a known distribution of Northeast Africa and the Arabian Peninsula. Body sizes of the three known oligopithecine species are estimated from allometric molar size regressions to be 700–1000 g forOligopithecus savagei, 600–900 g forCatopithecus browni, and 500 g for the least well-known and smallest species,Proteopithecus sylviae. Occlusal features of the molar teeth, considered in conjunction with body size, suggest that all three species were frugivorous and insectivorous. The orbital size ofCatopithecus indicates a diurnal activity cycle. A relatively broad interobital region in this species may indicate prosimian-like or callitrichid-like olfactory adaptations. Structural features of the crushed skull suggest thatCatopithecus had a smaller cranial capacity than those of extant anthropoids with a similar body size. Fossil plants and birds from localities yielding oligopithecines suggest a wet, warm, tropical, forested, swampy environment. These paleobiological inferences about the extinct oligopithecines are discussed in relation to questions about primate adaptations near the prosimian-anthropoid transition.