The history and composition of the Raymond A. Dart Collection of Human Skeletons at the University of the Witwatersrand,Johannesburg, South Africa

The Raymond A. Dart Collection of Human Skeletons (Dart Collection) is housed in the School of Anatomical Sciences at the University of the Witwatersrand, Johannesburg, South Africa, and comprises one of the largest documented cadaver‐derived human skeletal assemblages in the world. This collection originated in the early 1920s as a result of the efforts of Raymond Dart and continues to grow. The skeletons included represent varied indigenous and immigrant populations from southern Africa, Europe and Asia. This contribution documents the history of the collection and provides an updated inventory and demographic assessment of this valuable research collection. According to a recent inventory the Dart Collection currently comprises 2,605 skeletons representing individuals from regional SA African (76%), White (15%), Coloured (4%) and Indian (0.3%) populations. A large proportion of the skeletons (71%) represent males. The recorded ages at death range from the first year to over 100 years of age, but the majority of individuals died between the ages of 20 and 70. The Dart Collection has been affected by collection procedures based on availability. All of the cadavers collected before 1958, and large proportions subsequently, were derived from unclaimed bodies in regional South African hospitals. Some details of documentation (age at death, population group) are estimates and some aspects of the collection demographics (sex ratios) do not closely reflect any living South African population. Our inventory and analysis of the Dart Collection is aimed to assist researchers planning research on the materials from this collection. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Fetal ultrasonography: biometric data from four African primate species

BACKGROUND: Nonhuman primates are raised in large numbers in research centers and zoos. Reproductive monitoring is required to improve breeding performances. Ultrasonography is a safe method to determine gestational age and to estimate the date of parturition. However only few data are available in nonhuman primates. METHODS: Fetal biometric data were obtained throughout pregnancy on four African primate species, namely chimpanzee, gorilla, mandrill and patas monkey. Measurements included biparietal diameter, transverse abdominal diameter, femur and humerus length, external interorbital diameter, and fetal heart rate. Curves established from these data were compared with previously published data in chimpanzees and gorillas and with those for humans and other closely related primate species. RESULTS: The curves for the different hominids were very similar, while those for mandrills more closely resembled baboons and data for patas monkeys were comparable to those for macaques. CONCLUSIONS: These data, by providing a tool to evaluate precise gestational age, will be useful for centers raising these four primate species.     

Human Ability to Recognize Kin Visually Within Primates

The assessment of relatedness is a key determinant in the evolution of social behavior in primates. Humans are able to detect kin visually in their own species using facial phenotypes, and facial resemblance in turn influences both prosocial behaviors and mating decisions. This suggests that cognitive abilities that allow facial kin detection in conspecifics have been favored in the species by kin selection. We investigated the extent to which humans are able to recognize kin visually by asking human judges to assess facial resemblance in 4 other primate species (common chimpanzees, western lowland gorillas, mandrills, and chacma baboons) on the basis of pictures of faces. Humans achieved facial interspecific kin recognition in all species except baboons. Facial resemblance is a reliable indicator of relatedness in at least chimpanzees, gorillas, and mandrills, and future work should explore if the primates themselves also share the ability to detect kin facially.     

Trade-offs in skillacquisition and time allocation among juvenile chacma baboons

We hypothesize that juvenile baboons are less efficient foragers than adult baboons owing to their small size, lower level of knowledge and skill, and/or lesser ability to maintain access to resources. We predict that as resources are more difficult to extract, juvenile baboons will demonstrate lower efficiency than adults will because of their lower levels of experience. In addition, we hypothesize that juvenile baboons will be more likely to allocate foraging time to easier-to-extract resources owing to their greater efficiency in acquiring those resources. We use feeding efficiency and time allocation data collected on a wild, free-ranging, non-provisioned population of chacma baboons (Papio hamadryas ursinus) in the Moremi Wildlife Reserve, Okavango Delta, Botswana to test these hypotheses. The major findings of this study are: 1. Juvenile baboons are significantly less efficient foragers than adult baboons primarily for difficult-to-extract resources. We propose that this age-dependent variation in efficiency is due to differences in memory and other cognitive functions related to locating food resources, as is indicated by the greater amount of time juvenile baboons spend searching for food. There is no evidence that smaller body size or competitive disruption influences the differences in return rates found between adult and juvenile baboons in this study. 2. An individual baboon’s feeding efficiency for a given resource can be used to predict the duration of its foraging bouts for that resource. These results contribute both to our understanding of the ontogeny of behavioral development in nonhuman primates, especially regarding foraging ability, and to current debate within the field of human behavioral ecology regarding the evolution of the juvenile period in primates and humans. Sara E. Johnson is Assistant Professor of Anthropology at California State University, Fullerton. She received her Ph.D. in Anthropology (Human Evolutionary Ecology) from the University of New Mexico in 2001. She uses behavioral ecology and life history theory to address her research interests in the evolution of primate and human growth; ecological variation and phenotypic plasticity in growth and development; ecological variation in life course trajectories, including fertility, health, morbidity, and mortality differentials; food acquisition and production related to nutrition; societal transofmration and roles of the elderly among indigenous peoples; and women’s reproductive and productive roles in both traditional and nontraditional societies. For the past decade she has conducted research on these issues in several different populations, including chacma baboons in the Okavango Delta of Botswana, two multiethnic communities of forager/agropastoralists in the Okavango Delta of Botswana, and among New Mexican men. John Bock is Associate Professor of Anthropology at California State University at Fullerton and is Associate Editor of Human Nature. He received a Ph.D. in Anthropology (Human Evolutionary EcologY) from the University of New Mexico in 1995, and from 1995 to 1998 was an Andrew W. Mellon Foundation postdoctoral fellow in demography and epidemiology at the National Centre for Epidemiology and Population Health at Australian National University. His recent research has focused on applying life history theory to understanding the evolution of the primate and human juvenile period. Bock has been conducting research among the Okavango Delta peoples of Botswana since 1992, and his current research there is an examination of child development and family demography in relation to socioecology and the HIV/AIDS epidemic. Other research is focused on health disparties among minorities and indigenous peoples in Botswana and the United States related to differential access to health care.     

The subspecies concept in primatology: The case of mountain gorillas

The criteria for the application of subspecific units in living primate populations have received little attention relative to other vertebrate taxa, even though they have important implications for conservation strategies for many nonhuman primate populations. One of the most critically endangered primates is the mountain gorilla,Gorilla gorilla beringei, of which 600 animals exist in east-central Africa. FollowingSarmiento et al. (1996), taxonomists have proposed splitting these populations into two subspecies as part of a revised taxonomy of the genusGorilla. In this paper I review the application of the subspecies concept in primatology, using the gorillas of Bwindi Impenetrable National Park and the Virungas as case studies. An examination of genetic, morphological, biogeographic, ecological, and behavioral evidence indicates that reclassifying Bwindi gorillas as taxonomically distinct from those in the Virungas is not well supported and needs further study. Because taxonomy provides the basis of conservation management policies, a cautious and conservative approach to the subspecies question is warranted in the case of endangered primate populations.     

Estrogenic plant foods of red colobus monkeys and mountain gorillas in Uganda

Phytoestrogens, or naturally occurring estrogen-mimicking compounds, are found in many human plant foods, such as soybeans (Glycine max) and other legumes. Because the consumption of phytoestrogens may result in both health benefits of protecting against estrogen-dependent cancers and reproductive costs of disrupting the developing endocrine system, considerable biomedical research has been focused on the physiological and behavioral effects of these compounds. Despite this interest, little is known about the occurrence of phytoestrogens in the diets of wild primates, nor their likely evolutionary importance. We investigated the prevalence of estrogenic plant foods in the diets of two folivorous primate species, the red colobus monkey (Procolobus rufomitratus) of Kibale National Park and mountain gorilla (Gorilla beringei) of Bwindi Impenetrable National Park, both in Uganda. To examine plant foods for estrogenic activity, we screened 44 plant items (species and part) comprising 78.4% of the diet of red colobus monkeys and 53 plant items comprising 85.2% of the diet of mountain gorillas using transient transfection assays. At least 10.6% of the red colobus diet and 8.8% of the gorilla diet had estrogenic activity. This was mainly the result of the red colobus eating three estrogenic staple foods and the gorillas eating one estrogenic staple food. All estrogenic plants exhibited estrogen receptor (ER) subtype selectivity, as their phytoestrogens activated ERβ, but not ERα. These results demonstrate that estrogenic plant foods are routinely consumed by two folivorous primate species. Phytoestrogens in the wild plant foods of these two species and many other wild primates may have important implications for understanding primate reproductive ecology.     

The baboon as a non-human primate model of human schistosome infection

Over the past three decades, intensive studies of murine schistosomiasis have provided important clues to the understanding of the human disease, but growing evidence suggests that these results derived from highly inbred strains of mice might not have direct applicability to the human infection. Recent data based on the baboon indicate that infection in this non-human primate might mirror the human situation. In this review, Mramba Nyindo and Idle Farah demonstrate that baboons provide an excellent non-human primate model that produces pathology and disease closely resembling that observed in humans, and address how studies in baboons can provide insights into mechanisms regulating schistosomiasis mansoni pathology and immunity. They also address, in a general way, issues related to the use of non-human primates in biomedical research.     

Temporal variation selects for diet–microbe co-metabolic traits in the gut of Gorilla spp

Although the critical role that our gastrointestinal microbes play in host physiology is now well established, we know little about the factors that influenced the evolution of primate gut microbiomes. To further understand current gut microbiome configurations and diet–microbe co-metabolic fingerprints in primates, from an evolutionary perspective, we characterized fecal bacterial communities and metabolomic profiles in 228 fecal samples of lowland and mountain gorillas (G. g. gorilla and G. b. beringei, respectively), our closest evolutionary relatives after chimpanzees. Our results demonstrate that the gut microbiomes and metabolomes of these two species exhibit significantly different patterns. This is supported by increased abundance of metabolites and bacterial taxa associated with fiber metabolism in mountain gorillas, and enrichment of markers associated with simple sugar, lipid and sterol turnover in the lowland species. However, longitudinal sampling shows that both species'' microbiomes and metabolomes converge when hosts face similar dietary constraints, associated with low fruit availability in their habitats. By showing differences and convergence of diet–microbe co-metabolic fingerprints in two geographically isolated primate species, under specific dietary stimuli, we suggest that dietary constraints triggered during their adaptive radiation were potential factors behind the species-specific microbiome patterns observed in primates today.     

Cortical surface patterns in human and nonhuman primates

An analysis of skulls from several primate species shows that a “worm-track” surface pattern, first identified in the brow region in fossil adult hominids and subsequently in olive baboons, chimpanzees, and macaques, is also present in numerous other species. Fine cancellous bone and its attendant vermiculate surface pattern have been observed in subadult and adult gelada baboons, gibbons, gorillas, and orangutans as well as in modern Homo sapiens and several Plio-Pleistocene fossil hominids. In contemporary primates, fine cancellous bone has been identified not only in the brow region, but also along the zygomatic arch, on the pterygoid plates, on the maxilla, along the temporal line, on the mastoid process, and in the region of inion. Given the widespread distribution of this trait, caution is advised when using it as a diagnostic indicator of the evolutionary or functional significance of craniofacial morphology.     

Evolution of nuclear gene families in primates,Copy-number variation in the argininosuccinate synthetase (ASS) pseudogene family and the anonymous DNA sequence,D1S1

Changes in the copy number of nuclear genes provide the raw material for the creation of new gene functions. To better understand the mechanisms for such events, and their physiologic and evolutionary consequences, it is valuable to study a well characterized and closely related group of species such as primates. Fortuitously, most of the powerful molecular techniques and DNA probes developed for research in humans are equally applicable to non-human primates. We review what is known of copy number variation in primates and describe two informative DNA probes: pAS-1, a cDNA probe to the human urea cycle enzyme argininosuccinate synthetase (ASS), and an anonymous DNA probe, D1S1.In additon to the ASS structural locus on human chromosome 9, pAS-1 detects at least 14 dispersed, processed pseudogenes in humans. The number of pseudogene copies appears to be approximately the same in humans, chimpanzees, gorillas, orangutans and baboons; less in marmosets; and least in some rodents. Chimpanzees and gorillas appear to have all of the human pseudogenes though an Xp copy may be missing from gorillas. The Y pseudogene is apparently absent from orangutans and baboons, and, finally, a comparison of humans and chimpanzees revealed that the number of nucleotide substitutions in the Y chromosome pseudogenes is approximately 1 per 100.D1S1 maps to human chromosome 3 but also detects a high homology copy on chromosome 1. Chimpanzees, gorillas and orangutans all appear to have only the chromosome 3 homolog suggesting that this is the ancestral sequence and that the duplication occurred after separation of humans and the great apes.Both the ASS pseudogene family and the D1S1 system provide valuable information on the evolution of nuclear gene families in primates.     

A comparative study of the trabecular bony architecture of the talus in humans, non-human primates, and Australopithecus

This study tested the hypothesis that talar trabecular microarchitecture reflects the loading patterns in the primate ankle joint, to determine whether talar trabecular morphology might be useful for inferring locomotor behavior in fossil hominins. Trabecular microarchitecture was quantified in the anteromedial, anterolateral, posteromedial, and posterolateral quadrants of the talar body in humans and non-human primates using micro-computed tomography. Trabecular bone parameters, including bone volume fraction, trabecular number and thickness, and degree of anisotropy differed between primates, but not in a manner entirely consistent with hypotheses derived from locomotor kinematics. Humans have highly organized trabecular struts across the entirety of the talus, consistent with the compressive loads incurred during bipedal walking. Chimpanzees possess a high bone volume fraction, consisting of plate-like trabecular struts. Orangutan tali are filled with a high number of thin, connected trabeculae, particularly in the anterior portion of the talus. Gorillas and baboons have strikingly similar internal architecture of the talus. Intraspecific analyses revealed no regional differences in trabecular architecture unique to bipedal humans. Of the 22 statistically significant regional differences in the human talus, all can also be found in other primates. Trabecular thickness, number, spacing, and connectivity density had the same regional relationship in the talus of humans, chimpanzees, gorillas, and baboons, suggesting a deeply conserved architecture in the primate talus. Australopithecus tali are human-like in most respects, differing most notably in having more oriented struts in the posteromedial quadrant of the body compared with the posterolateral quadrant. Though this result could mean that australopiths loaded their ankles in a unique manner during bipedal gait, the regional variation in degree of anisotropy was similar in humans, chimpanzees, and gorillas. These results collectively suggest that the microarchitecture of the talus does not simply reflect the loading environment, limiting its utility in reconstructing locomotion in fossil primates.     

Antigenic similarities to HCG subunits among chorionic gonadotropins of nonhuman primates.

Comparison of antigenic similarity between human chorionic gonadotropin (HCG) subunits and the chorionic gonadotropins of six species of nonhuman primates indicates marked similarity of antigenic determinants between both subunits of HCG and the chorionic gonadotropins of chimpanzees, gorillas, and orangutans. Antisera to HCG subunits (alpha or beta) did not cross-react with the chorionic gonadotropins of baboons, macaques, or marmosets. Because of the relative availability of chimpanzees for laboratory studies, we suggest that chimpanzees may be the optimal nonhuman primate model for determining the advisability of vaccinations in man using conjugates of HCG fragments to achieve fertility control or for suppression of HCG-producing neoplasms.     

Nonhuman primate infections after organ transplantation

Nonhuman primates, primarily rhesus macaques (Macaca mulatta), cynomolgus macaques (Macaca fascicularis), and baboons (Papio spp.), have been used extensively in research models of solid organ transplantation, mainly because the nonhuman primate (NHP) immune system closely resembles that of the human. Nonhuman primates are also frequently the model of choice for preclinical testing of new immunosuppressive strategies. But the management of post-transplant nonhuman primates is complex, because it often involves multiple immunosuppressive agents, many of which are new and have unknown effects. Additionally, the resulting immunosuppression carries a risk of infectious complications, which are challenging to diagnose. Last, because of the natural tendency of animals to hide signs of weakness, infectious complications may not be obvious until the animal becomes severely ill. For these reasons the diagnosis of infectious complications is difficult among post-transplant NHPs. Because most nonhuman primate studies in organ transplantation are quite small, there are only a few published reports concerning infections after transplantation in nonhuman primates. Based on our survey of these reports, the incidence of infection in NHP transplant models is 14%. The majority of reports suggest that many of these infections are due to reactivation of viruses endemic to the primate species, such as cytomegalovirus (CMV), polyomavirus, and Epstein-Barr virus (EBV)-related infections. In this review, we address the epidemiology, pathogenesis, role of prophylaxis, clinical presentation, and treatment of infectious complications after solid organ transplantation in nonhuman primates.     

Advantages and limitations of nonhuman primates as animal models in genetic research on complex diseases

Abstract: The genetic similarity between humans and nonhuman primates makes nonhuman primates uniquely suited as models for genetic research on complex physiological and behavioral phenotypes. By comparison with human subjects, nonhuman primates, like other animal models, have several advantages for these types of studies: 1) constant environmental conditions can be maintained over long periods of time, greatly increasing the power to detect genetic effects; 2) different environmental conditions can be imposed sequentially on individuals to characterize genotype-environment interactions; 3) complex pedigrees that are much more powerful for genetic analysis than typically available human pedigrees can be generated; 4) genetic hypotheses can be tested prospectively by selective matings; and 5) essential invasive and terminal experiments can be conducted. Limitations of genetic research with nonhuman primates include cost and availability. However, the ability to manipulate both genetic and environmental factors in captive primate populations indicates the promise of genetic research with these important animal models for illuminating complex disease processes. The utility of nonhuman primates for biomedical research on human health problems is illustrated by examples concerning the use of baboons in studies of osteoporosis, alcohol metabolism, and lipoproteins.     

Major histocompatibility complex and microsatellite variation in two populations of wild gorillas

In comparison to their close relatives the chimpanzees and humans, very little is known concerning the amount and structure of genetic variation in gorillas. Two species of gorillas are recognized and while the western gorillas number in the tens of thousands, only several hundred representatives of the mountain gorilla subspecies of eastern gorillas survive. To analyse the possible effects of these different population sizes, this study compares the variation observed at microsatellite and major histocompatibility complex (MHC) loci in samples of wild western and mountain gorillas, collected using a sampling scheme that targeted multiple social groups within defined geographical areas. Noninvasive samples proved a viable source of DNA for sequence analysis of the second exon of the DRB loci of the MHC. Observed levels of variation at the MHC locus were similar between the two gorilla species and were comparable to those in other primates. Comparison of results from analysis of variation at multiple microsatellite loci found only a slight reduction in heterozygosity for the mountain gorillas despite the relatively smaller population size.     

Blood,sweat and plaster casts: Reviewing the history,composition, and scientific value of the Raymond A. Dart Collection of African Life and Death Masks

This paper addresses the history, composition and scientific value of one of the most comprehensive facemask collections in Africa, the Raymond A. Dart Collection of African Life and Death Masks. Housed within the School of Anatomical Sciences at the University of the Witwatersrand (South Africa), it comprises 1110 masks (397 life, 487 death, 226 unknown). Life masks represent populations throughout Africa; death masks predominately southern Africa. Males preponderate by 75%. Recorded ages are error prone, but suggest most life masks are those of <35 year-olds, death masks of 36+ year-olds. A total of 241 masks have associated skeletons, 209 presenting a complete skull.Life masks date between 1927 and c.1980s, death masks 1933 and 1963. This historical collection presents uncanny associations with outmoded typological and evolutionary theories. Once perceived an essential scientific resource, performed craniofacial superimpositions identify the nose as the only stable feature maintained, with the remaining face best preserved in young individuals with minimal body fat. The facemask collection is most viable for teaching and research within the history of science, specifically physical anthropology, and presents some value to craniofacial identification. Future research will have to be conducted with appropriate ethical considerations to science and medicine.     

Social structure and life-history patterns in western gorillas (Gorilla gorilla gorilla)

Life-history traits and ecological conditions have an important influence on primate social systems. Most of what we know about the life-history patterns and social structure of gorillas comes from studies of eastern gorillas (Gorilla beringei sp.), which live under dramatically different ecological conditions compared to western gorillas (Gorilla gorilla sp.). In this paper we present new data on western gorilla social structure and life histories from four study sites, and make comparisons with eastern gorilla populations. Data were obtained from two study sites with gorilla groups undergoing the habituation process (Lossi, Democratic Republic of Congo and Bai Hokou, Central African Republic) and two "bai" studies (Maya Nord and Mbeli Bai, Republic of Congo). The size and structure of these groups were similar to those seen in eastern gorillas. However, differences in the occurrence of various group transitions (group formations, changes between one-male and multimale composition, and group disintegrations) exist, and western gorillas notably exhibit much higher rates of male emigration and correspondingly fewer multimale groups compared to mountain gorillas. Certain phenomena have been observed only rarely, including predation by leopards. The preliminary data show no significant differences in birth rates between western gorillas and mountain gorillas. The ecological variability across gorilla habitats likely explains the flexibility in the social system of gorillas, but we need more information on the social relationships and ecology of western gorillas to elucidate the causes for the similarities and differences between western and eastern gorillas on the levels of individuals, social groups, and population dynamics.     

Geometry of nutrition in field studies: an illustration using wild primates

Nutritional geometry has shown the benefits of viewing nutrition in a multidimensional context, in which foraging is viewed as a process of balancing the intake and use of multiple nutrients. New insights into nutrient regulation have been generated in studies performed in a laboratory context, where accurate measures of amounts (e.g. eaten, converted to body mass, excreted) can be made and analysed using amounts-based nutritional geometry. In most field situations, however, proportional compositions (e.g. of foods, diets, faeces) are the only measures readily available, and in some cases are more relevant to the problem at hand. For this reason, a complementary geometric method was recently introduced for analysing multi-dimensional data on proportional compositions in nutritional studies, called the right-angled mixture triangle (RMT). We use literature data from field studies of primates to demonstrate how the RMT can provide insight into a variety of important concepts in nutritional ecology. We first compare the compositions of foods, using as an example primate milks collected in both the wild and the laboratory. We next compare the diets of different species of primates from the same habitat and of the same species (mountain gorillas) from two distinct forests. Subsequently, we model the relationships between the composition of gorilla diets in these two habitats and the foods that comprise these diets, showing how such analyses can provide evidence for active nutrient-specific regulation in a field context. We provide a framework to relate concepts developed in laboratory studies with field-based studies of nutrition.     

Brief communication: Hand preference for bimanual and unimanual feeding in captive gorillas: extension in a second colony of apes

Right-hand dominance is widely considered to be a uniquely human trait. Whether nonhuman primates exhibit similar population-level hand preferences remains a topic of considerable debate. Despite extensive research focusing on laterality in nonhuman primates, our interpretation of these studies is limited due to methodological issues including the lack of a common measure of hand preference and the use of tasks that may not be reliable indicators of handedness. The use of consistent methods between studies is necessary to enable comparisons within and between species and allow for more general conclusions to be drawn from these results. The present study replicates methods used in recent research reporting population-level right-handedness in captive gorillas (Meguerditchian et al.,2010). Observational data were collected on hand preference for unimanual and bimanual feeding in 14 captive western lowland gorillas (Gorilla gorilla gorilla). Individual-level preferences were found, primarily for bimanual feeding; however, the data reveal no group-level directional bias (contra Meguerditchian et al.). Like the study by Meguerditchian et al. (2010), though, bimanual feeding revealed significantly stronger hand preferences than unimanual reaching, and age, sex, group membership, or rearing history had no effect on hand preference. Finally, variations in diet and corresponding grip type between studies suggest that hand preferences may vary across bimanual tasks depending on grip morphology. This study aims to contribute to our existing knowledge of primate laterality by increasing the number of individuals investigated using methods that allow for comparisons with similar research.     

Failure to detect simian immunodeficiency virus infection in a large Cameroonian cohort with high non-human primate exposure

Hunting and butchering of wildlife in Central Africa are known risk factors for a variety of human diseases, including HIV/AIDS. Due to the high incidence of human exposure to body fluids of non-human primates, the significant prevalence of simian immunodeficiency virus (SIV) in non-human primates, and hunting/butchering associated cross-species transmission of other retroviruses in Central Africa, it is possible that SIV is actively transmitted to humans from primate species other than mangabeys, chimpanzees, and/or gorillas. We evaluated SIV transmission to humans by screening 2,436 individuals that hunt and butcher non-human primates, a population in which simian foamy virus and simian T-lymphotropic virus were previously detected. We identified 23 individuals with high seroreactivity to SIV. Nucleic acid sequences of SIV genes could not be detected, suggesting that SIV infection in humans could occur at a lower frequency than infections with other retroviruses, including simian foamy virus and simian T-lymphotropic virus. Additional studies on human populations at risk for non-human primate zoonosis are necessary to determine whether these results are due to viral/host characteristics or are indicative of low SIV prevalence in primate species consumed as bushmeat as compared to other retroviruses in Cameroon.