No abstract

Fossils and molecular data are two independent sources of information that should in principle provide consistent inferences of when evolutionary lineages diverged. Here we use an alternative approach to genetic inference of species split times in recent human and ape evolution that is independent of the fossil record. We first use genetic parentage information on a large number of wild chimpanzees and mountain gorillas to directly infer their average generation times. We then compare these generation time estimates with those of humans and apply recent estimates of the human mutation rate per generation to derive estimates of split times of great apes and humans that are independent of fossil calibration. We date the human-chimpanzee split to at least 7-8 million years and the population split between Neanderthals and modern humans to 400,000-800,000 y ago. This suggests that molecular divergence dates may not be in conflict with the attribution of 6-to 7-million-y-old fossils to the human lineage and 400,000-yold fossils to the Neanderthal lineage.hominin | molecular dating | primate | speciation O ver 40 y ago, Sarich and Wilson used immunological data to propose that humans and African great apes diverged only about 5 million y ago, some three to four times more recently than had been assumed on the basis of the fossil record (1). Although contentious at the time (e.g., ref. 2), this divergence has since been repeatedly estimated from DNA sequence data at 4-6 million years ago (Ma) (3-8). However, this estimate is incompatible with the attribution of fossils older than 6 Ma to the human lineage. Although the assignment of fossils such as the ∼6 Ma Orrorin (9) and the 6-7 Ma Sahelanthropus (10) to the human lineage remains controversial (11), it is also possible that the divergence dates inferred from DNA sequence data are too recent.The total amount of sequence differences observed today between two evolutionary lineages can be expressed as the sum of two values: the sequence differences that accumulated since gene flow ceased between the lineages ("split time") and the sequence differences that correspond to the diversity in the common ancestor of both lineages. The extent of variation in the ancestral species may be estimated from the variance of DNA sequence differences observed across different parts of the genome between the species today, which will be larger the greater the level of variation in the ancestral population. By subtracting this value from the total amount of sequence differences, the sequence differences accumulated since the split can be estimated. The rate at which DNA sequence differences accumulate in the genome ("mutation rate") is needed to then convert DNA sequence differences into split times.In prior research, mutation rates have been calculated using species split times estimated from the fossil record as calibration points. For calculating split times between present-day humans and great apes, calibration points that assume DNA sequence differences between humans and orangutans...

Genetic analysis using noninvasively collected samples such as faeces continues to pose a formidable challenge because of unpredictable variation in the extent to which usable DNA is obtained. We investigated the influence of multiple variables on the quantity of DNA extracted from faecal samples from wild mountain gorillas and chimpanzees. There was a small negative correlation between temperature at time of collection and the amount of DNA obtained. Storage of samples either in RNAlater solution or dried using silica gel beads produced similar results, but significantly higher amounts of DNA were obtained using a novel protocol that combines a short period of storage in ethanol with subsequent desiccation using silica.

The most important environmental factor explaining interspecies variation in ecology and sociality of the great apes is likely to be variation in resource availability. Relatively little is known about the activity patterns of western lowland gorillas (Gorilla gorilla gorilla), which inhabit a dramatically different environment from the well-studied mountain gorillas (G. beringei beringei). This study aims to provide a detailed quantification of western lowland gorillas' activity budgets using direct observations on one habituated group in Bai Hokou, Central African Republic. We examined how activity patterns of both sexes are shaped by seasonal frugivory. Activity was recorded with 5-min instantaneous sampling between December 2004 and December 2005. During the high-frugivory period the gorillas spent less time feeding and more time traveling than during the low-frugivory period. The silverback spent less time feeding but more time resting than both females and immatures, which likely results from a combination of social and physiological factors. When compared with mountain gorillas, western lowland gorillas spend more time feeding (67 vs. 55%) and traveling (12 vs. 6.5%), but less time resting (21 vs. 34%) and engaging in social/other activities (0.5 vs. 3.6%). This disparity in activity budgets of western lowland gorillas and mountain gorillas may be explained by the more frugivorous diet and the greater dispersion of food resources experienced by western lowland gorillas. Like other apes, western lowland gorillas change their activity patterns in response to changes in the diet.

Twenty-five years of demographic data on free-ranging mountain gorillas (Gorilla gorilla beringei) from the Karisoke Research Center, Rwanda were used to assess male life histories and the variation within the age-graded social structure. Group types include lone silver-backs, one-male, multimale, and all-male groups. Although 60% of gorilla groups in the Virunga population are one-male, a multimale structure may increase chances of survival and future reproductive success for males at three stages of their lives. Infants born in one-male groups appear more likely to die from infanticide than those in multimale groups. Immature males in one-male groups may face decreased future reproductive opportunities compared to males in multimale groups. Adult males in one-male groups lack possible partners for coalition formation during intergroup encounters. Demographic constraints, such as length of time to male maturation, coupled with intense male-male competition for mates may limit the number and duration of groups with a multimale structure. Individuals are not restricted to one group type for their entire adult lives and males that attain maturity in each group type may eventually reproduce. Variation in male reproductive success is based both on length of reproductive tenure and on the number of mates.