A missing piece of the Papio puzzle: Gorongosa baboon phenostructure and intrageneric relationships.

Most authors recognize six baboon species: hamadryas (Papio hamadryas), Guinea (Papio papio), olive (Papio anubis), yellow (Papio cynocephalus), chacma (Papio ursinus), and Kinda (Papio kindae). However, there is still debate regarding the taxonomic status, phylogenetic relationships, and the amount of gene flow occurring between species. Here, we present ongoing research on baboon morphological diversity in Gorongosa National Park (GNP), located in central Mozambique, south of the Zambezi River, at the southern end of the East African Rift System. The park exhibits outstanding ecological diversity and hosts more than 200 baboon troops. Gorongosa National Park baboons have previously been classified as chacma baboons (P. ursinus). In accordance with this, two mtDNA samples from the park have been placed in the same mtDNA clade as the northern chacma baboons. However, GNP baboons exhibit morphological features common in yellow baboons (e.g., yellow fur color), suggesting that parapatric gene flow between chacma and yellow baboons might have occurred in the past or could be ongoing. We investigated the phenostructure of the Gorongosa baboons using two approaches: 1) description of external phenotypic features, such as coloration and body size, and 2) 3D geometric morphometric analysis of 43 craniofacial landmarks on 11 specimens from Gorongosa compared to a pan-African sample of 352 baboons. The results show that Gorongosa baboons exhibit a mosaic of features shared with southern P. cynocephalus and P. ursinus griseipes. The GNP baboon phenotype fits within a geographic clinal pattern of replacing allotaxa. We put forward the hypothesis of either past and/or ongoing hybridization between the gray-footed chacma and southern yellow baboons in Gorongosa or an isolation-by-distance scenario in which the GNP baboons are geographically and morphologically intermediate. These two scenarios are not mutually exclusive. We highlight the potential of baboons as a useful model to understand speciation and hybridization in early human evolution.

Ontogenetic changes in limb postures and their impact on effective limb length in baboons (Papio cynocephalus).

OBJECTIVES: Digitigrade hand and foot postures and extended elbows and knees are considered adaptations to running in cursorial mammals because they increase effective limb lengths (ELLs). However, the relationship between digitigrady and ELL in primates is not well understood. We documented the ontogeny of limb postures in baboons to better understand the function of digitigrady during walking. We hypothesized that the hand and foot would become more elevated and the elbow and knee more extended, leading to increased relative ELLs throughout ontogeny. MATERIALS AND METHODS: Longitudinal kinematic data were collected on four infant yellow baboons (Papio cynocephalus) as they aged from two to nine months, and again at two to three years. Hand/foot postures, elbow/knee angles, relative fore/hind limb ELLs, and dimensionless velocity were measured for 404 symmetrical walking strides. RESULTS: Digitigrade hand and foot postures were preferred at all ages. The elbow extended slightly and the knee flexed slightly with age. Elevated proximal hands, extended elbows, and extended knees were associated with long relative ELLs. For a given age, relative hind limb ELL was longer than relative forelimb ELL. DISCUSSION: In the forelimb, digitigrade hand postures and extended elbows function to increase relative ELL at slow walking velocity. Increased forelimb ELL may be an attempt to equalize forelimb and hind limb ELLs in baboons with an absolutely longer hind limb. Pedal digitigrady is not a main contributing factor to hind limb ELL. Results suggest that manual and pedal digitigrady in terrestrial cercopithecoids does not function to increase velocity.

Sexual Dimorphism of Endocranial, Facial and Limb Measurements in the Yellow Baboon (Papio cynocephalus).

The present study focused on the sexual dimorphism of yellow baboon (Papio cynocephalus) to clarify its relationship with social behaviour. The degree of sexual dimorphism in the endocranial volume is the lowest among the investigated measurements. Among the facial measurements, the degree value of sexual dimorphism was the maximum (38.4%) for palate length and the minimum for palate breadth at the upper second molar (M2) (16.8%). Reduced major axis (RMA) regression analysis indicated that most positive allometry in relation to body mass was barely shown in the endocranial volume, palate length, palate breadth at M2 and mandibular ramal width. On the other hand, most negative allometry in relation to body mass was barely indicated in the bizygomatic breadth, skull length, humeral length and femoral length. The plate breadth at M2 in males was smaller than that in females in equivalent to body mass. The results of the present study suggest that more males have longer pointed muzzles than females, which is considered to create an impressive view of large canine teeth. This contributes to display among males and agonistic encounters rather than to necessity of increased facial size due to larger body size or dietary influences.

Tooth size variation related to age in Amboseli baboons.

We measured the molar size from a single population of wild baboons from Amboseli (Kenya), both females (n=57) and males (n=50). All the females were of known age; the males represented a mix of known-age individuals (n=31) and individuals with ages estimated to within 2 years (n=19). The results showed a significant reduction in the mesiodistal length of teeth in both sexes as a function of age. Overall patterns of age-related change in tooth size did not change whether we included or excluded the individuals of estimated age, but patterns of statistical significance changed as a result of changed sample sizes. Our results demonstrate that tooth length is directly related to age due to interproximal wearing caused by M2 and M3 compression loads. Dental studies in primates, including both fossil and extant species, are mostly based on specimens obtained from osteological collections of varying origins, for which the age at death of each individual in the sample is not known. Researchers should take into account the phenomenon of interproximal attrition leading to reduced tooth size when measuring tooth length for ondontometric purposes.

Nonhuman anthropoid primate femoral neck trabecular architecture and its relationship to locomotor mode.

Functional analyses of human and nonhuman anthropoid primate femoral neck structure have largely ignored the trabecular bone. We tested hypotheses regarding differences in the relative distribution and structural anisotropy of trabecular bone in the femoral neck of quadrupedal and climbing/suspensory anthropoids. We used high-resolution X-ray computed tomography to analyze quantitatively the femoral neck trabecular structure of Ateles geoffroyi, Symphalangus syndactylus, Alouatta seniculus, Colobus guereza, Macaca fascicularis, and Papio cynocephalus (n = 46). We analyzed a size-scaled superior and inferior volume of interest (VOI) in the femoral neck. The ratio of the superior to inferior VOI bone volume fraction indicated that the distribution of trabecular bone was inferiorly skewed in most (but not all) quadrupeds and evenly distributed the climbing/suspensory species, but interspecific comparisons indicated that all taxa overlapped in these measurements. Degree of anisotropy values were generally higher in the inferior VOI of all species and the results for the two climbing/suspensory taxa, A. geoffroyi (1.71 +/- 0.30) and S. syndactylus (1.55 +/- 0.04), were similar to the results for the quadrupedal anthropoids, C. guereza (male = 1.64 +/- 0.13; female = 1.68 +/- 0.07) and P. cynocephalus (1.47 +/- 0.13). These results suggest strong trabecular architecture similarity across body sizes, anthropoid phylogenetic backgrounds, and locomotor mode. This structural similarity might be explained by greater similarity in anthropoid hip joint loading mechanics than previously considered. It is likely that our current models of anthropoid hip joint mechanics are overly simplistic.

Effects of limb mass distribution on the ontogeny of quadrupedalism in infant baboons (Papio cynocephalus) and implications for the evolution of primate quadrupedalism.

Primate quadrupedal kinematics differ from those of other mammals. Several researchers have suggested that primate kinematics are adaptive for safe travel in an arboreal, small-branch niche. This study tests a compatible hypothesis that primate kinematics are related to their limb mass distribution patterns. Primates have more distally concentrated limb mass than most other mammals due to their grasping hands and feet. Experimental studies have shown that increasing distal limb mass by adding weights to the limbs of humans and dogs influences kinematics. Adding weights to distal limb elements increases the natural period of a limb's oscillation, leading to relatively long swing and stride durations. It is therefore possible that primates' distal limb mass is responsible for some of their unique kinematics. This hypothesis was tested using a longitudinal ontogenetic sample of infant baboons (Papio cynocephalus). Because limb mass distribution changes with age in infant primates, this project examined how these changes influence locomotor kinematics within individuals. The baboons in this sample showed a shift in their kinematics as their limb mass distributions changed during ontogeny. When their limb mass was most distally concentrated (at young ages), stride frequencies were relatively low, stride lengths were relatively long, and stance durations were relatively long compared to older ages when limb mass was more proximally concentrated. These results suggest that the evolution of primate quadrupedal kinematics was tied to the evolution of grasping hands and feet.

Ontogeny of limb mass distribution in infant baboons (Papio cynocephalus).

Primates have more distally distributed limb muscle mass compared to most nonprimate mammals. The heavy distal limbs of primates are likely related to their strong manual and pedal grasping abilities, and interspecific differences in limb mass distributions among primates are correlated with the amount of time spent on arboreal supports. Within primate species, individuals at different developmental stages appear to differ in limb mass distribution patterns. For example infant macaques have more distally distributed limb mass at young ages. A shift from distal to proximal limb mass concentrations coincides with a shift from dependent travel (grasping their mother's hair) to independent locomotion. Because the functional demands placed on limbs may differ between taxa, understanding the ontogeny of limb mass distribution patterns is likely an essential element in interpreting the diversity of limb mass distribution patterns present in adult primates. This study examines changes in limb inertial properties during ontogeny in a longitudinal sample of infant baboons (Papio cynocephalus). The results of this study show that infant baboons undergo a transition from distal to proximal limb mass distribution patterns. This transition in limb mass distribution coincides with the transition from dependent to independent locomotion during infant development. Compared to more arboreal macaques, infant baboons undergo a faster transition to more proximal limb mass distribution patterns. These results suggest that functional demands placed on the limbs during ontogeny have a strong impact on the development of limb mass distribution patterns.