Ontogeny and the evolution of adult body size dimorphism in apes

This analysis investigates the ontogeny of body size dimorphism in apes. The processes that lead to adult body size dimorphism are illustrated and described. Potential covariation between ontogenetic processes and socioecological variables is evaluated. Mixed-longitudinal growth data from 395 captive individuals (representing Hylobates lar [gibbon], Hylobates syndactylus [siamang], Pongo pygmaeus [orangutan], Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) form the basis of this study. Results illustrate heterogeneity in the growth processes that produce ape dimorphism. Hylobatids show no sexual differentiation in body weight growth. Adult body size dimorphism in Pongo can be largely attributed to indeterminate male growth. Dimorphism in African apes is produced by two different ontogenetic processes. Both pygmy chimpanzees (Pan paniscus) and gorillas (Gorilla gorilla) become dimorphic primarily through bimaturism (sex differences in duration of growth). In contrast, sex differences in rate of growth account for the majority of dimorphism in common chimpanzees (Pan troglodytes). Diversity in the ontogenetic pathways that produce adult body size dimorphism may be related to multiple evolutionary causes of dimorphism. The lack of sex differences in hylobatid growth is consistent with a monogamous social organization. Adult dimorphism in Pongo can be attributed to sexual selection for indeterminate male growth. Interpretation of dimorphism in African apes is complicated because factors that influence female ontogeny have a substantial effect on the resultant adult dimorphism. Sexual selection for prolonged male growth in gorillas may also increase bimaturism relative to common chimpanzees. Variation in female growth is hypothesized to covary with foraging adaptations and with differences in female competition that result from these foraging adaptations. Variation in male growth probably corresponds to variation in level of sexual selection. © 1995 Wiley-Liss, Inc.     

Epidermal patterns of the hands and feet of the pygmy chimpanzee (Pan paniscus)

The characteristics of the epidermal ridge system were studied in a series of eighteen lesser or pygmy chimpanzees (Pan paniscus). The general ridge alignments are very similar to those of the chimpanzee (Pan troglodytes); Biegert ('61). On the average the pattern intensity (P.I.) of the palm configurations is considerably higher in the pygmy chimpanzee than in the chimpanzee, thus representing the highest total palm pattern intensity of all species of the Hominoidea. The sole configurations show parallel main results to those of the palm; however, the decreased sole pattern frequency of the pygmy chimpanzee is of a smaller predominance only as compared to the values of the other species of this superfamily. The preliminary data on the finger tip patterns, translated into P.I. values, are much higher than in chimpanzees and within the range of the mean values of gorillas (Brehme, '73), while those of the toes of pygmy chimpanzees seem to possess the lowest P.I. values of the African apes.     

Scapula form and locomotion in chimpanzee evolution

A number of primatologists have followed Coolidge (Am. J. Phys. Anthropol. 18:1–57, 1933) in suggesting that 1) there are significant shape differences in scapula form between pygmy and common chimpanzees, 2) scapulae of P. paniscus resemble those of hylobatids more than do those of P. troglodytes, and 3) therefore pygmy chimpanzees may exhibit a greater component of arm-swinging and other arboreal behaviors than common chimpanzees. In this paper I utilize a comparative analysis of ontogenetic allometries of linear dimensions to determine shape differences in the scapulae of adult pygmy and common chimpanzees and to clarify size-related changes in shape resulting from ontogenetic scaling, i.e., the differential extension of common patterns of growth allometry. Results demonstrate that the scapulae of adult P. paniscus are relatively narrower (in a direction approximately perpendicular to the scapula spine) than those of P. troglodytes, supporting Coolidge's original claim. The allometric analysis further demonstrates, however, that the two chimpanzee species exhibit ontogenetic scaling for all proportions of the scapula examined. Thus, adult pygmy chimpanzees have the scapula proportions observed in small adult and subadult P. troglodytes of comparable scapula size. The implications of this finding for past claims concerning differences in locomotor behavior between the species are discussed. This work lends additional support to previous studies that have demonstrated a high frequency of ontogenetic scaling within the genus Pan and a pedomorphic or juvenilized morphology in the pygmy chimpanzee.     

Knuckle-walking and the evolution of hominoid hands

Knuckle-walking is a pattern of digitigrade locomotion unique to African apes among Primates. Only chimpanzees and gorillas are specially adapted for supporting weight on the dorsal aspects of middle phalanges of flexed hand digits II–V. When forced to the ground, most orangutans assume one of a variety of flexed hand postures, but they cannot knuckle-walk. Some orangutans place their hands in palmigrade postures which are impossible to African apes. The knuckle-walking hands and plantigrade feet of African apes are both morphologically and adaptively distinct from those of Pongo, their nearest relative among extant apes. These features are associated with a common adaptive shift to terrestrial locomotion and support placing chimpanzees and gorillas in the same genus Pan. It is further suggested than Pan comprises the subgenera (a) Pan, including P. troglodytes and pygmy chimpanzees, and (b) Gorilla, including mountain and lowland populations of P. gorilla. African apes probably diverged from ancestral pongids that were specially adapted for distributing their weight in terminal branches of the forest canopy. Early adjustments to terrestrial locomotion may have involved fist-walking which later evolved into knuckle-walking. Orangutans continued to adapt to feeding and locomotion in the forest canopy and their hands and feet became highly specialized for four-digit prehension. Although chimpanzees retained arboreal feeding and nesting habits, they moved from tree to tree by terrestrial routes and became less restricted in habitat. While adapting to a diet of ground plants gorillas increased in size to the point that arboreal nesting is less frequent among them than among chimpanzees and orangutans. Early hominids probably diverged from pongids that had not developed prospective adaptations to knuckle-walking, and therefore did not evolve through a knuckle-walking stage. Initial adjustments to terrestrial quadrupedal locomotion and resting stance probably included palmigrade hand posturing. Their thumbs may have been already well developed as an adaptation for grasping during arboreal climbing. A combination of selection pressures for efficient terrestrial locomotor support and for object manipulation further advanced early hominid hands toward modern human configuration.     

Masticatory form and function in the African apes

This study examines variability in masticatory morphology as a function of dietary preference among the African apes. The African apes differ in the degree to which they consume leaves and other fibrous vegetation. Gorilla gorilla beringei, the eastern mountain gorilla, consumes the most restricted diet comprised of mechanically resistant foods such as leaves, pith, bark, and bamboo. Gorilla gorilla gorilla, the western lowland gorilla subspecies, consumes leaves and other terrestrial herbaceous vegetation (THV) but also consumes a fair amount of ripe, fleshy fruit. In contrast to gorillas, chimpanzees are frugivores and rely on vegetation primarily as fallback foods. However, there has been a long-standing debate regarding whether Pan paniscus, the pygmy chimpanzee (or bonobo), consumes greater quantities of THV as compared to Pan troglodytes, the common chimpanzee. Because consumption of resistant foods involves more daily chewing cycles and may require larger average bite force, the mechanical demands placed on the masticatory system are expected to be greater in folivores as compared to primates that consume large quantities of fleshy fruit. Therefore, more folivorous taxa are predicted to exhibit features that improve load-resistance capabilities and increase force production. To test this hypothesis, jaw and skull dimensions were compared in ontogenetic series of G. g. beringei, G. g. gorilla, P. t. troglodytes, and P. paniscus. Controlling for the influence of allometry, results show that compared to both chimpanzees and bonobos, gorillas exhibit some features of the jaw complex that are suggestive of improved masticatory efficiency. For example, compared to all other taxa, G. g. beringei has a significantly wider mandibular corpus and symphysis, larger area for the masseter muscle, higher mandibular ramus, and higher mandibular condyle relative to the occlusal plane of the mandible. However, the significantly wider mandibular symphysis may be an architectural response to increasing symphyseal curvature with interspecific increase in size. Moreover, Gorilla and Pan do not vary consistently in all features, and some differences run counter to predictions based on dietary variation. Thus, the morphological responses are not entirely consonant with predictions based on hypothesized loading regimes. Finally, despite morphological differences between bonobos and chimpanzees, there is no systematic pattern of differentiation that can be clearly linked to differences in diet. Results indicate that while some features may be linked to differences in diet among the African apes, diet alone cannot account for the patterns of morphological variation demonstrated in this study. Allometric constraints and dental development also appear to play a role in morphological differentiation among the African apes.     

Relative growth of the limbs and trunk in the African apes

Examination of relative growth and allometry is important for our understanding of the African apes, as they represent a closely related group of species of increasing body size. This study presents a comparison of ontogenetic relative growth patterns of some postcranial dimensions in Pan paniscus, Pantroglodytes, and Gorilla gorilla. Interspecific proportion differences among the three species are also analyzed. It is stressed that reliable ontogenetic information can only be obtained if subadults are examined-growth data cannot be inferred from static adult scaling. Results indicate that some postcranial relative growth patterns are very similar in the three species, suggesting differential extrapolation of a common growth pattern, whereas for other proportion comparisons the growth trends differ markedly among the species, producing distinct shape differences in the adults Interspecific shape changes among the three species are characterized by positive allometry of chest girth and negative allometry of body height and leg length. It is suggested that relative decrease of leg length with increasing body size among the African pongids might be expected on biomechanical grounds, in order to maintain similar locomotor abilities of climbing arborealism and quadrupedal terrestrialism. Relative to body weight or trunk length, the limbs of the bonobo (Pan paniscus) are longer than in the common chimpanzee or the gorilla, with a lower intermembral index. This may most closely resemble the primitive condition for the African apes.     

Food Preferences Among Captive Western Gorillas (Gorilla gorilla gorilla) and Chimpanzees (Pan troglodytes)

I used a zoological park setting to address food preferences among gorillas (Gorilla gorilla gorill) and chimpanzees (Pan troglodytes). Gorillas and chimpanzees are different sizes, and consequently, have been traditionally viewed as ecologically distinct. Sympatric western gorillas and chimpanzees have proved difficult to study in the wild. Limited field data have provided conflicting information about whether gorillas are fundamentally different from chimpanzees in diet and behavior. Fruit eating shapes the behavior of most apes, but it is unclear whether the large-bodied gorillas are an exception to this rule, specifically whether they are less selective and more opportunistic fruit eaters than chimpanzees are. My research provides experimental observational data to complement field data and to better characterize the diets and food preferences of the African apes. During laboratory research at the San Francisco Zoological Gardens, I examined individual and specific differences in food preferences of captive gorillas and chimpanzees via experimental paired-choice food trials with foods that varied in nutritional content. During the study, I offered 2500 paired-food choices to 6 individual gorillas and 2000 additional pairs to them as a group. I also proffered 600 food pairs to 4 individual chimpanzees. Despite expectations of the implications of body size differences for diet, gorillas and chimpanzees exhibited similar food preferences. Both species preferred foods high in non-starch sugars and sugar-to-fiber ratios, and low in total dietary fiber. Neither species avoided foods containing tannins. These data support other suggestions of African apes sharing a frugivorous adaptation.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

African ape ancestry

It is commonly believed that the australopithecines are more closely related to humans than to African apes. This view is hardly compatible with the biomolecular data which place theHomo/Pan split at the beginning of the australopithecine period. Nothing in the fossil hominid morphology precludes the possibility that some australopithecines were ancestral to gorillas or chimpanzees and others to humans.     

Comparison of cranial ontogenetic trajectories among great apes and humans

Molecular data suggest that humans are more closely related to chimpanzees than either is to the gorillas, yet one finds the closest similarity in craniofacial morphology to be among the great apes to the exclusion of humans. To clarify how and when these differences arise in ontogeny, we studied ontogenetic trajectories for Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. A total of 96 traditional three-dimensional landmarks and semilandmarks on the face and cranial base were collected on 268 adult and sub-adult crania for a geometric morphometric analysis. The ontogenetic trajectories are compared by various techniques, including a new method, relative warps in size-shape space. We find that adult Homo sapiens specimens are clearly separated from the great apes in shape space and size-shape space. Around birth, Homo sapiens infants are already markedly different from the great apes, which overlap at this age but diverge among themselves postnatally. The results suggest that the small genetic differences between Homo and Pan affect early human ontogeny to induce the distinct adult human craniofacial morphology. Pure heterochrony does not sufficiently explain the human craniofacial morphology nor the differences among the African apes.     

Diet and Mandibular Morphology in African Apes

Investigations seeking to understand the relationship between mandibular form, function, and dietary behavior have focused on the mandibular corpus and symphysis. African apes vary along a gradient of folivory/frugivory, yet few studies have evaluated the morphology of the mandibular corpus and symphysis in these taxa, and the investigations have yielded mixed results. Specifically, studies using external metrics have identified differences in mandibular proportions that analysis of cortical bone distribution has not substantiated. I contribute to the ongoing debate on the relationship between jaw form and dietary behavior by comparing mandibular corporal and symphyseal shapes in African apes. Importantly, and in contrast to previous studies of African ape internal geometry, I include the Virunga mountain gorillas (Gorilla beringei beringei), the ape most specialized toward a folivorous diet. I test the hypotheses that 1) Gorilla beringei beringei always has significantly more robust mandibular corpora and symphyses, relative to mandibular length, than all other African apes and 2) all gorillas have significantly more robust mandibular corpora and symphyses, relative to mandibular length, than Pan. Results demonstrate that the folivorous mountain gorillas consistently exhibit a relatively wider mandibular symphysis and corpus than all other African apes. Furthermore, all gorillas consistently exhibit relatively more robust mandibular corporal and symphyseal dimensions than Pan. The results indicate that among African apes, mountain gorillas are better able to counter lateral transverse bending (wishboning) loads at the symphysis and torsional loads at the corpus. All gorillas are likewise better able to resist wishboning and vertical bending at the symphysis, and sagittal bending and torsion at the corpus, than Pan, findings that are consistent with masticating relatively tougher foods, repetitive loading of the jaws, or both. I offer possible explanations for the lack of concordance in results between studies that have analyzed the biomechanical properties of African ape mandibles and others that have relied on external metrics. More comprehensive study of the internal geometry of the mandible is needed to resolve whether African apes differ morphologically in ways predicted by dietary variation.

Densities,Distributions, and Seasonal Movements of Gorillas and Chimpanzees in Swamp Forest in Northern Congo

The conservation of gorillas (Gorilla spp.) and chimpanzees (Pan troglodytes) depends upon knowledge of their densities and distribution throughout their ranges. However, information about ape populations in swamp forests is scarce. Here we build on current knowledge of ape populations by conducting line transect surveys of nests throughout a reserve dominated by swamp forest: the Lac Télé Community Reserve in northern Congo. We estimated gorilla and chimpanzee densities, distributions across habitats, and seasonal changes in abundance. Gorilla density was 2.9 gorillas km^–2, but densities varied by habitat (0.3–5.4 gorillas km^–2) with highest densities in swamp forest and terra firma mixed forest. Average chimpanzee density is 0.7 chimpanzees km^–2 (0.1–1.3 chimpanzees km^–2), with highest densities in swamp forest. Habitat was the best predictor of ape nest abundance, as neither the number of human indices nor the distance from the nearest village predicted nest abundance. We recorded significantly greater numbers of apes in terra firma forest during the high-water season than the low-water season, indicating that many gorillas and chimpanzees are at times concentrated in terra firma forest amid a matrix of swamp forest. Seasonally high numbers of apes on terra firma forest islands easily accessible to local people may expose them to substantial hunting pressure. Conversely, the nearly impenetrable nature of swamp forests and their low value for logging makes them promising sites for the conservation of apes.     

Comparison of the communicative vocalizations and behaviors of group ranging in eastern gorillas,chimpanzees and pygmy chimpanzees

Field observations were carried out on chimpanzees, pygmy chimpanzees and eastern gorillas. Since the communicative behaviors which appear in group ranging are closely related to the grouping of the animals and to the social structure, the communicative behaviors of group ranging were compared in the above three species in order to elucidate the common and different characters of their three much diversified social structures. The following conclusions were drawn: (1) the common ancestor ofGorilla andPan had the territorial call “hoot” and behavior of display, and males were antagonistic between each other in the society; (2) the common ancestral species did not have any special long distance cohesive calling: the society was a small compact one, moving on the ground; and (3) the social structure of the pygmy chimpanzee is very different from the common ancestral social structure when compared with those of the gorilla and chimpanzee, in that the pygmy chimpanzee has lost the behaviors of strong antagonistic character between adult males.     

Quantitative and functional studies on the hands of the anthropoidea. I. The Hominoidea

The hands of the Hominoidea evidence four adaptive modes which distinguish the lesse apes (Hylobatidae), the orangutan (Pongo), the African apes (Pan), and man (Homo) from one another. The hands of the apes consist of compromises between manipulatory and locomotor functions because selection has operated for precision of grip as well as for special locomotor mechanisms. The human hand is almost totally devoted to manipulation. The hands of gibbons, orangutans and the African apes differ in many features that may be correlated with locomotion. The gibbons and siamang are specially adapted for ricochetal arm-swinging. The great apes possess morphological adaptations for arboreal foraging and climbing distinct from those of the hylobatids. In addition, the African apes have become secondarily adapted for terrestrial quadrupedal locomotion. Many features that distinguish the hands of chimpanzees and gorillas may be associated with the development of efficient knuckele-walking propulsive and support mechanisms.     

The role of taste in food selection by African apes: implications for niche separation and overlap in tropical forests

Ripe fruit eating shapes the behavior of most of the apes. Gorillas (Gorilla gorilla) and chimpanzees (Pan troglodytes) are very different sizes and, consequently, have been traditionally viewed as ecologically distinct, but few studies have explored the behavioral and physiological foundations of their diets. Debate continues on the extent that large-bodied gorillas may be less selective and more opportunistic fruit eaters than chimpanzees. Taste responses have been predicted to relate to body size and digestive strategies. This study employs laboratory research on taste perception and discrimination among captive zoo-housed chimpanzees and relates it to previous work on gorillas to better characterize diets and niche separation among these apes. During the captive trials, differences were recorded in consumption patterns of water and varying concentrations of dilute aqueous fructose (sweet) and tannic acid solutions (astringent), compounds commonly found in wild foods. The chimpanzees exhibited similar preference thresholds for fructose (50 mM) to other primates studied. They exhibited slightly lower inhibition thresholds for tannic acid solutions than gorillas, but higher than smaller primates studied to date. These preliminary findings suggest that tannin tolerance may well be mediated by body size, though possible species differences in salivary proteins or other sensory differences remain to be explored. This research furthers our efforts to understand the roles of body size and physiological adaptations in shaping diet and niche separation of chimpanzees and gorillas.     

Diet and seasonal changes in sympatric gorillas and chimpanzees at Kahuzi–Biega National Park

Based on 8 years of observations of a group of western lowland gorillas (Gorilla beringei graueri) and a unit-group of chimpanzees (Pan troglodytes schweinfurthii) living sympatrically in the montane forest at Kahuzi–Biega National Park, we compared their diet and analyzed dietary overlap between them in relation to fruit phenology. Data on fruit consumption were collected mainly from fecal samples, and phenology of preferred ape fruits was estimated by monitoring. Totals of 231 plant foods (116 species) and 137 plant foods (104 species) were recorded for gorillas and chimpanzees, respectively. Among these, 38% of gorilla foods and 64% of chimpanzee foods were eaten by both apes. Fruits accounted for the largest overlap between them (77% for gorillas and 59% for chimpanzees). Gorillas consumed more species of vegetative foods (especially bark) exclusively whereas chimpanzees consumed more species of fruits and animal foods exclusively. Although the number of fruit species available in the montane forest of Kahuzi is much lower than that in lowland forest, the number of fruit species per chimpanzee fecal sample (average 2.7 species) was similar to that for chimpanzees in the lowland habitats. By contrast, the number of fruit species per gorilla fecal sample (average 0.8 species) was much lower than that for gorillas in the lowland habitats. Fruit consumption by both apes tended to increase during the dry season when ripe fruits were more abundant in their habitat. However, the number of fruit species consumed by chimpanzees did not change according to ripe fruit abundance. The species differences in fruit consumption may be attributed to the wide ranging of gorillas and repeated usage of a small range by chimpanzees and/or to avoidance of inter-specific contact by chimpanzees. The different staple foods (leaves and bark for gorillas and fig fruits for chimpanzees) characterize the dietary divergence between them in the montane forest of Kahuzi, where fruit is usually scarce. Gorillas rarely fed on insects, but chimpanzees occasionally fed on bees with honey, which possibly compensate for fruit scarcity. A comparison of dietary overlap between gorillas and chimpanzees across habitats suggests that sympatry may not influence dietary overlap in fruit consumed but may stimulate behavioral divergence to reduce feeding competition between them.     

Sexual differences in the anterior dentition in African primates

A study was made of ranges of variation of the anterior dentition of various African nonhuman primates. Comparisons of the dentitions were made between different species and sex differences within each species were determined. Among the nonhuman primate groups studied were: Gorilla gorilla gorilla, Pan troglodytes, Pan paniscus, Cercopithecus nictitans, Cercocebus albigena and Colobus badius. In monkeys the canine teeth of the males are considerably larger than those of the females. There are also considerable differences in size in the rest of the anterior dentition. In apes, and specifically only gorillas, distinct sex differences are only found in the maxillary canines. In the chimpanzees, sex differences in the dentition are much smaller and there is considerable overlap in the ranges of variation. There are no fundamental differences in the size of the rest of the anterior dentition in the apes. The present study shows that differences due to sex in the anterior dentition, excluding the canine, are not as great as has been considered. If we consider the fossil record of man, whose morphological complex includes a much reduced canine, the probability will be that sex differences in the rest of the dentition will be negligible. Given the fragmentary nature of the fossil record, it is, therefore, highly unlikely that the determination of the sex of any fossil hominid specimen can be accurately made based solely on the evidence of its dentition.     

A comparative analysis of temporomandibular joint morphology in the African apes

A number of researchers have suggested a functional relationship between dietary variation and temporomandibular joint (TMJ) morphology, yet few studies have evaluated TMJ form in the African apes. In this study, I compare TMJ morphology in adults and during ontogeny in Gorilla (G.g. beringei, G.g. graueri, and G.g. gorilla) and Pan (P. paniscus, P. troglodytes troglodytes, P.t. schweinfurthii, and P.t. verus). I test two hypotheses: first, compared to all other African apes, G.g. beringei exhibits TMJ morphologies that would be predicted for a primate that consumes a diet comprised primarily of moderately to very tough, leafy vegetation; and second, all gorillas exhibit the same predicted morphologies compared to Pan. Compared to all adult African apes, G.g. beringei has higher rami and condyles positioned further above the occlusal plane of the mandible, relative to jaw length. Thus, mountain gorillas have the potential to generate relatively more muscle force, more evenly distribute occlusal forces along the postcanine teeth, and generate relatively greater jaw adductor moment. G.g. beringei also exhibits relatively wider mandibular condyles, suggesting these folivorous apes are able to resist relatively greater compressive loads along the lateral and/or medial aspect of the condyle. All gorillas likewise exhibit these same shape differences compared to Pan. These morphological responses are the predicted consequences of intensification of folivory and, as such, provide support for functional hypotheses linking these TMJ morphologies to degree of folivory. The African apes to not, however, demonstrate a systematic pattern of divergence in relative condylar area as a function of intensification of folivory. The ontogenetic trajectories for gorillas are significantly elevated above those of Pan, and to a lesser but still significant degree, mountain gorillas similarly deviate from lowland gorillas (G.g. gorilla and G.g. graueri). Thus, adult shape differences in ramal and condylar heights do not result from the simple extrapolation of common growth allometries relative to jaw length. As such, they are suggestive of an adaptive shift towards a tougher, more folivorous diet. However, the allometric patterning for condylar area and condylar width does not systematically conform to predictions based on dietary specialization. Thus, while differences in condylar shapes may confer functional advantages both during growth and as adults, there is no evidence to suggest selection for altered condylar proportions, independent of the effects of changes in jaw size.     

Apes in a changing world – the effects of global warming on the behaviour and distribution of African apes

Aim In this study we use a modelling approach to identify: (1) the factors responsible for the differences in ape biogeography, (2) the effects that global warming might have on distribution patterns of African apes, (3) the underlying mechanisms for these effects, and (4) the implications that behavioural flexibility might be expected to have for ape survival. All African apes are highly endangered, and the need for efficient conservation methods is a top priority. The expected changes in world climate are likely to further exacerbate the difficulties they face. Our study aims to further understand the mechanisms that link climatic conditions to the behaviour and biogeography of ape species. Location Africa. Method We use an existing validated time budgets model, derived from data on 20 natural populations of gorillas (Gorilla beringei and Gorilla gorilla) and chimpanzees (Pan troglodytes and Pan paniscus), which specifies the relationship between climate, group size, body weight and time available for various activities, to predict ape distribution across Africa under a uniform worst‐case climate change scenario. Results We demonstrate that a worst‐case global warming scenario is likely to alter the delicate balance between different time budget components. Our model points to the importance of annual temperature variation, which was found to have the strongest impact on ape biogeography. Our simulation indicates that rising temperatures and changes in rainfall patterns are likely to have strong effects on ape survival and distribution, particularly for gorillas. Even if they behaved with maximum flexibility, gorillas may not be able to survive in most of their present habitats if the climate was to undergo extreme changes. The survival of chimpanzees was found to be strongly dependent on the minimum viable group size required. Main conclusions Our model allows us to explore how climatic conditions, individual behaviour and morphological traits may interact to limit the biogeographical distributions of these species, thereby allowing us to predict the effects of climate change on African ape distributions under different climate change regimes. The model suggests that climate variability (i.e. seasonality) plays a more important role than the absolute magnitude of the change, but these data are not normally provided by climate models.