Kevin D. Hunt

Dietary Response of Chimpanzees and Cercopithecines to Seasonal Variation in Fruit Abundance. II. Macronutrients

In order to understand dietary differentiation among frugivorous primates with simple stomachs, we present the first comparison of plant diets between chimpanzees and cercopithecine monkeys that controls for food abundance. Our aim was to test the hypothesis that monkeys have a more diverse diet as a result of their dietary tolerance for chemical antifeedants. Our study species are chimpanzees, blue monkeys, redtail monkeys, and gray-cheeked mangabeys living in overlapping ranges in Kibale National Park, Uganda. We indexed food abundance by the percentage of trees having ripe fruit within the range of each group; it varied widely during the year. Chimpanzees spent almost 3 times as much of their feeding time eating ripe fruits as the monkeys did and confined their diets almost exclusively to ripe fruits when they were abundant. Monkeys maintained a diverse diet at all times. When ripe fruit was scarce chimpanzee and monkey diets diverged. Chimpanzees relied on piths as their main fallback food, whereas monkeys turned to unripe fruits and seeds. For each primate group we calculated the total weighted mean intake of 5 antifeedants; condensed tannins (CT), total tannins assayed by radial diffusion (RD), monoterpenoids (MT), triterpenoids (TT), and neutral-detergent fiber (NDF). Monkeys had absolutely higher intakes of CT, RD, MT, and TT than those of chimpanzees, and their intake of NDF did not differ from that of chimpanzees, appearing relatively high given their lower body weights. However contrary to expectation, dietary divergence during fruit scarcity was not associated with any change in absolute or relative intake of antifeedants. For example, fruit scarcity did not affect the relative intake of antifeedants by cercopithecines compared to chimpanzees. Our results establish chimpanzees as ripe-fruit specialists, whereas cercopithecines are generalists with a higher intake of antifeedants. The low representation of ripe fruits in the diets of cercopithecines has not been explained. An important next step is to test the hypothesis that the difference between Kibale chimpanzees and cercopithecines represents a more general difference between apes and monkeys.

Estimators of Fruit Abundance of Tropical Trees1

Many types of biological studies require the estimation of food abundance in tropical forests, and a variety of methods have been used to estimate this parameter. Here we compare the accuracy and precision of three methods for estimating the fruit abundance (biomass and number) of tropical tree species: tree diameter, crown volume, and visual estimation. Diameter at breast height (DBH) was the most consistently accurate method and exhibited low levels of interobserver variability. Generally, crown volume was neither precise nor accurate. The visual estimation method was accurate for trees with very large fruit, but exhibited high interobserver variability.

Standardized descriptions of primate locomotor and postural modes

As quantitative studies on primate positional behavior accumulate the lack of a standard positional mode terminology is becoming an increasingly serious deficiency. Inconsistent use of traditional terms and inappropriate conflation of mode categories hamper interspecific and interobserver comparisons. Some workers use common terms without definition, allowing at least the possibility of misunderstanding. Other researchers coin neologisms tailored to their study species and not clearly enough defined to allow application to other species. Such neologisms may overlap, may completely encompass, or may conflate previously defined labels. The result is, at best, the proliferation of synonyms and, at worst, the creation of confusion where clarity had existed. Historical precedents have sometimes resulted in “catch-all” terms that conflate any number of kinematically different behaviors (e.g. “brachiation,” “climbing,” and “quadrumanous climbing”). We recognize three areas where distinction of positional modes has some current importance: (1) Modes that require humeral abduction should be distinguished from adducted behaviors; (2) locomotor modes that involve ascent or descent should be distinguished from horizontal locomotor modes; and (3) suspensory modes should be distinguished from supported modes. We recommend a nomenclature that is not dedicated to or derived from any one taxonomic subset of the primates. Here we define 32 primate positional modes, divided more finely into 52 postural sub-modes and 74 locomotor sub-modes.

Gradual Change In Human Tooth Size In The Late Pleistocene And Post‐Pleistocene

Starting with the onset of the last glaciation approximately 100,000 years ago and continuing to the end of the Late Pleistocene approximately 10,000 years ago, human tooth size began to reduce at a rate of 1% every 2,000 years. Both the mesial‐distal and the buccal‐lingual dimensions of mandibular and maxillary teeth were undergoing the same rate of reduction. From the beginning of the Post‐Pleistocene until the present, the overall rate of dental reduction doubled, becoming approximately 1% per thousand years. Buccal‐lingual dimensions are now reducing twice as fast as mesial‐distal dimensions, and maxillary teeth are reducing at an even more rapid rate than mandibular teeth. Late Pleistocene rates are comparable in Europe and the Middle East. The Post‐Pleistocene rates are also the same for Europe, the Middle East, China, Japan, and Southeast Asia. It is suggested that the cookery at the beginning of the Late Pleistocene allowed the earlier changes to occur. The use of pottery within the last 10,000 years further reduced the amount of selection that had previously maintained usable tooth substance. Reduction then occurred as a consequence of the Probable Mutation Effect (Brace, 1963; McKee, 1984).

Positional behavior in the Hominoidea

Quantitative studies on the positional behavior of members of the Hominoidea are compared in order (1) to identify consistencies across the superfamily, (2) to contrast ape positional behavior with that of Old World monkeys (forest-livingPapio anubis were chosen for study to reduce body size effects), and (3) to identify distinctive behaviors in each of the ape taxa. Differences in the way behaviors were sampled in the various studies necessitated considering posture and locomotion separately. Unimanual arm-hanging and vertical climbing were the most distinctive shared postural and locomotor modes among the apes (the gorilla excepted), constituting ≥5.0% and ≥4.9% of all behavior in each species. Arm-hanging and brachiation (sensu stricto) frequencies were the highest by far in hylobatids. Hand-foot hanging, bipedal posture, and clambering, an orthograde suspensory locomotion assisted by the hindlimbs, were more common in orangutans than in any other hominoid. Sitting and walking were observed in the highest frequencies in the African apes but were no more common than in the baboon. Relatively high frequencies of brachiation (sensu stricto) were reported for all apes except chimpanzees and gorillas. Brachiation and arm-hanging were kinematically different in apes and baboons, involving complete humeral abduction only in the former, whereas vertical climbing appeared to be kinematically similar in apes and baboons. It is concluded that the morphological specializations of the apes may be adaptations to (1) the unique physical demands of arm-hanging and (2) less kinematically distinct, but still quantitatively significant, frequencies of vertical climbing.

The Value of Figs to Chimpanzees

Nine Ugandan figs have consistent differences in nutrient concentration between the pulp and seed fractions. Pulp has more water-soluble carbohydrates, complex carbohydrates, calories, and ash, while the seed fraction has more condensed tannins, lipids, and fiber. Because species differ, nutrient concentration in pulp could not be predicted from analysis of whole figs. Chimpanzees in Kibale Forest relied heavily on figs throughout 29 months, feeding relatively intensely at large trees. Fig size varied between species, between individuals of the same species, and between fruiting cycles of the same tree. Larger figs had higher water concentrations but still led to higher rates of nutrient intake per minute for chimpanzees, monkeys, and hornbills. Chimpanzees ate more than 40 cal/min, excluding calories derived from insoluble fiber, when harvesting large figs.

EMG study of hand muscle recruitment during hard hammer percussion manufacture of Oldowan tools

The activity of 17 hand muscles was monitored by electromyography (EMG) in three subjects during hard hammer percussion manufacture of Oldowan tools. Two of the subjects were archaeologists experienced in the replication of prehistoric stone tools. Simultaneous videotapes recorded grips associated with the muscle activities. The purpose of the study was to identify the muscles most likely to have been strongly and repeatedly recruited by early hominids during stone tool-making. This information is fundamental to the identification of skeletal features that may reliably predict tool-making capabilities in early hominids. The muscles most frequently recruited at high force levels for strong precision pinch grips required to control the hammerstone and core are the intrinsic muscles of the fifth finger and the thumb/index finger regions. A productive search for skeletal evidence of habitual Oldowan tool-making behavior will therefore be in the regions of the hand stressed by these intrinsic muscles and in the joint configurations affecting the relative lengths of their moment arms.

The Genetic Signature of Sex-Biased Migration in Patrilocal Chimpanzees and Humans

A large body of theoretical work suggests that analyses of variation at the maternally inherited mitochondrial (mt)DNA and the paternally inherited non-recombining portion of the Y chromosome (NRY) are a potentially powerful way to reveal the differing migratory histories of men and women across human societies. However, the few empirical studies comparing mtDNA and NRY variation and known patterns of sex-biased migration have produced conflicting results. Here we review some methodological reasons for these inconsistencies, and take them into account to provide an unbiased characterization of mtDNA and NRY variation in chimpanzees, one of the few mammalian taxa where males routinely remain in and females typically disperse from their natal groups. We show that patterns of mtDNA and NRY variation are more strongly contrasting in patrilocal chimpanzees compared with patrilocal human societies. The chimpanzee data we present here thus provide a valuable comparative benchmark of the patterns of mtDNA and NRY variation to be expected in a society with extremely female-biased dispersal.

A Thermodynamic Comparison of Arboreal and Terrestrial Sleeping Sites for Dry-Habitat Chimpanzees (Pan troglodytes schweinfurthii) at the Toro-Semliki Wildlife Reserve, Uganda

The nightly construction of an arboreal sleeping platform (SP) has been observed among every chimpanzees population studied to date. Here, we report on bioclimatic aspects of SP site choice among dry‐habitat chimpanzees (Pan troglodytes schweinfurthii) at the Toro‐Semliki Wildlife Reserve, Uganda. We placed a portable weather monitor within 1 m of chimpanzee SPs and compared the microenvironment of this site with terrestrial monitors placed 10 cm above the ground directly underneath the simultaneously studied SP. We calculated physical “comfort levels” of monitored sites using the RayMan thermophysiological model that we modified to take ape body proportions into account. The RayMan tool gauges energy balance using wind speed, temperature, relative humidity, and heat index in conjunction with the study subjects mass and stature to determine whether the individual is in energy balance or homeostasis. We found that (1) terrestrial microclimates have greater homeostatic potential than arboreal microclimates, and (2) there is a significant positive linear relationship between wind speed and height of SP in the forest canopy. Advantages of terrestrial sites are that they require lesser energetic expenditure to stabilize the body when the SP is under construction and perhaps during use as well. We found that terrestrial sites also had better homeostatic potentials. This combination of advantages explains why SPs are so often sited terrestrially in habitats where predation risk is low. Early hominins must have had technological or social measures to avoid or deter predators that were significantly advanced over those found among chimpanzees before they began sleeping on the ground. Am. J. Primatol. 74:811–818, 2012.

Social rank and body size as determinants of positional behavior inPan troglodytes

A yearlong study of the positional behavior ofPan troglodytes at the Mahale Mountains National Park yielded 571 hr of observation.Cant (1987) articulated four predictions concerning the relationship between body weight/branch diameter and positional behavior based on the classic suspensory-ape paradigm. He noted that only two were supported by orangutan data. Three of these four hypothesis were not supported by chimpanzee data, as follows: there was no significant difference between the three largest males and the four smallest males in (1) branch diameters (5.8 cm vs 5.2 cm) nor (2) in the percentage of arm-hanging (13.6% vs 12.1%); and (3) large males did not arm-hang significantly more often than small males in any of three support diameter categories. The fourth hypothesis, that arm-hanging should be more common among smaller branches, was supported: arm-hanging as a percentage of all posture rose from 2.5% to 8.3% to 24% as stratum size decreased from >10 cm to <3 cm. The possibility that the first three hypotheses failed because of confounding effects of a correlation between body size and social rank was examined. Multiple regressions were done on 6600 2-min instantaneous observations on focal individuals. With social rank effects factored out, larger individuals preferentially utilized smaller, rather than larger supports. When positional mode frequencies were compared between large and small males matched for social rank, large males exhibited a lower frequency of arm-hanging than small males. An unexpected result was that social rank more consistently predicted branch diameter choice than body size. The most profound trend was for high ranking males to use larger supports, even though they spent more time in the terminal branches. These results suggest that (1) suspensory behavior is functionally related to small branch diameters; (2) chimpanzees do not prefer smaller branches, rather they are forced into them by food choice limitations; and (3) social rank more profoundly affects chimpanzee behavior than body weight.