Linear enamel hypoplasia in the great apes: analysis by genus and locality

Most studies report a high prevalence of linear enamel hypoplasia (LEH) in the great apes relative to other nonhuman primates and some human populations. It is unclear if this difference is a direct result of poor health status for the great apes, or if it represents differential incidence due to a lower threshold (sensu Goodman and Rose, 1990 Am. J. Phys. Anthropol. [suppl.] 33:59-110) for the occurrence of enamel hypoplasia among great apes. This study uses the Smithsonian National Museum of Natural History's great ape collection to examine the prevalence of LEH, the most common type of hypoplasia observed. Frequencies of LEH are reported, as well as analyses by taxa and provenience. The study sample consists of 136 specimens and includes 41 gorillas, 25 chimpanzees, and 70 orangutans. Analyses of frequencies are presented for both individuals and teeth by taxonomic category and locality. Among the individuals in this study, 63.97% are affected by LEH. Overall, gorillas (29.27%) exhibit lower frequencies of LEH than chimpanzees (68.00%) and orangutans (82.86%). There is a marked difference in LEH frequencies between mountain and lowland gorillas. There is no difference in LEH frequencies between Sumatran and Bornean orangutans. A range of variation for the great apes in enamel hypoplasia frequencies is found when taxon and locality are considered. It is likely that both biological and environmental factors influence the high frequencies of enamel hypoplasia exhibited in the great apes.     

Enamel hypoplasia in deciduous teeth of great apes: do differences in defect prevalence imply differential levels of physiological stress?

This paper presents new data on enamel hypoplasia in the deciduous canine teeth of great apes. The enamel defect under consideration is known as localized hypoplasia of primary canines (LHPC), and is characterized by an area of thin or missing enamel on the labial surface of deciduous canine teeth (Skinner [1986a] Am. J. Phys. Anthropol. 69:59-69). Goals of this study are: 1) to determine if significant differences in the frequency of LHPC occur among three genera of great apes, and 2) to evaluate variation in LHPC prevalence among great apes as evidence of differential physiological stress. Infant and juvenile apes with deciduous teeth were examined at the Cleveland Museum of Natural History (n = 100) and at the Smithsonian Institution, National Museum of Natural History (n = 36). Deciduous teeth were observed under oblique incandescent light, with the naked eye and with a 10x hand lens. Enamel hypoplasia was scored using Federation Dentaire International (FDI)-Defects of Dental Enamel (DDE) standards. Hypoplasias were recorded by drawing defect location and size on a dental chart, and by measuring defect size and location with Helios needlepoint dial calipers. The prevalence of LHPC is reported by genus and sex, using two approaches: 1) the frequency of affected individuals-those having one or more deciduous canine teeth scored positive for LHPC; and 2) the number of canine teeth scored positive for LHPC as a percentage of all canine teeth observed. Variation in defect size and location will be described elsewhere. Localized hypoplasia of primary canine teeth was found in 62.5% of 128 individual apes, and in 45.5% of 398 great ape deciduous canines. As in humans, LHPC is the most common form of enamel hypoplasia in deciduous teeth of great apes, while LEH is rare or absent. The distribution and pattern of expression of LHPC in great apes is similar to that described in humans: side differences are not significant, but mandibular canines exhibit the defect two to five times more often than maxillary canine teeth. Differences in LHPC prevalence by sex are small and not significant. Intergeneric differences are large and non-random: chimpanzees (Pan) exhibit a significantly lower frequency of LHPC (22%, n = 50) by individual count, than either the orangutan (Pongo, 88.0%, n = 25) or the gorilla (Gorilla, 88.7%, n = 53). Tooth count prevalences exhibit a similar pattern of variation and are also statistically significant. These findings suggest that large bodied great apes (gorilla and orangutan) may be under greater physiological stress during perinatal and early postnatal development than the chimpanzee. The size, position, and timing of LHPC lesions are currently under analysis and may yield more insight into the etiological origin of this enamel defect.     

Enamel hypoplasia in the deciduous teeth of early Miocene catarrhines: evidence of perinatal physiological stress

Enamel hypoplasia (EH) is a deficiency in enamel thickness due to physiological insults that compromise ameloblast function during the secretory phase of amelogenesis. The prevalence of EH in the deciduous teeth of nonhuman primates is largely unknown. One exception is the recent discovery of EH in the deciduous teeth of extant great apes which exhibit significant differences in prevalence between genera (Lukacs, 1999 a, 2000 a, Am. J. phys. Anthrop.110, 351-363). EH in deciduous teeth of other primates, living and fossil, remain undocumented. This communication describes a "plane form" type of EH known as localized hypoplasia of primary canines (LHPC) (Skinner, 1986 a, Am. J. phys. Anthrop.69, 59-69) in early Miocene catarrhines from Kenya. Specimens were examined macroscopically, with a 10x hand lens and with a variable power (10-30x) binocular microscope. Fédération Dentaire International (FDI)/Defects of Dental Enamel (DDE) standards were employed in recognition and recording of enamel defects (Fédération Dentaire International, 1982, Int. Dent. J.32, 159-167; Clarkson, 1989, Adv. Dental Res.3, 104-109). Size, shape and location of defects were measured and recorded on an outline drawing of the tooth crown. The Kenya National Museum study sample includes six genera of early Miocene catarrhines (n=66 specimens, with n=80 teeth). Seven deciduous teeth were afflicted with EH, yielding an overall prevalence of 8.75%. Two taxa, Kalepithecus (n=1 deciduous canine) and Proconsul (n=3 deciduous canines), were affected with LHPC. Expression of LHPC in fossil catarrhines is consistent with the expression of EH observed in skeletal samples of extant great apes. This report establishes an approximately 17-23 Ma antiquity for EH among early catarrhines and suggests that the neonatal stage of ontogenetic development was sufficiently stressful physiologically to produce disruption in amelogenesis. These physiological stresses impacted neonates of fossil taxa with a wide range of adult body sizes, from large-bodied Proconsul major ( approximately 75 kg) to one of the smaller-bodied catarrhines, Kalepithecus ( approximately 5 kg).     

A quantitative and qualitative reanalysis of the endocast from the juvenile Paranthropus specimen l338y-6 from Omo, Ethiopia

Based on an analysis of its endocast, Holloway (1981 Am J Phys Anthropol 53:109-118) attributed the juvenile Omo L338y-6 specimen to Australopithecus africanus (i.e., gracile australopithecines) rather than to Paranthropus (Australopithecus) boisei (robust australopithecines) favored by other workers (Rak and Howell [1978] Am J Phys Anthropol 48:345-366). Holloway's attribution was based on the specimen's (1) low cranial capacity, (2) gracile-like meningeal vessels, (3) gracile-like cerebellar hemispheres, and (4) absence of an enlarged occipital/marginal (O/M) sinus system. Recent work, however, has shown that criteria 1 and 2 are not useful for sorting gracile from robust australopithecines (Culotta [1999] Science 284:1109-1111; Falk [1993] Am J Phys Anthropol 92:81-98). In this paper, we test criterion 3 by quantifying the endocranial cerebellar and occipital morphology reproduced on the Omo L338y-6 endocast, and comparing it to seven endocasts from South and East African early hominids. Our preliminary results show that metric analysis of this specimen cannot be used to sort it preferentially with either robust or gracile australopithecines. Finally, we demonstrate that, contrary to previous reports, the Omo L338y-6 endocast reproduces an enlarged left occipital sinus (criterion 4). This observation is consistent with the original attribution of the Omo specimen to robust australopithecines (Rak and Howell [1978] Am J Phys Anthropol 48:345-366). Furthermore, if Omo L338y-6 was a robust australopithecine, this discovery extends the occurrence of an enlarged O/M sinus system to one of the earliest known paranthropines. Am J Phys Anthropol 110:399-406, 1999.     

Enamel hypoplasia in the deciduous teeth of great apes: variation in prevalence and timing of defects

The prevalence of enamel hypoplasia in the deciduous teeth of great apes has the potential to reveal episodes of physiological stress in early stages of ontogenetic development. However, little is known about enamel defects of deciduous teeth in great apes. Unresolved questions addressed in this study are: Do hypoplastic enamel defects occur with equal frequency in different groups of great apes? Are enamel hypoplasias more prevalent in the deciduous teeth of male or female apes? During what phase of dental development do enamel defects tend to form? And, what part of the dental crown is most commonly affected? To answer these questions, infant and juvenile skulls of two sympatric genera of great apes (Gorilla and Pan) were examined for dental enamel hypoplasias. Specimens from the Powell‐Cotton Museum (Quex Park, UK; n = 107) are reported here, and compared with prior findings based on my examination of juvenile apes at the Cleveland Museum of Natural History (Hamman‐Todd Collection; n = 100) and Smithsonian Institution (National Museum of Natural History; n = 36). All deciduous teeth were examined by the author with a ×10 hand lens, in oblique incandescent light. Defects were classified using Fédération Dentaire International (FDI)/Defects of Dental Enamel (DDE) standards; defect size and location on the tooth crown were measured and marked on dental outline charts. Enamel defects of ape deciduous teeth are most common on the labial surface of canine teeth. While deciduous incisor and molar teeth consistently exhibit similar defects with prevalences of ～10%, canines average between 70–75%. Position of enamel defects on the canine crown was analyzed by dividing it into three zones (apical, middle, and cervical) and calculating defect prevalence by zone. Among gorillas, enamel hypoplasia prevalence increases progressively from the apical zone (low) to the middle zone to the cervical zone (highest), in both maxillary and mandibular canine teeth. Results from all three study collections reveal that among the great apes, gorillas (87–92%) and orangutans (91%) have a significantly higher prevalence of canine enamel defects than chimpanzees (22–48%). Sex differences in canine enamel hypoplasia are small and not statistically significant in any great ape. Factors influencing intergroup variation in prevalence of enamel defects and their distribution on the canine crown, including physiological stress and interspecific dento‐gnathic morphological variation, are evaluated. Am J Phys Anthropol 116:199–208, 2001. © 2001 Wiley‐Liss, Inc.     

Markers of physiological stress in juvenile bonobos (Pan paniscus): Are enamel hypoplasia,skeletal development and tooth size interrelated?

A reduction in enamel thickness due to disrupted amelogenesis is referred to as enamel hypoplasia (EH). Linear EH in permanent teeth is a widely accepted marker of systemic physiological stress. An enigmatic, nonlinear form of EH commonly manifest in great ape and human deciduous canines (dc) is known as localized hypoplasia of primary canines (LHPC). The etiology of LHPC and what it signifies—localized traumatic or systemic physiological stress—remains unclear. This report presents frequency data on LHPC, hypostotic cranial traits, and tooth size in a sample of juvenile bonobos, then tests hypotheses of intertrait association that improve knowledge of the etiology and meaning of LHPC. The fenestration hypothesis is tested using hypostotic cranial traits as a proxy for membrane bone ossification, and the relationship between tooth size, LHPC, and hypostosis is investigated. Macroscopic observations of EH, hypostotic traits, and measurements of buccolingual tooth size were conducted according to established standards. LHPC was found in 51.2% of bonobos (n = 86) and in 26% of dc teeth (n = 269). Hypostotic traits were observed in 55.2% of bonobos (n = 96). A test of the association between LHPC and hypostosis yielded nonsignificant results (χ² = 2.935; P = 0.0867). Primary canines were larger in specimens with LHPC than in unaffected specimens (paired samples t test; udc, P = 0.011; ldc, P = 0.018), a result consistent with the fenestration hypothesis of LHPC pathogenesis. Hypostosis was not associated with differences in tooth size (P > 0.05). LHPC may be an indirect indicator of physiological stress, resulting from large, buccally displaced primary canines. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Incidence and distribution of postcranial fractures in the prehistoric population of San Pedro de Atacama,northern Chile

Trauma incidence analysis in skeletal populations has been very popular among skeletal biologists during the last two decades. In this context, the work of Lovejoy and Heiple ([1981] Am. J. Phys. Anthropol. 55:529–541) has been quoted as a landmark because their analysis rested on a populational approach, avoiding simple assumptions about cause and etiology. In this study, we apply to the prehistoric population of San Pedro de Atacama, northern Chile, an approach similar to that carried out by Lovejoy and Heiple (1981). The results obtained point to a peak of risk of fractures among old people, estimated age around 45 years. The distribution of fractures by sex and age suggests that the prevailing etiology is related to accidents and not violence. When the frequencies of fractures are compared, the Libben population shows a much higher incidence than the Atacamenean population. It is suggested that this difference can be explained by peculiarities of the subsistence economies of the two populations. Am J Phys Anthropol 109:253–258, 1999. © 1999 Wiley-Liss, Inc.     

Hand preferences for coordinated bimanual actions in 777 great apes: implications for the evolution of handedness in hominins

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates.     

Canine tip wear in male and female anthropoids

One component of the “dual selection hypothesis” (Greenfield [1992a] Year. Phys. Anthropol. 35:153–185) is that the tips of female canines are commonly blunted and more frequently so than those of conspecific males. Data derived from two randomly selected age-graded samples of Macaca fascicularis (n = 70) and Colobus badius (n = 59) show that at least 80% of the females exhibit tip blunting on one or both canines and that frequencies of blunting are far greater than those of conspecific males in both jaws. Sexual dimorphism in mandibular canine morphology and wear and other recently critiqued aspects of the “dual selection hypothesis” (Plavcan and Kelley [1996] Am. J. Phys. Anthropol. 99:379–387.) are also discussed. Am J Phys Anthropol 107:87–97. © 1998 Wiley-Liss, Inc.     

Hepatic expression of cytochrome P450 enzymes in non‐human primate species

Cytochromes P450 (P450) largely remain to be characterized in great apes. Comparative immunochemical detection of drug metabolizing forms of P450s 1A, 2A, 2B, 2C, 2D, 2E, 2J, 3A, 4A, and 4F in liver microsomes from chimpanzees, gorillas, orangutans, gibbons, cynomolgus and rhesus macaques, and common marmosets were carried out.     

Test of revised method of age estimation from the auricular surface of the ilium

The objective of this paper is to test a revised method of age estimation based on the morphology of the auricular surface recently proposed by Buckberry and Chamberlain ([2002] Am. J. Phys. Anthropol. 119:231-239). The study sample consists of 309 individuals of known sex, age, and race from the Terry and Huntington Collections. Auricular surfaces were scored using the revised technique to determine whether it is equally applicable to both sexes as well as blacks and whites. The auricular surfaces of the same individuals were also scored using the original method of auricular surface scoring developed by Lovejoy et al. ([1985] Am. J. Phys. Anthropol. 68:15-28) to determine whether the revised technique is comparable to the original method in terms of accuracy. Results show that the revised method is equally applicable to males and females as well as blacks and whites. The revised method is less accurate than the original method for individuals between 20-49 years of age, but more accurate for individuals between 50-69 years of age.     

Interactions between zoo‐housed great apes and local wildlife

Although there are published reports of wild chimpanzees, bonobos, and orangutans hunting and consuming vertebrate prey, data pertaining to captive apes remain sparse. In this survey‐based study, we evaluate the prevalence and nature of interactions between captive great apes and various indigenous wildlife species that range into their enclosures in North America. Our hypotheses were threefold: (a) facilities housing chimpanzees will report the most frequent and most aggressive interactions with local wildlife; (b) facilities housing orangutans and bonobos will report intermediate frequencies of these interactions with low levels of aggression and killing; and (c) facilities housing gorillas will report the lowest frequency of interactions and no reports of killing local wildlife. Chimpanzees and bonobos demonstrated the most aggressive behavior toward wildlife, which matched our predictions for chimpanzees, but not bonobos. This fits well with expectations for chimpanzees based on their natural history of hunting and consuming prey in wild settings, and also supports new field data on bonobos. Captive gorillas and orangutans were reported to be much less likely to chase, catch and kill wildlife than chimpanzees and bonobos. Gorillas were the least likely to engage in aggressive interactions with local wildlife, matching our predictions based on natural history. However unlike wild gorillas, captive gorillas were reported to kill (and in one case, eat) local wildlife. These results suggest that some behavioral patterns seen in captive groups of apes may be useful for modeling corresponding activities in the wild that may not be as easily observed and quantified. Furthermore, the data highlight the potential for disease transmission in some captive settings, and we outline the associated implications for ape health and safety. Am. J. Primatol. 71:458–465, 2009. © 2009 Wiley‐Liss, Inc.     

Skull of early Eocene Cantius abditus (primates: Adapiformes) and its phylogenetic implications,with a reevaluation of “Hesperolemur” actius

A substantially complete skull and mandible of the primitive adapiform Cantius is reported from the Early Eocene Willwood Formation of Wyoming. The mandible contains an almost complete lower dentition in which the lower incisors are strongly inclined and have spatulate crowns, I₂ is larger than I₁, and the canine is large and projecting. The cranium shares many features with those of Notharctus and Smilodectes but differs in having nasals that broaden proximally. Presence of a prominent canine and strong sagittal crest may indicate that it represents a male. The basicranium preserves auditory structures almost identical to those in extant noncheirogaleid lemurs, including a large bony tube for the stapedial artery and a small, open sulcus for the distal portion of the promontorial artery. The dentition is sufficiently primitive to be compatible with a relationship to either strepsirrhines or anthropoids, but the anatomy of the auditory region is more consistent with either specific relationship to lemurs or, more likely, a basal position that approximates the euprimate morphotype. Certain features of the basicranium of “Hesperolemur” actius, described by Gunnell ([1995] Am. J. Phys. Anthropol. 98:447–470) as being unlike that of any other adapiform, were either misinterpreted or are apparently no longer present in the holotype. Reassessment of these and other features indicates that in fact “H.” actius differs little from Cantius and should not be separated from the latter at the genus level, although on dental grounds the species appears to be distinct (as C. actius). Am J Phys Anthropol 109:523–539, 1999. © 1999 Wiley-Liss, Inc.     

Variation in perikymata counts between repetitive episodes of linear enamel hypoplasia among orangutans from Sumatra and Borneo

The goal of this study is to evaluate whether repetitive linear enamel hypoplasia (rLEH) in apes is ecologically informative. LEH, which appears as grooves of thinner enamel often caused by malnutrition and/or disease, is a permanent record of departures from developmental homeostasis in infant and juvenile apes. Orangutans were selected for the study as they are a threatened species, have a remarkably high prevalence of rLEH, and because Sumatra is deemed a better habitat for orangutans than is Borneo, facilitating an ecological comparison. Objectives are to determine: a) whether periodicity of rLEH in orangutans corresponds to monsoon‐mediated cycles in precipitation or food; and b) whether patterning of rLEH supports the view that Borneo is an inferior habitat. This study compares the counts of perikymata between adjacent LEH from 9 Sumatran and 26 Bornean orangutans to estimate the periodicity of rLEH. A total of 131 nonredundant inter‐LEH perikymata counts were transformed to natural log values to reveal clusters of counts in a multiplicative series. Using a value of 10 days to form one perikyma, rLEH tends to recur semiannually in both populations. However, Sumatran orangutans show significantly fewer semiannual intervals and more annually recurring episodes. Bornean orangutans show mostly semiannual intervals and are more variable in inter‐LEH perikymata counts. It is concluded that: a) developmental conditions for infant orangutans in Sumatra protect them somewhat from seasonal and environmental variation; b) temporal patterning of rLEH indicates that Borneo is the poorer habitat for orangutans; and c) the study of rLEH can be ecologically informative. Am J Phys Anthropol 154:125–139, 2014. © 2014 Wiley Periodicals, Inc.     

Foraging and ranging behavior during a fallback episode: Hylobates albibarbis and Pongo pygmaeus wurmbii compared

Periodic episodes of food scarcity may highlight the adaptive value of certain anatomical traits, particularly those that facilitate the acquisition and digestion of exigent fallback foods. To better understand the selective pressures that favored the distinctive dental and locomotor morphologies of gibbons and orangutans, we examined the foraging and ranging behavior of sympatric Hylobates albibarbis and Pongo pygmaeus wurmbii during an episode of low fruit availability at Tuanan, Kalimantan Tengah, Indonesia. We found that Hylobates ranged 0.5 km day⁻¹ or 33% farther than did Pongo, but the overall daily ranging of both species did not vary as fruit availability decreased by as much as 50%. Among gibbons, we observed dietary switching to fallback foods; in particular, there was a progressively greater reliance on figs, liana products, and unripe fruit. Orangutans relied heavily on unripe fruit and fracture-resistant bark and pith tissues. Despite these divergent fallback patterns, the stiffness of fruit mesocarp consumed by Hylobates and Pongo did not differ. We discuss canine and molar functional morphology with respect to dietary mechanics. Next, to contextualize these results, we discuss our findings with respect to forest structure. The rain forests of Southeast Asia have been described as having open, discontinuous canopies. Such a structure may inform our understanding of the ranging behavior and distinctive locomotion of apes in the region, namely richochetal brachiation and quadrumanous clambering. Our approach of integrating behavioral ecology with physical measures of food may be a powerful tool for understanding the functional adaptations of primates. Am J Phys Anthropol 140:716–726, 2009. © 2009 Wiley-Liss, Inc.     

Quadrupedal locomotion in squirrel monkeys (Cebidae: Saimiri sciureus): a cineradiographic study of limb kinematics and related substrate reaction forces

Quadrupedal locomotion of squirrel monkeys on small-diameter support was analyzed kinematically and kinetically to specify the timing between limb movements and substrate reaction forces. Limb kinematics was studied cineradiographically, and substrate reaction forces were synchronously recorded. Squirrel monkeys resemble most other quadrupedal primates in that they utilize a diagonal sequence/diagonal couplets gait when walking on small branches. This gait pattern and the ratio between limb length and trunk length influence limb kinematics. Proximal pivots of forelimbs and hindlimbs are on the same horizontal plane, thus giving both limbs the same functional length. However, the hindlimbs are anatomically longer than the forelimbs. Therefore, hindlimb joints must be more strongly flexed during the step cycle compared to the forelimb joints. Because the timing of ipsilateral limb movements prevents an increasing amount of forelimb retraction, the forelimb must be more protracted during the initial stance phase. At this posture, gravity acts with long moment arms at proximal forelimb joints. Squirrel monkeys support most of their weight on their hindlimbs. The timing of limb movements and substrate reaction forces shows that the shift of support to the hindlimbs is mainly done to reduce the compressive load on the forelimb. The hypothesis of the posterior weight shift as a dynamic strategy to reduce load on forelimbs, proposed by Reynolds ([1985]) Am. J. Phys. Anthropol. 67:335-349; [1985] Am. J. Phys. Anthropol. 67:351-362), is supported by the high correlation of ratios between forelimb and hindlimb peak vertical forces and the range of motion of shoulder joint and scapula in primates.     

Heel contact as a function of substrate type and speed in primates

In this report we provide detailed data on the patterns and frequency of heel contact with terrestrial and arboreal supports in primates. These data can help resolve the question of whether African apes and humans are uniquely “plantigrade” (Gebo [1992] Am. J. Phys. Anthropol. 89:29–58; Gebo [1993a] Am. J. Phys. Anthropol. 97:382–385; Gebo [1993b] Postcranial Adaptation in Nonhuman Primates), or if plantigrady is common in other primates (Meldrum [1993] Am. J. Phys. Anthropol. 91:379–381). Using biplanar and uniplanar videotapes, we recorded the frequency and timing of heel contact for a variety of primates (32 species) walking on the ground and on simulated arboreal supports at a range of natural speeds. Our results indicate that Pongo as well as the African apes exhibit a “heel-strike” at the end of swing phase. Ateles and Hylobates make heel contact on all supports shortly after mid-foot contact, although spider monkeys do so only at slow or moderate speeds. Data available from uniplanar videotapes suggest that this pattern occurs in Alouatta and Lagothrix as well. No other New or Old World monkey or prosimian in this study made heel contact during quadrupedalism on any substrate. Thus, heel contact occurs in all apes and atelines, but only the great apes exhibit a heel-strike. We suggest that heel contact with the substrate is a by-product of an active posterior weight-shift mechanism involving highly protracted hindlimbs at touchdown. Force plate studies indicate that this mechanism is most extreme in arboreally adapted primate quadrupeds walking on arboreal supports. Although heel contact and heel-strike may have no evolutionary link, it is possible that both patterns are the result of a similar weight shift mechanism. Therefore, the regular occurrence of heel contact in a variety of arboreal primates, and the absence of a true biomechanical link between limb elongation, heel contact, and terrestriality, calls into question the claim that hominid foot posture was necessarily derived from a quadrupedal terrestrial ancestor. © 1995 Wiley-Liss, Inc.