Cortical surface patterns in human and nonhuman primates

An analysis of skulls from several primate species shows that a “worm-track” surface pattern, first identified in the brow region in fossil adult hominids and subsequently in olive baboons, chimpanzees, and macaques, is also present in numerous other species. Fine cancellous bone and its attendant vermiculate surface pattern have been observed in subadult and adult gelada baboons, gibbons, gorillas, and orangutans as well as in modern Homo sapiens and several Plio-Pleistocene fossil hominids. In contemporary primates, fine cancellous bone has been identified not only in the brow region, but also along the zygomatic arch, on the pterygoid plates, on the maxilla, along the temporal line, on the mastoid process, and in the region of inion. Given the widespread distribution of this trait, caution is advised when using it as a diagnostic indicator of the evolutionary or functional significance of craniofacial morphology.     

The bony labyrinth of Neanderthals

This paper presents a comprehensive comparative analysis of the Neanderthal bony labyrinth, a structure located inside the petrous temporal bone. Fifteen Neanderthal specimens are compared with a Holocene human sample, as well as with a small number of European Middle Pleistocene hominins, and early anatomically modern and European Upper Palaeolithic humans. Compared with Holocene humans the bony labyrinth of Neanderthals can be characterized by an anterior semicircular canal arc which is smaller in absolute and relative size, is relatively narrow, and shows more torsion. The posterior semicircular canal arc is smaller in absolute and relative size as well, it is more circular in shape, and is positioned more inferiorly relative to the lateral canal plane. The lateral semicircular canal arc is absolutely and relatively larger. Finally, the Neanderthal ampullar line is more vertically inclined relative to the planar orientation of the lateral canal. The European Upper Palaeolithic and early modern humans are most similar, although not fully identical to Holocene humans in labyrinthine morphology. The European Middle Pleistocene hominins show the typical semicircular canal morphology of Neanderthals, with the exception of the arc shape and inferiorly position of the posterior canal and the strongly inclined ampullar line. The marked difference between the labyrinths of Neanderthals and modern humans can be used to assess the phylogenetic affinities of fragmentary temporal bone fossils. However, this application is limited by a degree of overlap between the morphologies. The typical shape of the Neanderthal labyrinth appears to mirror aspects of the surrounding petrous pyramid, and both may follow from the phylogenetic impact of Neanderthal brain morphology moulding the shape of the posterior cranial fossa. The functionally important arc sizes of the Neanderthal semicircular canals may reflect a pattern of head movements different from that of modern humans, possibly related to aspects of locomotor behaviour and the kinematic properties of their head and neck.     

Krapina, “Classic” Neanderthals,and the evolution of the European face

Except for the front end of the dental arch, tooth size remained at approximately the same level throughout the Middle Pleistocene. The Krapina Neanderthals at the end of the last interglacial differed from Homo erectus only in having larger front teeth. From that time on, tooth size in populations at the northern edge of the area of human occupation in the Old World has reduced approximately in proportion to the time elapsed. The “Classic” Neanderthals of western Europe, in fact, have teeth that are 15% smaller than those of the earlier Krapina Neanderthals and only 5% larger than the early Upper Palaeolithic. Reduction since the early Upper Palaeolithic has proceeded another full 20%. It is suggested that the development of heated stone cooking in the Mousterian, originally for the purpose of thawing frozen food, reduced the forces of selection that had previously maintained tooth size during the Middle Pleistocene. The operation of the Probable Mutation Effect, then produced the observed reductions.     

Masticatory-stress hypotheses and the supraorbital region of primates

The purpose of this study is to test various masticatory-stress hypotheses about the evolution and function of well-developed browridges of higher primates. This was done by measuring and analyzing patterns of in vivo bone strain recorded from three-element rosette strain gages bonded to the supraorbital region and to other portions of the bony face of Macaca fascicularis and Papio anubis during mastication and incision. The magnitude and direction of the principal strains recorded support Endo's hypothesis that the supraorbital region during mastication and incision is bent in the frontal plane (Endo, 1966). Our data do not, however, support his hypothesis that the supraorbital region is bent more during incision than during mastication. The data also demonstrate that overall levels of supraorbital strain are not larger in more prognathic subjects. Most importantly, the data indicate that the supraorbital region of nonhuman catarrhines is strained very little during mastication and incision. This indicates that there is much more supraorbital bone than is necessary both to counter masticatory loads and to provide an adequate safety factor to failure for these loads. This in turn suggests that the macaque and baboon browridges can be considerably reduced in size and still maintain these required structural characteristics. Thus, our experiments provide no support whatsoever for those hypotheses that directly link browridge morphology to masticatory stress (cf. Endo, 1966; Russell, 1983, 1985). A recent review of Endo's original work indicates that this latter statement is also true for humans (Picq and Hylander, 1989). We conclude, therefore, that there is no good reason to believe that enlarged browridges in living and/or fossil primates are structural adaptations to counter intense masticatory forces. The evolution of browridge morphology in primates is best explained on the basis of factors related to the position of the brain relative to the orbits (Moss and Young, 1960). When these structures are widely separated, as in gorillas, the large intervening space must be bridged with bone. In addition, enough bone must be present within the supraorbital and bridged regions to prevent structural failure due to non-masticatory external forces associated with highly active primates (e.g., accidental traumatic forces applied to the orbits and neurocranium). This requirement results in both pronounced browridges and in much more supraorbital bone than is necessary to counter routine cyclical stress during mastication and incision. This in turn explains why bone strains recorded from the supraorbital region are extremely small relative to other portions of the primate face during mastication and incision.     

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.     

Neocortical development and social structure in primates

The relationships between the relative size of the neocortex and differences in social structures were examined in prosimians and anthropoids. The relative size of the neocortex (RSN) of a given congeneric group in each superfamily of primates was measured based on the allometric relationships between neocortical volume and brain weight for each superfamily, to control phylogenetic affinity and the effects of brain size. In prosimians, “troop-making” congeneric groups (N=3) revealed a significantly larger RSN than solitary groups (N=6), and there was a significant, positive correlation between RSN and troop size. In the case of anthropoids, polygynous/frugivorous groups (N=5) revealed a significantly larger RSN than monogynous/frugivorous groups (N=8). Furthermore, a significant, positive correlation between RSN and troop size was found for frugivorous congeneric groups of the Ceboidea. These results suggest that neocortical development is associated with differences in social structure among primates.     

The dentinoenamel junction and the hypocone in primates

Enamel has been stripped from primate teeth (especially humans and ceboids) with special reference to the comparative form of the hypocone. Dentally reduced species show variable developments not always expected of a hypothetical ontogenetically prior stage.     

Paternity exclusion in primates: Two strategies

Two strategies for the use of polymorphic biochemical and serological markers in paternity testing problems in non-human primate groups, where pedigree information is incomplete, are discussed. The positive approach, of attempting to prove paternity, is shown to be impracticable given the levels of detectable genetic variation among primates. The more conventional approach of paternity exclusion is examined and found to be useful under certain conditions. This approach is illustrated using the published data on the levels of biochemical and serological variation in Macaca nemestrina.     

Human Behavioral Organization in the Middle Paleolithic: Were Neanderthals Different?

Interwoven with the debate regarding the biologic replacement of Neanderthals by modern humans is the question of the degree to which Neanderthals and modern foragers differed behaviorally. We consider this question through a detailed spatial analysis of artifacts and related evidence from stratified living floors within a 49–69 k.y.a. rock shelter site, Tor Faraj, in southern Jordan. The study involves a critical evaluation of living floors, the identification of site structure, and the decoding of the site structure in an effort to understand how the inhabitants of the shelter organized their behaviors. The site structure of Tor Faraj is also compared to occupations of modern foragers in ethnographic and archaeological contexts. When the information from the excavation of Tor Faraj is considered with evidence from other late Middle Paleolithic sites, there seems to be little basis for the claims that constraints in the behavioral organization of Neanderthals led to their replacement by modern foragers.     

Femoral curvature in Neanderthals and modern humans: a 3D geometric morphometric analysis

Since their discovery, Neanderthals have been described as having a marked degree of anteroposterior curvature of the femoral shaft. Although initially believed to be pathological, subsequent discoveries of Neanderthal remains lead femoral curvature to be considered as a derived Neanderthal feature. A recent study on Neanderthals and middle and early Upper Palaeolithic modern humans found no differences in femoral curvature, but did not consider size-corrected curvature. Therefore, the objectives of this study were to use 3D morphometric landmark and semi-landmark analysis to quantify relative femoral curvature in Neanderthals, Upper Palaeolithic and recent modern humans, and to compare adult bone curvature as part of the overall femoral morphology among these populations.Comparisons among populations were made using geometric morphometrics (3D landmarks) and standard multivariate methods. Comparative material involved all available complete femora from Neanderthal and Upper Palaeolithic modern human, archaeological (Mesolithic, Neolithic, Medieval) and recent human populations representing a wide geographical and lifestyle range. There are significant differences in the anatomy of the femur between Neanderthals and modern humans. Neanderthals have more curved femora than modern humans. Early modern humans are most similar to recent modern humans in their anatomy. Femoral curvature is a good indicator of activity level and habitual loading of the lower limb, indicating higher activity levels in Neanderthals than modern humans. These differences contradict robusticity studies and the archaeological record, and would suggest that femoral morphology, and curvature in particular, in Neanderthals may not be explained by adult behavior alone and could be the result of genetic drift, natural selection or differences in behavior during ontogeny.     

Variation in protein structure and function: Primate hemoglobins

Summary Variation in structure among primate hemoglobins is associated with variation in function. This supports the hypothesis that most substitutions observed among homologous proteins in different species have been fixed by natural selection because they contribute to the fitness of the genotype. It does not support the concept that most substitutions result from the fixation of neutral alleles by genetic drift.     

Multivariate dental allometry in primates

Disagreement is current over the question of whether relatively large teeth in some large primates are a natural outcome of growth trends instead of an indication of intrinsic differences. A cross-primate survey of dental scaling relative to skull (and inferred body) size is given in this study, using a principal component technique to measure the multivariate growth relation between two sets of data: dental size and cranial size. Cheek teeth are strongly positively allometric in restriced taxonomic groups, especially in cercopithecoids. Conversely, the allometry drops to an almost linear proportional growth relation when variation in diet is controlled.     

Postcanine tooth size in female primates

The results of many allometric studies of postcanine tooth size in mammals have not corresponded to expectations of tooth size based on energy requirements and dental function. The purpose of this study is to investigate the relationship between postcanine occlusal surface area, body size, and the metabolic demands of pregnancy and lactation in female primates. Tooth and body sizes from 38 primate species were taken from the literature to test two hypotheses: 1) females should have relatively larger teeth than males in order to masticate additional food for the energetic costs of reproduction; 2) taxa with the largest neonatal size (a measure of average metabolic costs of pregnancy and lactation) should have females with a greater degree of relative dental enlargement. The results show that relatively large female teeth are not found consistently in primate species. Females have less occlusal surface area than expected on the basis of the male tooth and body size regression in 21% of the species, and there is no correlation between relative female tooth size and relative newborn size across higher primate taxa. The degree of female dental enlargement is most closely related to degree of sexual dimorphism in body weight. The correlation between degree of body weight dimorphism and relatively larger postcanine teeth in females than in males is 0.87 in the 38 species. Species that are monomorphic in weight tend to be monomorphic in tooth size even though females apparently require more food than males.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sexual dimorphism and dental variability in platyrrhine primates

Leutenegger and Cheverud (1982, 1985) propose a hypothesis to explain why larger primates are more sexually dimorphic in body weight and canine size. Their hypothesis states that any factor selecting for an evolutionary increase in body size will produce an increase in sexual dimorphism in any character if either heritability or phenotypic variability is greater in males than in females for that character. They cite no evidence for heritability but give some data to suggest that males are, in fact, more variable than females. We test the latter proposition more fully using measurements on the dentitions of platyrrhine primates. Male and female phenotypic variances are not significantly different in most cases. Cases of greater male phenotypic variance are not limited to sexually dimorphic species. We conclude that the hypothesis of Leutenegger and Cheverud does not explain the observed patterns of dental sexual dimorphism, at least in platyrrhines.     

Twinning frequency in catarrhine primates

It has been repeatedly suggested that twinning frequency in most catarrhine primates is approximately the same as in humans, whereas the frequency in the chimpanzee and the gorilla might be higher. This study presents a re-evaluation of the evidence from the pertinent literature. It can be demonstrated that most data on twinning frequency in Old World monkeys and apes should not be used because of their small sample size. A lower limit of 1500 pregnancies is suggested here. If all frequency estimates taken from smaller samples are rejected, only four estimates forMacaca mulatta andPapio hamadryas remain. The estimates range from 0.19 to 0.35% and are in fact lower than the frequencies of most (but necessarily all) human populations. The published birth samples for apes are, however, relatively small, and the resulting twinning rates may not be reliable. A republication, occasioned by publisher's errors, of this article that originally appeared in Human Evolution 2: 547–555 (1987). It is recommended that all references to this article be cited from the present publication.     

Twinning frequency in catarrhine primates

It has been repeatedly suggested that twinning frequency in most catarrhine primates is approximately the same as in humans, whereas the frequency in the chimpanzee and the gorilla might be higher. This study presents a re-evaluation of the evidence from the pertinent literature. It can be demonstrated that most data on twinning frequency in Old World monkeys and apes should not be used because of their small sample size. A lower limit of 1500 pregnancies is suggested here. If all frequency estimates taken from smaller samples are rejected, only four estimates forMacaca mulatta andPapio hamadryas remain. The estimates range from 0.19 to 0.35% and are in fact lower than the frequencies of most (but necessarily all) human populations. The published birth samples for apes are, however, relatively small, and the resulting twinning rates may not be reliable.     

Immunoglobulin allotypes (Gm,Am, and Km) in non-human primates

The immunoglobulin (Ig) allotypes (Gm, Am, and Km systems) are the genetic markers of the human IgG1, IgG2, IgG3(Gm), IgA2(Am), and kappa light chain(Km). The Gm system, with 18 allotypes shows the greatest degree of polymorphism and we define two Am and three Km allotypes. In this review, we report all the results observed in non-human primates belonging to Hominoidea, Cercopithecoidea, Ceboidea, Lorisoidea, Lemuroidea, and Tupaoidea superfamilies (72 species and subspecies). They concern published data and new unpublished ones. The distribution of the human allotypes and their localization are reported, as well as discordant results observed in some cases with anti-allotype reagents of the same specificity (human and animal origin). Some allotypes are restricted to man. Hominoidea have the greatest number of Gm allotypes and the richest polymorphism. Relatively few allotypes have been found in Cercopithecoidea and Prosimians; most Platyrrhinian species have no allotype. The epitopic polymorphism has been interpreted in terms of evolution of Ig allotypes from primate to man and of the phylogenetic relationships of non-human primate species.     

Canine size and shape in male anthropoid primates

While studies of canine dimorphism in primates are common, only a few have examined canine tooth size independently within each sex. Recently, Greenfield and Washburn (Am. J. Phys. Anthropol. 84:17–34, 1991) proposed that there are two types of male canines which reflect different allometric scaling patterns of canine crown height against canine occlusal dimensions. They also suggest that proportional canine shape, measured as canine crown height (or projection) relative to the occlusal dimensions, provides an estimate of the canine's function as a weapon, though they provide no test of this hypothesis. This analysis critically examines the claim that there are two types of male canines among anthropoids. It then tests the hypothesis that relative male canine size (measured against body weight) and proportional canine shape are related to estimates of intermale competition, diet, and substrate (used as a surrogate measure of predation pressure). While there is strong taxonomic variation in canine size and shape among male anthropoids, no evidence is found for two discrete canine types. Rather, within families and subfamilies, canine dimensions scale isometrically against body weight and against each other, with a continuum of canine shapes among different taxa. While variation in male canine size is associated with intermale competition and substrate, even when taxonomic variation is controlled, variation in proportional canine shape is not. Neither canine size nor shape are generally associated with variation in diet. © 1993 Wiley-Liss, Inc.