Testicular Size,Mating System,and Maturation Schedules in Wild Anubis and Hamadryas Baboons

We report body mass and testicular size in 258 anubis (Papio anubis or P. hamadryas anubis) and 59 hamadryas (P. hamadryas or P. h. hamadryas) baboons, live-trapped in Ethiopia. As predicted by theories of sexual selection by sperm competition, among hamadryas baboons, which are monandrous, fully adult males have absolutely and relatively smaller testes than those of comparable males among anubis baboons, which are polyandrous. Male hamadryas are also ca. 10% smaller in bodily mass as adults. The intertaxonal difference in adults is due entirely to the fact that in male anubis baboons, testicular and bodily mass continue to grow up to full adulthood–the age at which most males emigrate from their natal troop and initiate a confrontational breeding strategy among unrelated animals. By contrast, male hamadryas baboons, which are usually philopatric, attain adult body mass and testicular size as subadults. In both species, juveniles experience rapid testicular growth peaking in rate at ca. 12kg body mass, but testicular descent and growth starts earlier in hamadryas than in anubis baboons. Juvenile hamadryas baboons have relatively larger testes than their anubis equivalents, perhaps because male philopatry allows the mating strategy of male hamadryas baboons to be initiated during juvenile life and therefore permits some sperm competition between juveniles and adults.     

Behavioral variation and reproductive success of male baboons (Papio anubis x Papio hamadryas) in a hybrid social group

We take advantage of an array of hybrid baboons (Papio anubis x Papio hamadryas) living in the same social group to explore the causes and consequences of different male mating strategies. Male hamadryas hold one-male units and exhibit a sustained, intense interest in adult females, regardless of the latter's reproductive state. Anubis baboons, by contrast, live in multi-male, multi-female groups where males compete for females only when the latter are estrous. These two taxa interbreed to form a hybrid zone in the Awash National Park, Ethiopia, where previous work has suggested that hybrid males have intermediate and ineffective behavior. Here, we first examine male mating strategies with respect to morphological and genetic measures of ancestry. We found significant relationships between behavioral measures and morphology; males with more hamadryas-like morphology had more hamadryas-like behavior. However, genetic ancestry was not related to behavior, and in both cases intermediates displayed a previously unreported level of behavioral variation. Furthermore, male behavior was unrelated to natal group. Second, we evaluated reproductive success by microsatellite-based paternity testing. The highest reproductive success was found for individuals exhibiting intermediate behaviors. Moreover, over nine years, some genetically and morphologically intermediate males had high reproductive success. We conclude that the behavior of hybrid males is therefore unlikely to be an absolute barrier to admixture in the region.     

Sociological study of a wild group of hybrid baboons betweenPapio anubis andP. hamadryas in the Awash Valley,Ethiopia

A group of hybrid baboons between anubis and hamadryas named the Gorge group was studied for five months in the Awash Valley, Ethiopia. Both morphologically and genetically, anubis features were prevalent in the Gorge group. Three types of subgroups within the group were distinguished: one-male groups, pair groups, and multi-male groups. The joining and parting of subgroups is examined by cluster-analysis. The Gorge group is concluded to be a single social unit, though it was composed of two main clusters. Grooming relations, spacing mechanisms, and sexual relations among its members were analyzed in detail. The dominance relationships among males are clarified in order to be related with their social interactions. Various aspects of the social organization are compared with those of anubis or hamadryas. It is demonstrated that the social structure of the Gorge group showed an intricate mixture of anubis and hamadryas characteristics. It is discussed how such a structure had been formed through hybridization, and what were the factors causing the differences among three types of subgroups.     

Migration of male hamadryas baboons into anubis groups in the Awash National Park,Ethiopia

Among “savanna” baboons, males are the dispersing sex, and females are philopatric. Despite clear evidence for migration of adult males at Erer-Gota, Ethiopia (Abegglen, 1984), it is generally believed that a different pattern-dispersal only by female transfer-is found in hamadryas baboons,Papio hamadryas hamadryas (Pusey and Packer, 1987; Pusey, 1988; Stammbach, 1987). Since the late 1960's, there have been isolated observations of hamadryas males migrating into anubis groups in the Awash National Park, Ethiopia (Nagel, 1973; Kawai and Sugawara, 1976; Sugawara, 1982). Since 1983, we have observed 11 individually identified adult hamadryas immigrants in four anubis groups above the Awash Falls and have trapped and tagged 9 of them. One subadult male was also captured and marked. Repeated visits to the study site allow us to document long-term residence of these “cross-migrant” males in their host groups. The longest-resident male has been in the same group for 5 years or more; a conservative estimate of the average length of residence is 3 years. We estimate that 25 hamadryas males have moved into this ozne over the last 15 years. Although larger than the hamadryas males captured in 1973, all but one of our cross-migrants appear phenotypically hamadryas. By comparing, the ages of our cross-migrants with Abegglen's account of the typical hamadryas male life-history, we have found that the adult hamadryas males seem to immigrate at ages consistent with having left their     

High Prevalence of Simian T-Lymphotropic Virus Type L in Wild Ethiopian Baboons

Simian T-cell leukemia viruses (STLVs) are the simian counterparts of human T-cell leukemia viruses (HTLVs). A novel, divergent type of STLV (STLV-L) from captive baboons was reported in 1994, but its natural prevalence remained unclear. We investigated the prevalence of STLV-L in 519 blood samples from wild-living nonhuman primates in Ethiopia. Seropositive monkeys having cross-reactive antibodies against HTLV were found among 22 out of 40 hamadryas baboons, 8 of 96 anubis baboons, 24 of 50 baboons that are hybrids between hamadryas and anubis baboons, and 41 of 177 grivet monkeys, but not in 156 gelada baboons. A Western blotting assay showed that sera obtained from seropositive hamadryas and hybrid baboons exhibited STLV-L-like reactivity. A PCR assay successfully amplified STLV sequences, which were subsequently sequenced and confirmed as being closely related to STLV-L. Surprisingly, further PCR showed that nearly half of the hamadryas (20 out of 40) and hybrid (19 out of 50) baboons had STLV-L DNA sequences. In contrast, most of the seropositive anubis baboons and grivet monkeys carried typical STLV-1 but not STLV-L. These observations demonstrate that STLV-L naturally prevails among hamadryas and hybrid baboons at significantly high rates. STLV-1 and -2, the close relative of STLV-L, are believed to have jumped across simian-human barriers, which resulted in widespread infection of HTLV-1 and -2. Further studies are required to know if STLV-L is spreading into human populations.     

Age at maturity in wild baboons: genetic, environmental and demographic influences

The timing of early life-history events, such as sexual maturation and first reproduction, can greatly influence variation in individual fitness. In this study, we analysed possible sources of variation underlying different measures of age at social and physical maturation in wild baboons in the Amboseli basin, Kenya. The Amboseli baboons are a natural population primarily comprised of yellow baboons (Papio cynocephalus) that occasionally hybridize with anubis baboons (Papio anubis) from outside the basin. We found that males and females differed in the extent to which various factors influenced their maturation. Surprisingly, we found that male maturation was most strongly related to the proportion of anubis ancestry revealed by their microsatellite genotypes: hybrid males matured earlier than yellow males. In contrast, although hybrid females reached menarche slightly earlier than yellow females, maternal rank and the presence of maternal relatives had the largest effects on female maturation, followed by more modest effects of group size and rainfall. Our results indicate that a complex combination of demographic, genetic, environmental, and maternal effects contribute to variation in the timing of these life-history milestones.     

Characteristics of hamadryas-like male baboons living in anubis baboon troops in the Awash hybrid zone, Ethiopia.

In the twenty-two years since the Awash baboon hybrid zone was first described, about 25-30 hamadryas or hamadryas-like hybrid males have taken up residence in anubis baboon groups. Most enter as mature adults, though some may immigrate as juvenile followers. Long-term tracking of known immigrants indicates that they may reside for four years or more in their adopted troop, during which time they establish a series of comparatively short-term harems, and probably sire hybrid offspring. "Abductions" of anubis females to hamadryas troops, as postulated by early observers, were not seen, though their occurrence cannot be ruled out. Younger, more recent immigrants tend to be "purer" hamadryas in phenotype, perhaps reflecting a more distant origin, or a change in the composition of neighboring hamadryas populations. Coexisting hamadryas and anubis show no signs of reproductive isolation by behavioral barriers, and evidently belong to a single "recognition species."     

Social System of a Hybrid Baboon Group (Papio anubis × P. hamadryas)

We describe the social organization, mating system, and social structure of a group of hybrid baboons (Papio anubis×P. hamadryas) in Ethiopia's Awash National Park. The group contained elements of both hamadryas and anubis societies. Overall, the group was a multimale, multifemale group that lacked cohesion and frequently formed subgroups. Subgroup formation was more strongly associated with predation risk than food availability. Although there were several hamadryas-like one-male units OMUs within the group, there was no evidence of a hamadryas multilevel society. Male and female members of OMUs were phenotypically more hamadryas-like than non-OMU individuals. The group contained substantial variation in the strength of inter- and intrasexual bonds: some females primarily groomed males while other females primarily groomed females, and the patterns were consistent with the OMU substructure. Despite some promiscuous mating, mating was biased towards the hamadryas condition for all group members. Additionally, rates of immigration and emigration were very low, and mean pairwise relatedness within the group is rising. For measures of intersexual bonding, all members of the group were intermediate between anubis and hamadryas individuals in less hybridized groups. The group was phenotypically and behaviorally more intermediate than it was in the 1970s (Sugawara, K. (1988). Primates 29: 429–448.) and the changes may indicate a relatively young and dynamic hybrid zone.     

Affiliation Among Females in Wild Hamadryas Baboons (Papio hamadryas hamadryas)

Previous researchers of hamadryas baboons have described a star-shaped sociogram, whereby the strongest social bonds within hamadryas one-male units are between a leader male and his females and bonds among females are weak by comparison. This type of social organization is also known as cross-bonding to distinguish it from the female-bonding found in most papionin monkeys. Models of female primate socioecology suggest that hamadryas baboons lack female bonding due to their reliance on scarce, widely-dispersed food resources. Here, I report observational data from a wild population of hamadryas baboons in Ethiopia indicating that, while females varied widely in their frequency of social interaction with other females, most females spent about as much social time with other females as they did with the leader male and some females even crossed unit boundaries to interact with one another. The size of a unit was positively correlated with the tendency of its females to interact with other females and was negatively correlated with the tendency of its females to interact with the leader male. Females were equally likely to spend social time with other females whether or not the leader male was available for social interaction at the time. Overall, this study suggests that a star-shaped sociogram does not characterize all hamadryas baboons and that female hamadryas may be, to some extent, female-bonded as well as cross-bonded. The lack of more pronounced female bonding in hamadryas is probably due to the behavior of males rather than to ecological factors.     

Hepatocystis in populations of baboons (Papio hamadryas s.l.) of Tanzania and Ethiopia

Bloodsmears from 119 yellow baboons (Papio hamadryas cynocephalus) in six groups in Mikumi National Park, Tanzania, and from 189 olive (P.h. anubis) hamadryas (P.h. hamadryas) and hybrid baboons in three groups in Awash National park, Ethiopia showed differing levels of infection with Hepatocystis simiae. 0% to 42% in the Tanzanian groups were carriers of the parasite but none of the Ethiopian baboons was positive. The remarkable absence of H. simiae in the Ethiopian baboons is believed to be related to environmental factors that exclude transmission in the absence of the Culicoides vector.     

Fetal and infant head circumference sexual dimorphism in primates

Studies have shown that after controlling for the effects of body size on brain size, the brains of adult humans, rhesus monkeys, and chimpanzees differ in relative size, where males have a greater volume of cerebral tissue than females. We assess whether head circumference sexual dimorphism is present during early development by evaluating sex differences in relative head circumference in living fetuses and infants within the first year of life. Head circumference is used as a proxy for brain size in the fetus and infant. Femur length is used as a proxy for body length in the fetus. Ultrasonography was used to obtain fetal measures, and anthropometry was used to obtain postnatal measures in humans, rhesus monkeys, baboons, and common marmosets. We show that statistically significant but low levels of head circumference sexual dimorphism are present in humans, rhesus monkeys, and baboons in early life. On average, males have head circumferences about 2% larger than females of comparable femur/body length in humans, rhesus monkeys, and baboons. No evidence for head circumference sexual dimorphism in the common marmoset was found. Dimorphism was present across all body size ranges. We suggest that head circumference sexual dimorphism emerges largely postnatally and increases throughout maturation, particularly in humans who reach adult dimorphism values greater than the monkeys. We suggest that brain dimorphism is not likely to impose an additional energetic burden to the gestating or lactating mother. Finally, some of the problems with ascribing functional significance to brain size sexual dimorphism are discussed, and the energetic implications for brain size sexual dimorphism in infancy are assessed.     

Changes in the structure of the baboon hybrid zone in the Awash National Park,Ethiopia

The zone of hybridization between anubis and hamadryas baboons in the Awash National Park, Ethiopia, evidently changed significantly in the early 1970s. When first studied, by Nagel in 1968, the distribution of adult male phenotypes suggested that the hybrid zone was a narrow, static, genetic sink. In contrast, studies during the 1970s found evidence for a broad hybrid zone intergrading with parental populations and probably expanding. A more detailed investigation of data collected in 1973, presented here, shows that between 1968 and 1973 the distribution of phenotypes changed to a more clinal pattern because of the maturation and successful integration of an extensive cohort of hybrid males. We present a provisional model that accounts for the changes in terms of climatic oscillations, with the hybrid zone tending to expand during periods of climatic instability.     

Ethological study of the social behavior of hybrid baboons betweenPapio anubis andP. hamadryas in free-ranging groups

The male-female and male-male interactions of hybrid baboons betweenPapio anubis andP. hamadryas were analyzed quantitatively in two free-ranging groups in the Awash Valley, Ethiopia. Morphologically, one group (the Kerrayu group) was closer to hamadryas, while the other (the Gorge group) was closer to anubis. The adult males were classified into four categories; i.e., leaders of one-male units, males in pair units (both in the Kerrayu group), males with a closer appearance to hamadryas, and males with an anubis-like appearance (both in the Gorge group). These categories coincided with the morphological gradient from hamadryas to anubis. Social behavior was described in terms of 53 kinds of behavioral elements identified. In both groups, the closer a male was to being hamadryas in morphology, the more frequent was the aggressive behavior it tended to address toward partner females. The patterns of male-female interactions were compared between one-male units and pair units in the Kerrayu group. In one-male units, females responded immediately with obedient behavior to the males' behavior of directing visual attention toward them. In pair units, females responded with affinitive or avoiding behavior to physical contact by males. The flow of male social behavior was resolved into transitions from one behavioral element to another, and the succession rates for all the observed transitions were independently computed within each male category. By comparing the principal pathways of the behavioral sequences between male categories, 31 “basic pathways” were identified, which were presumed to be common to bothP. anubis andP. hamadryas. The basic pathways were composed of four main behavioral categories, i.e.,Contact, Visual-spacing, Aggressive behavior toward females, andAppeasing behavior toward males. In one-male units, transitions fromVisual-spacing toAggressive behavior were clearly recognized, whereas the principal pathways in pair units lacked such transitions. The frequency distributions of behavioral elements among the above behavioral categories were compared among the four male categories. The more prevalent anubis characters were in morphology, the larger was the proportion thatContact behavior occupied in the total amount of social behavior. The more prevalent hamadryas characters were, the larger was the proportion thatVisual-spacing behavior occupied. The leaders of one-male units showed a higher percentage ofAggressive behavior than did males of the other three categories. A conspicuous difference was observed in the percentage ofAppeasing behavior between the Gorge and Kerrayu groups. It appeared that the high frequency ofAppeasing behavior among males in the Kerrayu group contributed to the spatial cohesion of this group. The similarities of morphology, principal pathways, and frequency distribution of behavior were calculated for every combination of the four male categories. The morphological similarities correlated best with the similarities of principal pathways. It is concluded that the sequential structure of the male social behavior was strongly correlated with the genetic background, while the frequency of social behavior itself was subject to factors other than innate behavioral tendencies, such as the characteristics of the social environment and social context.     

Cerebrospinal Fluid Monoaminergic Metabolites in Wild Papio anubis and P. hamadryas are Concordant with Taxon-specific Behavioral Ontogeny

We used a cross-sectional sample to compare ontogenetic trajectories in the concentrations of monoamine neurotransmitter metabolites in cerebrospinal fluid of wild anubis (Papio anubis, n = 49) and hamadryas (P. hamadryas, n = 54) baboons to test the prediction that they would differ, especially in males, in association with their distinct behavioral ontogenies. Values of all 3 metabolites [3-methoxy-4-hydroxyphenylglycol (MHPG), the norepinephrine metabolite; 5-hydroxyindoleacetic acid (5-HIAA), the serotonin metabolite; and homovanillic acid (HVA), the dopamine metabolite] declined consistently with dentally-calibrated maturation, and few taxon-related differences were apparent among juveniles. Adult females were too few for adequate comparison, but a discriminant function suggested that they might differ by taxon. Adult males of the 2 species differed strikingly from juveniles and from each other. Contrary to our initial hypothesis, adult male anubis had significantly lower HVA and MHPG, and higher 5-HIAA levels, than predicted from the overall, age-related trend, and MHPG continued to decline with age among adults. As young adults, male hamadryas had low 5-HIAA and a high HVA/5-HIAA ratio, while older males [normatively one-male unit (OMU) leaders] showed a reversal in the trend, with 5-HIAA rising and the HVA/5-HIAA ratio tending to fall. We speculate that the results are related to the dispersing and philopatric ontogenies of anubis and hamadryas males, respectively. Adult male anubis, whose fitness depends on building social networks with nonkin, have high relative serotonin activity, commonly associated with greater social circumspection and skill. Young adult male hamadryas, living among agnatic kin and mating opportunistically, exhibit low 5-HIAA levels, generally associated with impulsivity and social irresponsibility. This reverses as a male approaches the age at which he is normatively the leader of a one-male unit (OMU), and his fitness depends on his maintaining stable relationships with other leaders and with females. An erratum to this article can be found at     

Distribution and Habitat Associations of Baboons (Papio hamadryas) in Central Eritrea

At least three diurnal primate taxa are still present in Eritrea, NE Africa: hamadryas baboons (Papio hamadryas hamadryas), olive baboons (Papio h. anubis) and grivet monkeys (Chlorocebus aethiops aethiops). However, information on status and distribution of primates and their habitats in Eritrea is outdated and incomplete. We conducted a primate survey, focussing on hamadryas baboons, to obtain data which will be integrated in a national wildlife management and conservation plan in Eritrea. We obtained information about the geographical distribution and abundance of baboons, their altitudinal range, habitat quality of their home-ranges, aggregation sizes at sleeping cliffs and predator presence. We described habitat quality via the Normalized Difference Vegetation Index (NDVI), a vegetation classification of Landsat MSS satellite data. Hamadryas and olive baboons are still present in Eritrea in ample numbers. Their geographical distributions in 1997 and 1998 did not deviate significantly from their historical distributions. An estimated 15,000 Papio hamadryas hamadryas lived in the 25,000-km² area of survey (0.58 baboons/km²). Population densities of hamadryas baboons in many parts of the survey area are higher than at Kummer's (1968) study site in Ethiopia. Hamadryas baboons live at all altitudes in four of five ecogeographical zones of Eritrea. Olive baboons replaced them in the western lowlands. Both baboon taxa tend to select better quality habitats, characterized by a higher normalized difference vegetation index (NDVI) than the average for the respective ecogeographical zones. Hamadryas baboons show a greater ecological plasticity than olive baboons, which are confined to riverbeds with extended gallery forest. By the end of 1999, a hybrid zone could not be confirmed.     

Female "dispersal" in hamadryas baboons: transfer among social units in a multilevel society

Unlike most cercopithecines, hamadryas baboons (Papio hamadryas hamadryas) are characterized by female-biased dispersal. To clarify this pattern within the context of their hierarchical social system (comprising one-male units, clans, bands, and troops), we report here 7 years of data on female transfers among social units in wild hamadryas baboons in Ethiopia. Female tenure in one-male units (OMUs) ranged from 1 to 2,556 days (N = 208) and survival analysis revealed a median tenure length of 1,217 days (40 months). Changes in OMU membership consisted almost exclusively of takeovers by males, not voluntary transfer. Of 130 takeovers, 67% occurred within the band and 33% across bands, and, of the 22 takeovers for which we have clan membership data, 77% occurred within, not between, clans. These results reinforce the notion that hamadryas female dispersal is not analogous to sex-biased dispersal in other taxa, because (1) at least in Ethiopian populations, females do not disperse voluntarily but are transferred, often forcibly, by males; (2) only dispersal between bands will promote gene flow, whereas females are most often rearranged within bands; (3) hamadryas females undergo social dispersal but not usually locational dispersal; and (4) while male hamadryas are far more philopatric than females, they have been observed to disperse. It thus appears that the ancestral baboon pattern of female philopatry and male dispersal has evolved into a system in which neither sex is motivated to disperse, but females are forcibly transferred by males, leading to female-mediated gene flow, and males more rarely disperse to find females.     

Mitochondrial evidence for the origin of hamadryas baboons

Baboons (Mammalia: Primates, Papio) are found primarily on the continent of Africa, but the range of hamadryas baboons (Papio hamadryas) extends to the Arabian Peninsula, and the origin of Arabian populations is unclear. To estimate the timing of the divergence between Arabian and African hamadryas populations we analyzed mitochondrial DNA (mtDNA) sequences from individuals of Arabian and African origin, and from representatives of the other major baboon taxa. The oldest hamadryas mitochondrial lineages in the Arabian Peninsula form an ancient trichotomy with the two major African lineages. This suggests that Arabia was colonized by hamadryas very soon after the appearance of the distinctive hamadryas phenotype, both events perhaps coinciding with a mid-Pleistocene stage of dry climate and low sea-level. The most closely related Arabian and African mtDNA haplotypes coalesce at approximately 35 ka, suggesting that no gene flow between African and Arabian baboons has occurred since the end of the last ice age, when a land bridge at the southern sill of the Red Sea was submerged. The mitochondrial paraphyly of Ethiopian hamadryas and anubis (P. anubis) baboons suggests an extensive and complex history of sex-specific introgression.     

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.     

Big-bodied males help us recognize that females have big pelves

Schultz ([1949] Am. J. Phys. Anthropol. 7:401-424) presented a conundrum: among primates, sexual dimorphism of the pelvis is a developmental adjunct to dimorphism in other aspects of the body, albeit in the converse direction. Among species in which males are larger than females in body size, females are larger than males in some pelvic dimensions; species with little sexual dimorphism in nonpelvic size show little pelvic dimorphism. Obstetrical difficulty does not explain this relationship. The present study addresses this issue, evaluating the relationship between pelvic and femoral sexual dimorphism in 12 anthropoid species. The hypothesis is that species in which males are significantly larger than females in femoral size will have a higher incidence, magnitude, and variability of pelvic sexual dimorphism, with females having relatively larger pelves than males, compared with species monomorphic in femoral size. The results are consistent with the hypothesis. The proposed explanation is that the default pelvic anatomy in adulthood is that of the female; testosterone redirects growth from the default type to that of the male by differentially enhancing and repressing growth among the pelvic dimensions. Testosterone also influences sexual dimorphism of the femur. The magnitude of the pelvic response to testosterone is greater in species that are sexually dimorphic in the femur than in those that are monomorphic.     

The relationship between sexual dimorphism in human faces and fluctuating asymmetry

Previous studies have found both support and lack of support for a positive relationship between masculinity and symmetry, two putative signs of mate quality, in male faces. We re-examined this relationship using an explicit measure of facial fluctuating asymmetry, as well as other measures of asymmetry, and measures of facial masculinity/femininity. We also used ratings of these traits for faces. Further, we examined the relationship between facial sexual dimorphism and body asymmetry. We found no significant correlations between facial masculinity and any of our measures of asymmetry or ratings of symmetry in males. Facial femininity was not consistently associated with facial symmetry in females, but was associated with body symmetry. Therefore, for females, but not males, facial femininity and body symmetry may reflect similar aspects of mate quality. We also examined the relationships between trait ratings and measurements. Our results provide validation of our ability to measure aspects of asymmetry that are perceived to be symmetrical, and aspects of sexual dimorphism that are perceived as feminine in females and masculine in males.