RESUMEN
Since Washburn's New Physical Anthropology, researchers have sought to understand the complexities of morphological evolution among anatomical regions in human and non-human primates. Researchers continue, however, to preferentially use comparative and functional approaches to examine complex traits, but these methods cannot address questions about evolutionary process and often conflate function with fitness. Moreover, researchers also tend to examine anatomical elements in isolation, which implicitly assumes independent evolution among different body regions. In this paper, we argue that questions asked in primate evolution are best examined using multiple anatomical regions subjected to model-bound methods built from an understanding of evolutionary quantitative genetics. A nascent but expanding number of studies over the last two decades use this approach, examining morphological integration, evolvability, and selection modeling. To help readers learn how to use these methods, we review fundamentals of evolutionary processes within a quantitative genetic framework, explore the importance of neutral evolutionary theory, and explain the basics of evolutionary quantitative genetics, namely the calculation of evolutionary potential for multiple traits in response to selection. Leveraging these methods, we demonstrate their use to understand non-independence in possible evolutionary responses across the limbs, limb girdles, and basicranium of humans. Our results show that model-bound quantitative genetic methods can reveal unexpected genetic covariances among traits that create a novel but measurable understanding of evolutionary complexity among multiple traits. We advocate for evolutionary quantitative genetic methods to be a standard whenever appropriate to keep studies of primate morphological evolution relevant for the next seventy years and beyond.
Asunto(s)
Evolución Biológica , Hominidae , Animales , Flujo Genético , Fenotipo , Primates/genéticaRESUMEN
OBJECTIVES: Though recent quantitative genetic analyses have indicated that directional selection appears to be acting on limb lengths and measures of body size in modern humans, these studies assume equal evolvability across modern human groups. However, differences in trait covariance structure due to ancient migration patterns and/or selection may limit the evolvability of populations further from Africa. This study therefore explores patterns of human evolvability across ecogeographic regions. MATERIALS AND METHODS: Mean evolvability, respondability, conditional evolvability, and autonomy were calculated from variance-covariance matrices of limb length and body size measures representing 14 human groups spanning four ecogeographic regions. Measures of evolvability were compared across groups and regions, and the minimum sample size, inaccuracy, and bias were calculated for each. RESULTS: When compared between regions, humans demonstrate significant differences between indices of evolvability across regions. Despite the relatively recent evolution of modern humans, several measures of evolvability show a strong negative correlation with latitude across regions, demonstrating a reduction in genetic variance that is potentially reflective of human migration and/or response to selection. CONCLUSIONS: These results demonstrate the importance of establishing patterns of evolvability prior to additional quantitative genetic analyses, and emphasize the influence of sample size on the accuracy of estimated evolvability measures. These findings also suggest that while modern human groups share similar covariance structures, there is evidence for emergent differentiation in evolvability and respondability between human groups across ecogeographic regions, further complicating our ability to apply results derived from modern human groups to ancient hominin lineages.
Asunto(s)
Evolución Biológica , Huesos/anatomía & histología , África del Sur del Sahara , África del Norte , Antropología Física , Regiones Árticas , Tamaño Corporal , Europa (Continente) , HumanosRESUMEN
Considerable research has shown that modern human pelvic dimensions, especially of the birth canal, are sexually dimorphic. Studies also suggest that females with younger ages-at-death have narrower canal dimensions than those who die at older ages, possibly due to continued independent growth of the pubis. A recent examination of this pattern argued that it is unlikely that these differences relate to mortality, but the source of the difference in pelvic dimensions with age remains unresolved. We use pelvic dimensions to assess differences in magnitudes of morphological integration between adult females and males across ages-at-death. We first ascertain whether the sexes have different strengths of integration, and then assess if differences in magnitudes of integration are associated with age-at-death. Pelvic dimensions of all groups were moderately integrated. Females and males have similar magnitudes of integration, and there is no change in the strength of integration with age. Examining individual regions of the pelvis indicates that the ilium, pubis, and pelvic inlet and outlet have stronger integration than the overall pelvis. This was particularly true of the pelvic outlet, which demonstrated the strongest integration. Our findings suggest that regions of the pelvis are more strongly integrated internally, and less integrated with each other, which would allow for proportional growth among regions of the pelvis with age that do not affect its overall integration. No single region of the pelvis appears to be motivating the difference in pelvic dimensions between age groups. We further consider the implications of these findings on evolutionary constraints. Anat Rec, 300:666-674, 2017. © 2017 Wiley Periodicals, Inc.
Asunto(s)
Huesos Pélvicos/anatomía & histología , Pelvis/anatomía & histología , Adulto , Factores de Edad , Femenino , Humanos , Masculino , Factores Sexuales , Adulto JovenRESUMEN
Variation in body form among human groups is structured by a blend of natural selection driven by local climatic conditions and random genetic drift. However, attempts to test ecogeographic hypotheses have not distinguished between adaptive traits (i.e., those that evolved as a result of selection) and those that evolved as a correlated response to selection on other traits (i.e., nonadaptive traits), complicating our understanding of the relationship between climate and morphological distinctions among populations. Here, we use evolutionary quantitative methods to test if traits previously identified as supporting ecogeographic hypotheses were actually adaptive by estimating the force of selection on individual traits needed to drive among-group differentiation. Our results show that not all associations between trait means and latitude were caused by selection acting directly on each individual trait. Although radial and tibial length and biiliac and femoral head breadth show signs of responses to directional selection matching ecogeographic hypotheses, the femur was subject to little or no directional selection despite having shorter values by latitude. Additionally, in contradiction to ecogeographic hypotheses, the humerus was under directional selection for longer values by latitude. Responses to directional selection in the tibia and radius induced a nonadaptive correlated response in the humerus that overwhelmed its own trait-specific response to selection. This result emphasizes that mean differences between groups are not good indicators of which traits are adaptations in the absence of information about covariation among characteristics.