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Morphological Integration and the Interpretation of Fossil Hominin Diversity

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Abstract

The fossil record of primate and human evolution cannot provide accurate estimates of within species variation and integration. This means that we cannot directly observe how patterns of integration have evolved over time in this lineage. And yet, our interpretations of fossil diversity are awash with assumptions about variation patterning in precisely these fossil taxa. Most commonly, researchers rely on extant models of variation for interpreting past diversity, by assuming equality of variation (and occasionally covariation) among extant and fossil populations. Yet one of the things we know from studies of integration in primates is that patterns of morphological covariation can differ among even closely related taxa, indicating that they have diverged over evolutionary time, either in response to selection or as the result of neutral evolution. At the same time, overall patterns of integration remain remarkably similar, meaning that in many respects they are highly conserved evolutionarily. Taken together, these seemingly contradictory observations offer an important conceptual framework for interpreting patterns that we observe in the fossil past. This framework dictates that while we can use patterns of covariation in extant taxa as proxies for extinct diversity, and indeed their conserved nature makes them superior to approaches that rely on variation alone, we also need to account for the fact that such patterns change over time, and incorporate that into our models. Here I provide examples using covariation patterns estimated from modern humans and African great apes to demonstrate the extent to which divergence in covariance structure might affect our interpretations of hominin diversity.

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Notes

  1. In its simplest definition, morphological integration is the connectivity or relationship among parts, and can be measured as trait covariation.

  2. As opposed to other measures of fossil sample size, which include total fossils and estimates of minimum numbers of individuals, effective sample size is an expression of known trait variability in the sample. Therefore, a sample with 10 fossils, representing five individuals, can still have an effective sample size of one if no skeletal element is shared among individuals.

  3. These data consist of 10 linear distances taken on the face, drawn from a world-wide human sample, as described in “Materials and Methods”; matrix correlations were measured via Pearson’s cross product.

  4. Such an approach would be specific to change seen in the fossil record. For example, a researcher might examine whether strains of mice with very different brain sizes have different patterns of cranial integration, in order to understand how evolutionary changes in patterns of integration—and presumably changes in underlying genetic/developmental architecture—might have corresponded with the emergence of a larger brain in the genus Homo.

  5. And indeed, visual inspection of the covariance matrices confirms that matrices accounting for sexual dimorphism plus subspecific structure and those accounting for subspecific structure alone differ very little.

  6. A case could be made for either. We might expect the larger Neanderthals as a taxon to be more variable than modern humans, who have gone through an evolutionary bottleneck; this would act to increase the pairwise Neanderthal distances. Conversely, since human morphology reflects world-wide diversity and the Neanderthals are a geographically restricted group, the opposite might be true.

  7. Indeed, Pearson correlations between the pooled-sex human, chimpanzee, and gorilla covariance matrices used here also support this notion (human–chimpanzee correlation = 0.945; human–gorilla = 0.932; chimpanzee–gorilla = 0.956), and moreover suggest that humans differ primarily in pattern, as size differences would be reflected in a closer link between humans and chimpanzees to the exclusion of gorillas.

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Acknowledgements

I wish to thank Katherine Willmore and Campbell Rolian for organizing the AAPA symposium. Thanks as well to all of the curators who have allowed access to specimens in their care over the years. Gail Krovitz was particularly generous in supplying her Neanderthal data, and I thank her for that. Finally, special thanks to Jim Cheverud for being a wonderful mentor, colleague, and friend.

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Correspondence to Rebecca Rogers Ackermann.

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Ackermann, R.R. Morphological Integration and the Interpretation of Fossil Hominin Diversity. Evol Biol 36, 149–156 (2009). https://doi.org/10.1007/s11692-009-9050-2

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