Heather F. Smith

Which cranial regions reflect molecular distances reliably in humans? Evidence from three-dimensional morphology.

Knowledge of the degree to which various subsets of morphological data reflect molecular relationships is crucial for studies attempting to estimate genetic relationships from patterns of morphological variation. This study assessed the phylogenetic utility of six different human cranial regions, plus the entire cranium. Three‐dimensional landmark data were collected for 83 landmarks from samples of skulls from 14 modern human populations. The data were subsequently divided into anatomical regions: basicranium, upper face, mandible, temporal bone, upper jaw, cranial vault, and a subset of points from around the entire cranium. Depictions of population molecular distances were calculated using published data on microsatellites for the same or closely related populations. Distances based on morphological variation of each of the anatomical regions were compared with molecular distances, and the correlations assessed. The morphology of the basicranium, temporal bone, upper face, and entire cranium demonstrated the highest correlations with molecular distances. The morphology of the mandible, upper jaw, and cranial vault, as measured here, were not significantly correlated with molecular distances. As the three‐dimensional morphology of the temporal bone, upper face, basicranium, and entire cranium appear to consistently reflect genetic relationships in humans, especially with more reliability than the cranial vault, it would be preferable to focus on these regions when attempting to determine the genetic relationships of human specimens with no molecular data. Am. J.Hum. Biol., 2009.

Comparative anatomy and phylogenetic distribution of the mammalian cecal appendix

A recently improved understanding of gut immunity has merged with current thinking in biological and medical science, pointing to an apparent function of the mammalian cecal appendix as a safe‐house for symbiotic gut microbes, preserving the flora during times of gastrointestinal infection in societies without modern medicine. This function is potentially a selective force for the evolution and maintenance of the appendix, and provides an impetus for reassessment of the evolution of the appendix. A comparative anatomical approach reveals three apparent morphotypes of the cecal appendix, as well as appendix‐like structures in some species that lack a true cecal appendix. Cladistic analyses indicate that the appendix has evolved independently at least twice (at least once in diprotodont marsupials and at least once in Euarchontoglires), shows a highly significant (P < 0.0001) phylogenetic signal in its distribution, and has been maintained in mammalian evolution for 80 million years or longer.

Cladistic analysis of early Homo crania from Swartkrans and Sterkfontein, South Africa

The phylogenetic relationships of early Pleistocene Homo crania from the South African sites of Swartkrans and Sterkfontein were investigated through cladistic analyses of 99 morphological characters. The Swartkrans Member 1 specimen SK 847 and the Stw 53 cranium from Sterkfontein Member 5A were treated as separate operational taxonomic units (OTUs), distinct from the three species of early Homo-H. erectus, H. habilis, and H. rudolfensis-that are recognized from the Plio-Pleistocene deposits of East Africa. The cladistic analyses differed in the treatment of the South African OTUs (separate Swartkrans and Sterkfontein OTUs vs. a single Swartkrans+Sterkfontein OTU). PAUP 4.0 was used to construct cladograms and address hypotheses about relationships. In the analysis that treated the South African specimens as a single OTU, the position of that OTU was stable as a separate branch on the Homo clade between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)]. When SK 847 and Stw 53 were treated as separate OTUs, the majority of most parsimonious trees indicated that they were positioned in similar positions as the combined South African Homo OTU; that is, as separate branches between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)], with the Swartkrans OTU generally occupying a more derived position. The position of the Sterkfontein OTU was more stable than that of the Swartkrans OTU, which was found in several other positions among the minimum length trees. Running the analyses with only those characters preserved by SK 847 and Stw 53 resulted in similar topologies for minimum length trees, although the positions of Stw 53, SK 847, and H. habilis exchanged places in some trees. In no case was an exclusive sister relationship between either South African OTU and a particular species of Homo supported statistically. Both South African OTUs differ from H. habilis in the fewest number of cladistic characters.

The Role of Genetic Drift in Shaping Modern Human Cranial Evolution: A Test Using Microevolutionary Modeling

The means by which various microevolutionary processes have acted in the past to produce patterns of cranial variation that characterize modern humans is not thoroughly understood. Applying a microevolutionary framework, within- and among-population variance/covariance (V/CV) structure was compared for several functional and developmental modules of the skull across a worldwide sample of modern humans. V/CV patterns in the basicranium, temporal bone, and face are proportional within and among groups, which is consistent with a hypothesis of neutral evolution; however, mandibular morphology deviated from this pattern. Degree of intergroup similarity in facial, temporal bone, and mandibular morphology is significantly correlated with geographic distance; however, much of the variance remains unexplained. These findings provide insight into the evolutionary history of modern human cranial variation by identifying signatures of genetic drift, gene flow, and migration and set the stage for inferences regarding selective pressures that early humans encountered since their initial migrations around the world.

The relative congruence of cranial and genetic estimates of hominoid taxon relationships: Implications for the reconstruction of hominin phylogeny

Previous analyses of extant catarrhine craniodental morphology have often failed to recover their molecular relationships, casting doubt on the accuracy of hominin phylogenies based on anatomical data. However, on the basis of genetic, morphometric and environmental affinity patterns, a growing body of literature has demonstrated that particular aspects of cranial morphology are remarkably reliable proxies for neutral modern human population history. Hence, it is important to test whether these intra-specific patterns can be extrapolated to a broader primate taxon level such that inference rules for understanding the morphological evolution of the extinct hominins may be devised. Here, we use a matrix of molecular distances between 15 hominoid taxa to test the genetic congruence of 14 craniomandibular regions, defined and morphometrically delineated on the basis of previous modern human analyses. This methodology allowed us to test directly whether the cranial regions found to be reliable indicators of population history were also more reliable proxies for hominoid genetic relationships. Cranial regions were defined on the basis of three criteria: developmental-functional units, individual bones, and regions differentially affected by masticatory stress. The results found that all regions tested were significantly and strongly correlated with the molecular matrix. However, the modern human predictions regarding the relative congruence of particular regions did not hold true, as the face was statistically the most reliable indicator of hominoid genetic distances, as opposed to the vault or basicranium. Moreover, when modern humans were removed from the analysis, all cranial regions improved in their genetic congruence, suggesting that it is the inclusion of morphologically-derived humans that has the largest effect on incongruence between morphological and molecular estimates of hominoid relationships. Therefore, it may be necessary to focus on smaller intra-generic taxonomic levels to more fully understand the effects of neutral and selective evolutionary processes in generating morphological diversity patterns.

Controllability in Environmental Enrichment for Captive Chimpanzees (Pan troglodytes)

This study considers the use of nonsocial environmental enrichment by captive chimpanzees at the Primate Foundation of Arizona. The goal was to determine whether a relationship existed between controllability of enrichment items by captive chimpanzees and frequency of use. The study measured controllability, the ability of nonhuman animals to alter aspects of their environment by the potential destructibility of the enrichment item. This study examined additional factors that may affect enrichment use: individual age, sex, rearing history, social group composition, and availability of outdoor access. The chimpanzees in the study used destructible items—the enrichment category with the highest level of controllability—more than indestructible items across all age, sex, and rearing classes. Thus, controllability seems to be an important factor in chimpanzee enrichment. Younger individuals and groups with outdoor access used enrichment more than did older individuals and groups with indoor-only access. Individual sex, rearing history, and social group composition had minimal effects on enrichment use. These results support the importance of control to captive chimpanzees and further enable captive management to customize enrichment programs to the needs of particular animals.

A 3-D geometric morphometric study of intraspecific variation in the ontogeny of the temporal bone in modern Homo sapiens

This study addresses how the human temporal bone develops the population-specific pattern of morphology observed among adults and at what point in ontogeny those patterns arise. Three-dimensional temporal bone shape was captured using 15 landmarks on ontogenetic series of specimens from seven modern human populations. Discriminant function analysis revealed that population-specific temporal bone morphology is evident early in ontogeny, with significant shape differences among many human populations apparent prior to the eruption of the first molar. As early as five years of age, temporal bone shape reflects population history and can be used to reliably sort populations, although those in closer geographic proximity and molecular affinity are more likely to be misclassified. The deviation of cold-adapted populations from this general pattern of congruence between temporal bone morphology and genetic distances, identified in previous work, was confirmed here in adult and subadult specimens, and was revealed to occur earlier in ontogeny than previously recognized. Significant differences exist between the ontogenetic trajectories of some pairs of populations, but not among others, and the angles of these trajectories do not reflect genetic relationships or final adult temporal bone size. Significant intrapopulation differences are evident early in ontogeny, with differences becoming amplified by divergent trajectories in some groups. These findings elucidate how the congruence between adult human temporal bone morphology and population history develops, and reveal that this pattern corresponds closely to that described previously for facial ontogeny.

Population genetic structure of traditional populations in the Peruvian Central Andes and implications for South American population history.

ABSTRACT Molecular-based characterizations of Andean peoples are traditionally conducted in the service of elucidating continent-level evolutionary processes in South America. Consequently, genetic variation among “western” Andean populations is often represented in relation to variation among “eastern” Amazon and Orinoco River Basin populations. This west-east contrast in patterns of population genetic variation is typically attributed to large-scale phenomena, such as dual founder colonization events or differing long-term microevolutionary histories. However, alternative explanations that consider the nature and causes of population genetic diversity within the Andean region remain underexplored. Here we examine population genetic diversity in the Peruvian Central Andes using data from the mtDNA first hypervariable region and Y-chromosome short tandem repeats among 17 newly sampled populations and 15 published samples. Using this geographically comprehensive data set, we first reassessed the currently accepted pattern of western versus eastern population genetic structure, which our results ultimately reject: mtDNA population diversities were lower, rather than higher, within Andean versus eastern populations, and only highland Y-chromosomes exhibited significantly higher within-population diversities compared with eastern groups. Multiple populations, including several highland samples, exhibited low genetic diversities for both genetic systems. Second, we explored whether the implementation of Inca state and Spanish colonial policies starting at about ad 1400 could have substantially restructured population genetic variation and consequently constitute a primary explanation for the extant pattern of population diversity in the Peruvian Central Andes. Our results suggest that Peruvian Central Andean population structure cannot be parsimoniously explained as the sole outcome of combined Inca and Spanish policies on the regions population demography: highland populations differed from coastal and lowland populations in mtDNA genetic structure only; highland groups also showed strong evidence of female-biased gene flow and/or effective sizes relative to other Peruvian ecozones. Taken together, these findings indicate that population genetic structure in the Peruvian Central Andes is considerably more complex than previously reported and that characterizations of and explanations for genetic variation may be best pursued within more localized regions and defined time periods.

Anatomical variation in the anterolateral ligament of the knee and a new dissection technique for embalmed cadaveric specimens

Claes et al. recently documented and described the anterolateral ligament (ALL) of the knee, demonstrating its existence in 97% of their samples. Here, we further examined the anatomy of this ligament, documented its morphological variation, and assessed the feasibility of its dissection in preserved cadaveric specimens. To achieve this, we dissected 53 preserved cadaveric knees and documented their morphological variation in the anterolateral ligament. The originally described dissection technique for identifying and following the ALL requires flexion of the knee, a state which is often not possible in stiff, preserved cadavers. Here, we describe and confirm the feasibility of an alternate dissection technique in which the quadriceps femoris tendon is incised, for use on specimens in which flexion of the undissected knee is not possible. We also identify a novel technique for assessing whether the anterolateral ligament is absent from a specimen or has simply been obliterated or overlooked, using the lateral inferior genicular vasculature. These dissection techniques have great potential for the dissection of preserved cadavers used in gross anatomy laboratories, and we discuss the applications of such an approach in student-led dissections. Our dissections also uncovered noticeable variation in the anterolateral ligament course and position. Most notably, it often inserts significantly more laterally than the classical presentation (30.2%), or originates more proximally with superficial fibers extending superiorly and laterally over the distal femur (7.5%).

Myology of the Head, Neck, and Thoracic Region of the Lesser Grison ( Galictis cuja ) in Comparison with the Red Panda ( Ailurus fulgens ) and Other Carnivorans: Phylogenetic and Functional Implications

The lesser grison (Galictis cuja) and the red panda (Ailurus fulgens) represent two opposed morpho-functional musteloid extremes. The mid-sized lesser grison is primarily terrestrial, a frequent burrow-dweller, and carnivorous, while the larger, scansorial red panda eats bamboo. This study documents the axial myology of these species, including muscle descriptions, weights, and optimizations. Muscle maps are also provided for the lesser grison, representing the first axial maps for a wild-caught carnivoran. The functional analyses revealed that G. cuja, contrary to A. fulgens, possesses longer, stronger, and subdivided neck muscles. It also possesses a thoraco-lumbar iliocostalis system that is more developed than the longissimus complex, and numerous, robust, and laterally inserted deep bellies of the cervical and thoracic transversospinalis systems. These specializations allow powerful neck movement during hunting and transport of heavy prey as well as axial flexibility, facilitating bounding gaits and lateral movements while navigating subterranean galleries. Some myological traits of the red panda differ from those expected in a highly herbivorous taxon (e.g., m. sternocephalicus, m. masseter), and may reflect its depredatory ancestry. The optimization analysis revealed phylogenetically informative traits across Carnivora, including the absence of m. longissimus capitis in Mephitidae, the absence of spinous thoracic origins for m. biventer cervicis in Musteloidea, and the presence of a relatively lateral insertion of m. rectus dorsalis capitis intermedius in the clade Ictonychinae+Lutrinae+Mustelinae. This study reveals key associations between axial myological and osteological features that will prove useful for future studies of carnivorans.