Using geometric morphometrics to determine the "fittest" floral shape: A case study in large-flowered, buzz-pollinated Meriania hernandoi.

PREMISE: Floral shape (relative arrangement and position of floral organs) is critical in mediating fit with pollinators and maximizing conspecific pollen transfer particularly in functionally specialized systems. To date, however, few studies have attempted to quantify flowers as the inherently three-dimensional (3D) structures they are and determine the effect of intraspecific shape variation on pollen transfer. We here addressed this research gap using a functionally specialized system, buzz pollination, in which bees extract pollen through vibrations, as a model. Our study species, Meriania hernandoi (Melastomataceae), undergoes a floral shape change from pseudocampanulate corollas with more actinomorphically arranged stamens (first day) to open corollas with a more zygomorphic androecium (second day) over anthesis, providing a natural experiment to test how variation in floral shape affects pollination performance. METHODS: In one population of M. hernandoi, we bagged 51 pre-anthetic flowers and exposed half of them to bee pollinators when they were in either stage of their shape transition. We then collected flowers, obtained 3D flower models through x-ray computed tomography for 3D geometric morphometric analyses, and counted the pollen grains remaining per stamen (male pollination performance) and stigmatic pollen loads (female pollination performance). RESULTS: Male pollination performance was significantly higher in open flowers with zygomorphic androecia than in pseudo-campanulate flowers. Female pollination performance did not differ among floral shapes. CONCLUSIONS: These results suggest that there is an "optimal" shape for male pollination performance, while the movement of bees around the flower when buzzing the spread-out stamens results in sufficient pollen deposition regardless of floral shape.

Transgenerational effects of parental crude oil exposure on the morphology of adult Fundulus grandis.

The current study involved exposing adult F0 Gulf killifish (Fundulus grandis) to Macondo-252 oil for 36 to 44 days and assessing the effects of this oiling on the swimming performance and morphology in two generations of progeny reared in clean water. Following exposure to oil, the F0 fish were used as broodstock to generate four lineages of F1 fish using a full-matrix mating design derived from the gametes of clean and oil-exposed parents. Later, the four lineages of F1 fish were used as broodstock to create an F2 generation of the same four lineages. We found few differences in embryonic outcome (% dead,% hatched, and% unhatched) in any of the four lineages of F1 and F2 fish. However, as adults, F1 and F2 fish derived from oil-exposed males from the F0 generation had significantly lower critical swimming speeds (Ucrit) than both the control and maternally oil-exposed lineages. Additionally, progeny of oil-exposed fish had altered body shape based on the statistical analysis of two-dimensional landmark-based geometric morphometrics. Fish from oil-exposed lineages showed increased body depth, altered spinal curvature, and changes in the upward angle of projection of the head. Both generations had a significant main effect of maternal and paternal oil exposure on shape; however, F0 paternal oil exposure explained more of the variance in shape across both generations relative to F0 maternal exposure. Our findings demonstrate that parental exposure to oil can impact the shape and aerobic swimming capacity of offspring for at least two generations after the original paternal oiling.

Vertebral pneumaticity is correlated with serial variation in vertebral shape in storks.

Birds and their ornithodiran ancestors are unique among vertebrates in exhibiting air-filled sinuses in their postcranial bones, a phenomenon called postcranial skeletal pneumaticity. The factors that account for serial and interspecific variation in postcranial skeletal pneumaticity are poorly understood, although body size, ecology, and bone biomechanics have all been implicated as influencing the extent to which pneumatizing epithelia invade the skeleton and induce bone resorption. Here, I use high-resolution computed-tomography to holistically quantify vertebral pneumaticity in members of the neognath family Ciconiidae (storks), with pneumaticity measured as the relative volume of internal air space. These data are used to describe serial variation in extent of pneumaticity and to assess whether and how pneumaticity varies with the size and shape of a vertebra. Pneumaticity increases dramatically from the middle of the neck onwards, contrary to previous predictions that cervical pneumaticity should decrease toward the thorax to maintain structural integrity as the mass and bending moments of the neck increase. Although the largest vertebrae sampled are also the most pneumatic, vertebral size cannot on its own account for serial or interspecific variation in extent of pneumaticity. Vertebral shape, as quantified by three-dimensional geometric morphometrics, is found to be significantly correlated with extent of pneumaticity, with elongate vertebrae being less pneumatic than craniocaudally short and dorsoventrally tall vertebrae. Considered together, the results of this study are consistent with the hypothesis that shape- and position-specific biomechanics influence the amount of bone loss that can be safely tolerated. These results have potentially important implications for the evolution of vertebral morphology in birds and their extinct relatives.

The human remains from Axlor (Dima, Biscay, northern Iberian Peninsula).

OBJECTIVES: We provide the description and comparative analysis of all the human fossil remains found at Axlor during the excavations carried out by J. M. de Barandiarán from 1967 to 1974: a cranial vault fragment and seven teeth, five of which likely belonged to the same individual, although two are currently lost. Our goal is to describe in detail all these human remains and discuss both their taxonomic attribution and their stratigraphic context. MATERIALS AND METHODS: We describe external and internal anatomy, and use classic and geometric morphometrics. The teeth from Axlor are compared to Neandertals, Upper Paleolithic, and recent modern humans. RESULTS: Two teeth (a left dm2 , a left di1 ) and the parietal fragment show morphological features consistent with a Neandertal classification, and were found in an undisturbed Mousterian context. The remaining three teeth (plus the two lost ones), initially classified as Neandertals, show morphological features and a general size that are more compatible with their classification as modern humans. DISCUSSION: A left parietal fragment (Level VIII) from a single probably adult Neandertal individual was recovered during the old excavations performed by Barandiarán. Additionally, two different Neandertal children lost deciduous teeth during the formations of levels V (left di1 ) and IV (right dm2 ). In addition, a modern human individual is represented by five remains (two currently lost) from a complex stratigraphic setting. Some of the morphological features of these remains suggest that they may represent one of the scarce examples of Upper Paleolithic modern human remains in the northern Iberian Peninsula, which should be confirmed by direct dating.

Short communication: Selection for persistency by changing the shape of lactation curves with geometric morphometrics.

The objective was to simulate the response to selection by modifying the shape of the lactation curve to improve persistency in Israeli first-parity Holstein cattle using geometric morphometrics and multivariate quantitative genetics. The 2-dimensional Cartesian coordinates representing the shape of the lactation curve were characterized using geometric morphometrics. Genetic parameters of daily milk yield were estimated for the individual animal model using REML. Results of selection were simulated using the multivariate breeders' equation. We defined 6 selection regimens: (1) decrease in the third month in milk (MIM), (2) increase in the ninth MIM, (3) decrease in the third MIM and increase in the ninth MIM, (4) decrease in the first MIM, (5) decrease in the second MIM, and (6) decrease in the fourth MIM. The first 3 selection criteria targeted changes in the MIM of previous measurements of persistency in the Israeli Holstein population, and the last 3 targeted the months with the highest variability. All selection regimens attempted to modify the shape of the lactation curve while keeping total milk yield constant. The first 3 selection regimens showed greater total responses, with the highest response for decrease in the third MIM and increase in the ninth MIM. Magnitudes of correlated responses consistently surpassed direct responses in all selection criteria, suggesting that selection on specific parts of the lactation curve would produce correlated responses in other parts. Predicted responses to selections were depicted as shape changes from the mean shape in the data set. Most predicted responses demonstrated a fairly similar shape compared with the lactation curve. The predicted responses of lactation shapes of the last 3 selection regimens were overall lower in the first 3 MIM and decreased less in the last MIM, demonstrating a much flatter shape and thus improved persistency.

Diachronic changes in craniofacial morphology among the middle-late Holocene populations from Hehuang region, Northwest China.

OBJECTIVES: This study analyzes craniofacial shape variation in the Hehuang region of Northwest China within a population genetic framework, and takes a diachronic approach to explore the relationship betwee cultural discontinuity and biological continuity/discontinuity in the Hehuang region during the middle to late Holocene. MATERIALS AND METHODS: The sample comprises 76 adult skulls from five archaeological sites, ranging from 4,500 to 1,530 BP. 3D geometric morphometrics, multivariate statistics, quantitative evolutionary genetic and biodistance analyses were performed to study the diachronic variation in craniofacial morphology. Analyses were performed on two cranial modules: the face and the cranial vault, across three major diachronic groups representing the late Neolithic (LNA), the Bronze Age (BA), and the Han-Jin dynasty (HD). RESULTS: Average regional FST for both cranial modules was low, indicating relatively greater variation within diachronic groups than among them. While the LNA and BA groups did not show any significant differences in facial and vault shape, significant craniofacial shape changes were detected between the BA and HD groups. DISCUSSION: The consistent craniofacial morphology during the LNA and the BA, and the shift in morphology between the BA and the HD indicates that cultural discontinuity does not always coincide with biological discontinuity. The Hehuang population evolved in situ with few changes, despite cultural and dietary changes, until the HD when migrations from the Central Plains are associated with extra-local gene flow to the area.

Describing the mandible in patients with craniofacial microsomia based on principal component analysis and thin plate spline video analysis.

Craniofacial microsomia (CFM) is most often described as a unilateral malformation of derivatives of the first and second branchial arches. The mandible has been classified using several classification systems. However, all are based on two-dimensional imaging. The aim of this study was to mathematically describe the deformed mandible based on principal component analysis (PCA) in a three-dimensional way. This may aid in defining the flaws in existing surgical corrections of the mandible through the identification of the differences in shape compared with a normal mandible in a holistic view with the help of videos. Forty-three homologous landmarks were defined to describe a mandible with CFM. Computed tomography scans of 22 patients and 30 controls were marked manually. The changes in shape between the mandibles were visualized using videos. A lateral rotation with increase in posterior rotation of the condyle due to shortening of the condyle-gonial height and a longitudinal rotation with outward bending of the mandibular angle were noted on the affected side, as well as an inward bending of the angle on the unaffected side. Due to the compensatory remodelling of the mandible on the unaffected side, one could suggest that CFM is never truly unilateral.

Bilateral asymmetry of the humerus in Neandertals, Australian aborigines and medieval humans.

OBJECTIVES: Bilateral asymmetry of diaphyseal shape and size may be a reflection of relative activity levels and patterns of habitual biomechanical stress in the upper arms of Neandertals and Homo sapiens. The main purpose of our study was to assess the level of directional asymmetry of humeral cross sections in Neandertals, recent Australian aborigines, and medieval farmers. MATERIALS AND METHODS: Indices of directional and absolute asymmetry (%DA and %AA) of humeral cross-sectional properties in Neandertals and recent Homo sapiens were calculated. Evenly distributed semilandmarks around the external and internal borders of cortical bone were digitized in the course of computed tomography for analysis of shape differences between sides of the body. RESULTS: The medieval farmers were characterized by significant %DA and %AA for polar second moment of area (J), ratio of maximum to minimum second moments of area, and ratio of antero-posterior to medio-lateral bending strength. In Australian aborigines, only J in males shows significant %DA and %AA, while Neandertals exhibit no significant asymmetry of any cross-sectional properties. Differences in cross-sectional shape between sides of the body were established in all three analyzed groups. DISCUSSION: High levels of directional asymmetry of cross-sectional shape and properties in medieval farmers may be caused by the performance of more physically demanding tasks using one side of the body from an early age in that population. Various patterns of asymmetry in Neandertals and modern humans may be caused by different habitual behaviors during growth, eco-geographic patterns in body proportions, genetic factors, and differences in ontogeny.

Evolutionary population history of early Paleoamerican cranial morphology.

The nature and timing of the peopling of the Americas is a subject of intense debate. In particular, it is unclear whether high levels of between-group craniometric diversity in South America result from multiple migrations or from local diversification processes. Previous attempts to explain this diversity have largely focused on testing alternative dispersal or gene flow models, reaching conflicting or inconclusive results. Here, a novel analytical framework is applied to three-dimensional geometric morphometric data to partition the effects of population divergence from geographically mediated gene flow to understand the ancestry of the early South Americans in the context of global human history. The results show that Paleoamericans share a last common ancestor with contemporary Native American groups outside, rather than inside, the Americas. Therefore, and in accordance with some recent genomic studies, craniometric data suggest that the New World was populated by multiple waves of dispersion from northeast Asia throughout the late Pleistocene and early Holocene.

Precuneus proportions and cortical folding: A morphometric evaluation on a racially diverse human sample.

Recent analyses have suggested that the size and proportions of the precuneus are remarkably variable among adult humans, representing a major source of geometrical difference in midsagittal brain morphology. The same area also represents the main midsagittal brain difference between humans and chimpanzees, being more expanded in our species. Enlargement of the upper parietal surface is a specific feature of Homo sapiens, when compared with other fossil hominids, suggesting the involvement of these cortical areas in recent modern human evolution. Here, we provide a survey on midsagittal brain morphology by investigating whether precuneus size represents the largest component of variance within a larger and racially diverse sample of 265 adult humans. Additionally, we investigate the relationship between precuneus shape variation and folding patterns. Precuneus proportions are confirmed to be a major source of human brain variation even when racial variability is considered. Larger precuneus size is associated with additional precuneal gyri, generally in its anterior district. Spatial variation is most pronounced in the dorsal areas, with no apparent differences between hemispheres, between sexes, or among different racial groups. These dorsal areas integrate somatic and visual information together with the lateral elements of the parietal cortex, representing a crucial node for self-centered mental imagery. The histological basis and functional significance of this intra-specific variation in the upper precuneus remains to be evaluated.

Holocene population history of the Sabana de Bogotá region, Northern South America: An assessment of the craniofacial shape variation.

OBJECTIVES: Several authors using multiple and independent lines of evidence investigating the biocultural continuity versus discontinuity in the Sabana de Bogotá region, in the eastern highlands of Colombia, have arrived at contradictory conclusions supporting either scenarios. This study analyzes the craniofacial size and shape variation of diachronic samples from the study region to test distinct population history scenarios that support continuity or, alternatively, divergence. MATERIALS AND METHODS: A total of 92 adult skulls belonging to five chronological groups, ranging from c. 10,100 to 350 14 C YBP, were analyzed through Procrustean geometric morphometric techniques. Matrix correlation analysis, multivariate exploratory (PCA, FDA), and evolutionary quantitative genetic methods (R-matrix analysis and ß-test) were used to study the diachronic craniofacial shape variation. RESULTS: A model that supports strong evolutionary diversification over the Holocene better explains the patterns of morphological variation observed. At least two periods of significant craniofacial size and shape change were detected: one during the middle to initial late Holocene transition (c. 4,000-3,200 14 C YBP) and other toward the final late Holocene (post-2,000 14 C YBP), which exhibit differences in the pattern and magnitude of cranial divergence. In addition, the differentiation viewed between early and mid-Holocene foragers could mark the initial entry of non-local populations into the region toward the beginnings of the middle Holocene. DISCUSSION: Distinct to previous investigations the present study supports a more complex regional population history where multiple population contractions/extinctions, dispersals and assimilations along with dietary adaptations took place during the last 10,000 years. These results are in agreement with the archaeological and paleoecological record which suggests marked periods of change rather than temporal stability.

Small reductions in corolla size and pollen: ovule ratio, but no changes in flower shape in selfing populations of the North American Arabidopsis lyrata.

The shift from outcrossing to selfing is often accompanied by striking changes in floral morphology towards a "selfing syndrome", which is characterized by flowers with reduction in size, pollen: ovule (P/O) ratio, and herkogamy. This study aims to test whether such changes have occurred in the North American Arabidopsis lyrata, which is of particular interest because of the relatively recent transitions to selfing in this system. Flower size, flower shape, herkogamy levels, P/O ratio, and floral integration of six self-incompatible (outcrossing) and six self-compatible (selfing) populations of A. lyrata were measured in a common environment using conventional and geometric morphometrics methods. Although selfers had on average 9.2% smaller corollas, 8.4% longer pistils, and 21.5% lower P/O ratios than outcrossers, there were no differences in shape, floral integration, and herkogamy between outcrossing and selfing populations. Moreover, most variation in floral traits was explained by population genetic background rather than by mating system. We conclude that selfing populations in A. lyrata have not evolved a selfing syndrome.

The Neanderthal in the karst: First dating, morphometric, and paleogenetic data on the fossil skeleton from Altamura (Italy).

In 1993, a fossil hominin skeleton was discovered in the karst caves of Lamalunga, near Altamura, in southern Italy. Despite the fact that this specimen represents one of the most extraordinary hominin specimens ever found in Europe, for the last two decades our knowledge of it has been based purely on the documented on-site observations. Recently, the retrieval from the cave of a fragment of bone (part of the right scapula) allowed the first dating of the individual, the quantitative analysis of a diagnostic morphological feature, and a preliminary paleogenetic characterization of this hominin skeleton from Altamura. Overall, the results concur in indicating that it belongs to the hypodigm of Homo neanderthalensis, with some phenetic peculiarities that appear consistent with a chronology ranging from 172 ± 15 ka to 130.1 ± 1.9 ka. Thus, the skeleton from Altamura represents the most ancient Neanderthal from which endogenous DNA has ever been extracted.

Perceived aggressiveness predicts fighting performance in mixed-martial-arts fighters.

Accurate assessment of competitive ability is a critical component of contest behavior in animals, and it could be just as important in human competition, particularly in human ancestral populations. Here, we tested the role that facial perception plays in this assessment by investigating the association between both perceived aggressiveness and perceived fighting ability in fighters' faces and their actual fighting success. Perceived aggressiveness was positively associated with the proportion of fights won, after we controlled for the effect of weight, which also independently predicted perceived aggression. In contrast, perception of fighting ability was confounded by weight, and an association between perceived fighting ability and actual fighting success was restricted to heavyweight fighters. Shape regressions revealed that aggressive-looking faces are generally wider and have a broader chin, more prominent eyebrows, and a larger nose than less aggressive-looking faces. Our results indicate that perception of aggressiveness and fighting ability might cue different aspects of success in male-male physical confrontation.