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Wing geometry as a tool for studying the Lutzomyia longipalpis (Diptera: Psychodidae) complex.

Toro Toro (T) and Yungas (Y) have been described as genetically well differentiated populations of the Lutzomyia longipalpis (Lutz & Neiva, 1912) complex in Bolivia. Here we use geometric morphometrics to compare samples from these populations and new populations (Bolivia and Nicaragua), representing distant geographical origins, qualitative morphological variation ("one-spot" or "two-spots" phenotypes), ecologically distinct traits (peridomestic and silvatic populations), and possibly different epidemiological roles (transmitting or nor transmitting Leishmania chagasi). The Nicaragua (N) (Somotillo) sample was "one-spot" phenotype and a possible peridomestic vector. The Bolivian sample of the Y was also "one-spot" phenotype and a demonstrated peridomestic vector of visceral leishmaniasis (VL). The three remaining samples were silvatic, "two-spots" phenotypes. Two of them (Uyuni and T) were collected in the highlands of Bolivian where VL never has been reported. The last one (Robore, R) came from the lowlands of Bolivia, where human cases of VL are sporadically reported. The decomposition of metric variation into size and shape by geometric morphometric techniques suggests the existence of two groups (N/Y/R, and U/T). Several arguments indicate that such subdivision of Lu. longipalpis could correspond to different evolutionary units.

Animals↗

Bilateral symmetry of the human metacarpal: implications for sample size calculations.

OBJECTIVE: The aim of this study was to assess the three-dimensional mechanical symmetry of the human second metacarpal and provide sample size estimates for future mechanical intervention studies of the metacarpal. DESIGN: Bone densitometry and digital image analysis were used to assess the morphometric, geometric and densitometric symmetry of the second human metacarpal. BACKGROUND: An assessment of the left-right mechanical symmetry of the human metacarpal is important in considering the suitability of using the contralateral metacarpal as a control and in providing sample size calculations for future studies involving a mechanical intervention to the metacarpal such as implantation of a metacarpophalangeal prosthesis. METHODS: Metaphyseal sectional areas, diaphyseal cortical sectional areas, second moments of area, average periosteal and medullary radii and bone densities were measured at nine transverse levels for each of seven pairs of index metacarpals using computed tomography and bone densitometry. Polar Fourier regression was used to assess the morphometry of sectional periosteal and endosteal boundaries. Differences between clinically important left-right parameters were assessed. RESULTS: Mean differences between clinically important left-right parameters were small (<3%) and similar to the degree of experimental precision. There were strong significant left-right correlations for the morphometric, geometric and densitometric parameters considered, indicating a high degree of bilateral mechanical symmetry. CONCLUSIONS: The contralateral bone is a suitable control for mechanical intervention studies of the human metacarpal, and the use of bilateral pairing results in an important reduction in sample size. RELEVANCE: Responses to mechanical interventions on the human metacarpal, such as implantation of a metacarpophalangeal prosthesis, are generally unknown. The degree of left-right mechanical symmetry in the human metacarpal provides a measure of the advantage of using paired design studies to address these questions.

Anatomy, Cross-Sectional↗

Quantitative genetics of plastron shape in slider turtles (Trachemys scripta).

Shape variation is widespread in nature and embodies both a response to and a source for evolution and natural selection. To detect patterns of shape evolution, one must assess the quantitative genetic underpinnings of shape variation as well as the selective environment that the organisms have experienced. Here we used geometric morphometrics to assess variation in plastron shell shape in 1314 neonatal slider turtles (Trachemys scripta) from 162 clutches of laboratory-incubated eggs from two nesting areas. Multivariate analysis of variance indicated that nesting area has a limited role in describing plastron shape variation among clutches, whereas differences between individual clutches were highly significant, suggesting a prominent clutch effect. The covariation between plastron shape and several possible maternal effect variables (yolk hormone levels and egg dimensions) was assessed for a subset of clutches and found to be negligible. We subsequently employed several recently proposed methods for estimating heritability from shape variables, and generalized a univariate approach to accommodate unequal sample sizes. Univariate estimates of shape heritability based on Procrustes distances yielded large values for both nesting populations (h2 approximately 0.86), and multivariate estimates of maximal additive heritability were also large for both nesting populations (h2max approximately 0.57). We also estimated the dominant trend in heritable shape change for each nesting population and found that the direction of shape evolution was not the same for the two sites. Therefore, although the magnitude of shape evolution was similar between nesting populations, the manner in which plastron shape is evolving is not. We conclude that the univariate approach for assessing quantitative genetic parameters from geometric morphometric data has limited utility, because it is unable to accurately describe how shape is evolving.

Animals↗

Damage to the human cerebellum from prenatal alcohol exposure: the anatomy of a simple biometrical explanation.

Since 1973, it has become clear that exposure of otherwise normal human fetuses to high levels of alcohol damages a substantial number of the exposed brains in a wide variety of ways nowadays referred to collectively as the fetal alcohol spectrum disorders (FASDs). Averages of images and measurements of brains with these disorders are quantitatively different from normal, and the cerebellum is one of the structures at which differences are typically noted. The present article extends these techniques to a simple, practical, and enlightening detection rule for fetal alcohol damage in adolescents and adults known to have been heavily exposed. The data arise from 180 clinical MR brain images (half of adolescents, half of adults; half male, half female; one-third each fetal alcohol syndrome, fetal alcohol effects, and unexposed). The 180 cerebellums were represented by 328-semilandmark triangulations covering most of the cerebellar surface. Statistical analysis exploited the now-conventional methods of Procrustes analysis in three dimensions, along with a recent extension to incorporate size information explicitly. If we reduce the data complexity even further, to just 23 points along the silhouette of the cerebellum as viewed from above along the aqueductal axis, the analysis becomes more precise. Now a single multivariate summary score, very strongly correlated with size, supports a discrimination (diagnosed vs. unexposed) with about 75% accuracy. About one-quarter of our FASD sample overlaps with the central range of the unexposed in the distribution of this size-based score, with the other three-quarters distinctly showing cerebellar damage. The finding, which corresponds quite closely to the fundamental finding of cerebellar hypoplasia in animal experiments, bears implications for fetal alcohol epidemiology, for geometric morphometrics, and for the geometric complexity of useful data structures derived from clinical brain imaging.

Adolescent↗

Adaptive or non-adaptive? Cranial evolution in a radiation of miniaturized day geckos.

Lygodactylus geckos represent a well-documented radiation of miniaturized lizards with diverse life-history traits that are widely distributed in Africa, Madagascar, and South America. The group has diversified into numerous species with high levels of morphological similarity. The evolutionary processes underlying such diversification remain enigmatic, because species live in different ecological biomes, ecoregions and microhabitats, while suggesting strikingly high levels of homoplasy. To underscore this evolutionary pattern, here we explore the shape variation of skull elements (i.e., cranium, jaw and inner ear) using 3D geometric morphometrics and phylogenetic comparative methods on computed tomography scans (CT-scan) of a sample encompassing almost all recognized taxa within Lygodactylus. The results of this work show that skull and inner ear shape variation is low (i.e., there is high overlapping on the morphospace) across geographic regions, macrohabitats and lifestyles, implying extensive homoplasy. Furthermore, we also found a strong influence of allometry shaping cranial variation both at intra and interspecific levels, suggesting a major constraint underlying skull architecture, probably as a consequence of its miniaturization. The remaining variation that is not allometric is independent of phylogeny and ecological adaptation and can probably be interpreted as the result of intrinsic developmental plasticity. This, in turn, supports the interpretation that speciation in this group is largely concordant with a non-adaptive hypothesis, which results mainly from vicariant processes.

Animals↗

Shape analysis of female facial attractiveness.

Previous studies have suggested that female facial attractiveness is associated with exaggerated sex-specific facial traits and averageness. Here we applied geometric morphometrics, a method for multivariate statistical analysis of shape, to measure geometric averageness and geometric sexual dimorphism of natural female face profiles. Geometric averageness and geometric sexual dimorphism correlate with attractiveness ratings. However, principal component analysis extracted a shape component robustly correlated with attractiveness but independent of sexual dimorphism. The shape differences between attractive- and hyperfeminine traits are localised: attractive facial shape and sexual dimorphism are similar in the upper face, but are markedly distinct in the jaw and chin.

Adolescent↗

Wing shape divergence between Rhodnius prolixus from Cojedes (Venezuela) and Rhodnius robustus from Mérida (Venezuela).

The existence of Rhodnius robustus as a species distinct from Rhodnius prolixus has long been the main epidemiological question about Chagas disease transmission in Venezuela and surrounding countries. These two taxa are morphologically and genetically very similar, but only R. prolixus is assumed to colonize houses and transmit Chagas disease to humans. R. robustus is assumed to be an exclusively sylvatic species, restricted to palm trees. If robustus and prolixus are actually the same species, the theoretical possibility exists of sylvatic specimens invading houses, even after insecticide application, and a control strategy similar to that of the successful Southern Cone Initiative against Triatoma infestans would be difficult to consider. Since no valid alternative control strategy exists, the answer to this biological question could be decisive about the future of vector control in this region. Although we believe genetic techniques are best suited to define species boundaries, we present here an example of the relevance of modern morphometrics in dealing with such an issue. Using both traditional and geometric morphometrics, we compared the wing size and shape in both sexes of these two taxa reared in the same laboratory for one generation. R. robustus specimens were collected from palm trees in the state of Mérida (Venezuela), and R. prolixus were collected from houses in the state of Cojedes (Venezuela). Our study provided no argument to question their specific status. Even after one generation of living in the same laboratory conditions, the two lines showed clear size differences, divergent allometric trends, and significant allometry-free differences in shape. These results suggest that R. robustus (Mérida, Venezuela) and R. prolixus (Cojedes, Venezuela) are distinct evolutionary units. Due to the epidemiological importance of this question, further studies in other geographic areas of Venezuela are required to accurately define the relationships of R. robustus and R. prolixus.

Animals↗

A cephalometric comparison of skulls from different time periods--the Bronze Age, the 19th century and the present.

The aim of this study was to evaluate secular trends by means of orthodontic measurements on lateral cephalograms. We use roentgenograms from three populations: 22 Bronze Age skulls from a cemetery near Hainburg/Austria, 140 soldiers who served in the Hapsburg Imperial Army in the late 19th century, and 154 contemporary recruits of the Austrian Federal Army. Using conventional morphometric analysis, no statistically significant differences could be established. But applying geometric morphometrics to the 2D-coordinates of the pentagon composed of the landmarks Sella, Nasion, Articulare, Gonion and Menton, some biologically interpretable differences were detected, the size allometry between the 19th- and 20th-century populations being the only notable one. We conclude that landmarks should be digitized directly (and many more of them) and that conventional methods used in clinical orthodontics are inappropriate for addressing the scientific questions approached here.

Anthropology, Physical↗

Morphometric analysis of the mandible in subjects with Class III malocclusion.

This study evaluated the deformations that contribute to Class III mandibular configuration, employing geometric morphometric analysis. Lateral cephalograms of male and female groups of 100 young adults and 70 children with Class III malocclusion were compared to those of counterparts with normal occlusion. The sample included an equal number of both genders. The cephalographs were traced, and 12 homologous landmarks were identified and digitized. Average mandibular geometries were generated by means of Procrustes analysis. Thin-plate spline analysis was then applied to mandibular configurations to determine local form differences in male and female groups of adults and children with normal occlusion and Class III mal-occlusion. The mandibular morphology was significantly different between these two groups of male and female adults, and children (p < 0.0001). This spline analysis revealed an anteroposterior elongation of the mandible along the condylion-gnathion axis, showing an extension in the regions of the mandibular condyle and ramus, and of the anteroinferior portion of the mandibular symphysis in Class III groups. More extension was evident in Class III adults. The deformations in subjects with Class III malocclusion may represent a developmental elongation of the mandible anteroposteriorly, which leads to the appearance of a prognathic mandibular profile.

Adolescent↗

Geometric estimates of heritability in biological shape.

The recently developed geometric morphometrics methods represent an important contribution of statistics and geometry to the study of biological shapes. We propose simple protocols using shape distances that incorporate geometric techniques into linear quantitative genetic models that should provide insights into the contribution of genetics to shape variation in organisms. The geometric approaches use Procrustes distances in a curved shape space and distances in tangent spaces within and among families to estimate shape heritability. We illustrate the protocols with an example of wing shape variation in the honeybee, Apis mellifera. The heritability of overall shape variation was small, but some localized components depicting shape changes on distal wing regions showed medium to large heritabilities. The genetic variance-covariance matrix of the geometric shape variables was significantly correlated with the phenotypic shape variance-covariance matrix. A comparison of the results of geometric methods with the traditional multivariate analysis of interlandmark distances indicated that even with a larger dimensionality, the interlandmark distances were not as rich in shape information as the landmark coordinates. Quantitative genetics studies of shape should greatly benefit from the application of geometric methods.

Animals↗

Craniofacial morphometric analysis of mandibular prognathism.

The purpose of this study was to provide more information about the morphological characteristics of the craniofacial complex in mandibular prognathism. Forty young adult males having mandibular prognathism were compared with 40 having normal occlusion. This was conducted to carry out geometric morphometric assessments to localize alterations, using Procrustes analysis and thin-plate spline analysis, in addition to conventional cephalometric techniques. Procrustes analysis indicated that the mean craniofacial, midfacial and mandibular morphology was significantly different in prognathic subjects compared with normal controls. This finding was corroborated by the multivariate Hotelling T(2)-test of cephalometric variables. Mandibular prognathism demonstrated a shorter and slightly retropositioned maxilla, a greater total length and anterior positioning of the mandible. Thin-plate spline analysis revealed a developmental diminution of the palatomaxillary region anteroposteriorly and a developmental elongation of the mandible anteroposteriorly, leading to the appearance of a prognathic mandibular profile. In conclusion, thin-plate spline analysis seems to provide a valuable supplement for conventional cephalometric analysis because the complex patterns of craniofacial shape change are visualized suggestive by means of grid deformations.

Adult↗

New approaches in hominoid taxonomy: morphometrics.

We report here on new cranial data relevant to hominoid taxonomic analyses, based on a study of 438 skulls belonging to 13 nonhuman living hominoid taxa. Nineteen landmarks were selected to describe the overall shape of the maxillofacial complex, in order to investigate its discriminative power in taxonomic analyses. We used a geometric morphometrics approach to depict morphological variation from the genus down to the subspecific level, and we evaluated whether our morphologic criteria are relevant to discriminating species and subspecies among living hominoids. Considering previous genetic studies, we discuss whether our results can be extrapolated to the hominin fossil record, providing a reference for species and subspecies morphologic differentiation. Our results indicate that the relative warp method, as applied to facial landmarks, provides a powerful tool to discriminate taxa down to a subspecific level. Results show a noticeable divergence of P. t. verus compared to P. t. troglodytes and P. t. schweinfurthii. According to our data, the distance between eastern and western gorilla populations as well as between Bornean and Sumatran orangutan subspecies is as great as between the two species of Pan. In the same manner, differences between Hylobates and Symphalangus are similar to those between Pan and Gorilla genera. Congruence between the morphological distances computed in this study and previous morphological and genetical studies strongly supports their relevance for morphological species recognition in paleoanthropology. Our data provide an objective standard for assessing taxonomic differences among hominoids, and will enable us to define more precisely the significance of morphological differences in the fossil record.

Animals↗

3D morphometrics of craniofacial dysmorphology reveals sex-specific asymmetries in schizophrenia.

Over early fetal life cerebral and craniofacial morphogenesis proceed in embryological intimacy. Therefore, craniofacial shape differences between schizophrenia patients and controls are informative of developmental disturbance(s) in cerebral-craniofacial morphogenesis. 3D craniofacial coordinates were calculated from interlandmark distances for 169 patients with DSM-III-R schizophrenia and 78 matched normal controls. These were analysed using geometric morphometrics with visualisation of the resultant statistical models. Patients of both sexes were characterised by an intricate topography of 3D shape change involving lengthened lower mid-facial height, shortened upper mid-facial height, nasion located posteriorly and a wider face posteriorly; there was sex-specific rotation of the midface such that the base of the nose is more anterior in female patients but more posterior in male patients. Importantly, there were sex-specific asymmetries: in males, controls evidenced marked directional asymmetry while patients showed reduced directional asymmetry; conversely, in females controls evidenced little directional asymmetry while patients showed marked directional asymmetry. In schizophrenia, the topography of craniofacial dysmorphology appears to reflect subtle disruption to a critical 3D trajectory of embryonic-fetal craniofacial growth, particularly along the midline, with disturbance to the establishment of normal asymmetries in a sex-related manner.

Adult↗

Morphometrics and hominoid phylogeny: Support for a chimpanzee-human clade and differentiation among great ape subspecies.

Taxonomic and phylogenetic analyses of great apes and humans have identified two potential areas of conflict between molecular and morphological data: phylogenetic relationships among living species and differentiation of great ape subspecies. Here we address these problems by using morphometric data. Three-dimensional landmark data from the hominoid temporal bone effectively quantify the shape of a complex element of the skull. Phylogenetic analysis using distance-based methods corroborates the molecular consensus on African ape and human phylogeny, strongly supporting a Pan-Homo clade. Phenetic differentiation of great ape subspecies is pronounced, as suggested previously by mitochondrial DNA and some morphological studies. These results show that the hominoid temporal bone contains a strong phylogenetic signal and reveal the potential for geometric morphometric analysis to shed light on phylogenetic relationships.

Animals↗

Life history dependent morphometric variation in stream-dwelling Atlantic salmon.

The time course of morphometric variation among life histories for stream-dwelling Atlantic salmon ( Salmo salar L.) parr (age-0+ to age-2+) was analyzed. Possible life histories were combinations of parr maturity status in the autumn (mature or immature) and age at outmigration (smolt at age-2+ or later age). Actual life histories expressed with enough fish for analysis in the 1997 cohort were immature/age-2+ smolt, mature/age-2+smolt, and mature/age-2+ non-smolt. Tagged fish were assigned to one of the three life histories and digital pictures from the field were analyzed using landmark-based geometric morphometrics. Results indicated that successful grouping of fish according to life history varied with fish age, but that fish could be grouped before the actual expression of the life histories. By March (age-1+), fish were successfully grouped using a descriptive discriminant function and successful assignment ranged from 84 to 97% for the remainder of stream residence. A jackknife of the discriminant function revealed an average life history prediction success of 67% from age-1+ summer to smolting. Low sample numbers for one of the life histories may have limited prediction success. A MANOVA on the shape descriptors (relative warps) also indicated significant differences in shape among life histories from age-1+ summer through to smolting. Across all samples, shape varied significantly with size. Within samples, shape did not vary significantly with size for samples from December (age-0+) to May (age-1+). During the age-1+ summer however, shape varied significantly with size, but the relationship between shape and size was not different among life histories. In the autumn (age-1+) and winter (age-2+), life history differences explained a significant portion of the change in shape with size. Life history dependent morphometric variation may be useful to indicate the timing of early expressions of life history variation and as a tool to explore temporal and spatial variation in life history expression.

Adaptation, Physiological↗

Functional and historical determinants of shape in the scapula of Xenarthran mammals: evolution of a complex morphological structure.

The mammalian scapula is a complex morphological structure, composed of two ossification plates that fuse into a single structure. Most studies on morphological differentiation in the scapula have considered it to be a simple, spatially integrated structure, primarily influenced by the important locomotor function presented by this element. We used recently developed geometric morphometric techniques to test and quantify functional and phylogenetic influences on scapular shape variation in fossil and extant xenarthran mammals. The order Xenarthra is well represented in the fossil record and presents a stable phylogenetic hypothesis for its genealogical history. In addition, its species present a large variety of locomotor habits. Our results show that approximately half of the shape variation in the scapula is due to phylogenetic heritage. This is contrary to the view that the scapula is influenced only by functional demands. There are large-scale shape transformations that provide biomechanical adaptation for the several habits (arboreality, terrestriality, and digging), and small scale-shape transformations (mostly related to the coracoid process) that are not influenced by function. A nonlinear relationship between morphometric and phylogenetic distances indicates the presence of a complex mixture of evolutionary processes acting on shape differentiation of the scapula.

Animals↗

Variation and causal factors of craniofacial robusticity in Patagonian hunter-gatherers from the late Holocene.

Fueguian-Patagonian skulls have been characterized as some of the most robust of any modern crania. However, the causal factors of such robusticity remain unsettled. We assess within- and among-sample cranial robusticity of seven samples from continental Patagonia and Tierra del Fuego, using geometric morphometric techniques. In addition, the biomechanical, phylogenetic, and climatic hypotheses proposed to account for robusticity in such samples are discussed. Two Amerindian samples of farmers and two early middle Holocene samples from South America were included. The results show: 1) large variation in craniofacial robusticity among Patagonian samples, with the highest robusticity in samples from south continental Patagonia and Isla Grande of Tierra del Fuego, whereas central and north Patagonian samples display the same degree of robusticity as farmer samples; 2) that early middle Holocene samples display lower levels of robusticity than South Patagonian samples; and 3) strong association between latitude and craniofacial robusticity, with the most robust craniofacial morphologies occurring at the highest latitudes. In consequence, neither masticatory stress nor retention of ancestral features is supported by the morphological evidence analyzed. Hence it is hypothesized that endocrine changes related to cold climate may be a plausible explanation for several craniofacial features found in Fueguian and south continental Patagonian samples, such as their large masticatory component, and pronounced supraorbital ridge and glabellar region.

Adaptation, Physiological↗

Thin-plate spline analysis of allometry and sexual dimorphism in the human craniofacial complex.

The relationship between allometry and sexual dimorphism in the human craniofacial complex was analyzed using geometric morphometric methods. Thin-plate splines (TPS) analysis has been applied to investigate the lateral profile of complete adult skulls of known sex. Twenty-nine three-dimensional (3D) craniofacial and mandibular landmark coordinates were recorded from a sample of 52 adult females and 52 adult males of known age and sex. No difference in the influence of size on shape was detected between sexes. Both size and sex had significant influences on shape. As expected, the influence of centroid size on shape (allometry) revealed a shift in the proportions of the neurocranium and the viscerocranium, with a marked allometric variation of the lower face. Adjusted for centroid size, males presented a relatively larger size of the nasopharyngeal space than females. A mean-male TPS transformation revealed a larger piriform aperture, achieved by an increase of the angulation of the nasal bones and a downward rotation of the anterior nasal floor. Male pharynx expansion was also reflected by larger choanae and a more posteriorly inclined basilar part of the occipital clivus. Male muscle attachment sites appeared more pronounced. In contrast, the mean-female TPS transformation was characterized by a relatively small nasal aperture. The occipital clivus inclined anteriorly, and muscle insertion areas became smoothed. Besides these variations, both maxillary and mandibular alveolar regions became prognathic. The sex-specific TPS deformation patterns are hypothesized to be associated with sexual differences in body composition and energetic requirements.

Adult↗