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Swift laboratory thermal evolution of wing shape (but not size) in Drosophila subobscura and its relationship with chromosomal inversion polymorphism.

Latitudinal clinal variation in wing size and shape has evolved in North American populations of Drosophila subobscura within about 20 years since colonization. While the size cline is consistent to that found in original European populations (and globally in other Drosophila species), different parts of the wing have evolved on the two continents. This clearly suggests that 'chance and necessity' are simultaneously playing their roles in the process of adaptation. We report here rapid and consistent thermal evolution of wing shape (but not size) that apparently is at odds with that suggestion. Three replicated populations of D. subobscura derived from an outbred stock at Puerto Montt (Chile) were kept at each of three temperatures (13, 18 and 22 degrees C) for 1 year and have diverged for 27 generations at most. We used the methods of geometric morphometrics to study wing shape variation in both females and males from the thermal stocks, and rates of genetic divergence for wing shape were found to be as fast or even faster than those previously estimated for wing size on a continental scale. These shape changes did not follow a neat linear trend with temperature, and are associated with localized shifts of particular landmarks with some differences between sexes. Wing shape variables were found to differ in response to male genetic constitution for polymorphic chromosomal inversions, which strongly suggests that changes in gene arrangement frequencies as a response to temperature underlie the correlated changes in wing shape because of gene-inversion linkage disequilibria. In fact, we also suggest that the shape cline in North America likely predated the size cline and is consistent with the quite different evolutionary rates between inversion and size clines. These findings cast strong doubts on the supposed 'unpredictability' of the geographical cline for wing traits in D. subobscura North American colonizing populations.

Analysis of Variance↗

Differential correlates of diet and phylogeny on the shape of the premaxilla and anterior tooth in sparid fishes (Perciformes: Sparidae).

We explore the correlational patterns of diet and phylogeny on the shape of the premaxilla and anterior tooth in sparid fishes (Perciformes: Sparidae) from the western Mediterranean Sea. The premaxilla is less variable, and in spite of the presence of species-specific features, a common structural pattern is easily recognizable in all species (i.e. the ascending and the articular processes are fused in a single branch, as in many percoid fishes). In contrast, tooth shape is more variable, and different structural types can be recognized (e.g. canine-like or incisive). Coupling geometric morphometric and comparative methods we found that the relationship between shape, diet and phylogeny also differs between premaxilla and tooth. Thus, the shape of the premaxilla is significantly correlated with food type, whereas the shape of the teeth is not correlated with diet, and probably reflects the species phylogenetic relationships. Two biological roles, resistance against compressive forces generated in the buccal cavity and the size of the oral gape, would explain the ecomorphological patterns of the premaxilla. The premaxilla and anterior tooth appear to evolve at different rates (mosaic evolution) and represent an example of morphological traits belonging to the same functional unit but following uncoupled evolutionary pathways.

Adaptation, Biological↗

Heat stress and age induced maternal effects on wing size and shape in parthenogenetic Drosophila mercatorum.

Maternal effects on progeny wing size and shape in a homozygous parthenogenetic strain of Drosophila mercatorum were investigated. The impact of external maternal factors (heat stress) and the impact of internal maternal factors (different maternal and grand maternal age) were studied. The offspring developed under identical environmental conditions, and due to lack of genetic variation any phenotypic difference among offspring could be ascribed to maternal effects. Wing size was estimated by centroid size, shape was analysed with the Procrustes geometric morphometric method and variation in landmark displacement was visualized by principal component analysis. Both kinds of maternal effects had a significant impact on progeny wing size and shape. Maternal heat stress led to the same pattern of response in size and shape among the progeny, with increased difference between the control group and progeny from heat stressed flies in both size and shape with increased maternal heat stress temperature. The effects of maternal age, however, led to different responses in size and shape between the different progeny groups. The observed variation in landmark displacements was similar, and in both cases mainly associated with shape differences of the posterior part of the wing. Finally, our results suggest that maternal effect has some evolutionary implications by altering the genetic correlations among traits, which can affect the response to selective pressures.

Age Factors↗

Parallel evolution of the sexes? Effects of predation and habitat features on the size and shape of wild guppies.

Environmental gradients often lead to the parallel evolution of populations and species. To what extent do such gradients also lead to parallel evolution of the sexes? We used guppies (Poecilia reticulata) to examine the parallel and independent (sex-specific) aspects of population divergence in response to predation and habitat features. Geometric morphometrics was used to analyse size and shape variation for 1335 guppies from 27 to 31 sites sampled in each of 2 years. Body size showed strong parallel population divergence; both sexes were larger at sites with a more open canopy and with higher flow. Body shape showed a mixture of parallel and independent population divergence. The strongest and most consistent effects were (1) high-predation sites had males with smaller heads and deeper caudal peduncles, (2) open-canopy sites had females with smaller heads and more distended abdomens and (3) high-flow sites had males and females with smaller heads and deeper caudal peduncles.

Animals↗

Adaptive radiation in African weakly electric fish (Teleostei: Mormyridae: Campylomormyrus): a combined molecular and morphological approach.

We combined multiple molecular markers and geometric morphometrics to revise the current taxonomy and to build a phylogenetic hypothesis for the African weakly electric fish genus Campylomormyrus. Genetic data (2039 bp DNA sequence of mitochondrial cytochrome b and nuclear S7 genes) on 106 specimens support the existence of at least six species occurring in sympatry. We were able to further confirm these species by microsatellite analysis at 16 unlinked nuclear loci and landmark-based morphometrics. We assigned them to nominal taxa by comparisons to type specimens of all Campylomormyrus species recognized so far. Additionally, we showed that the shape of the elongated trunk-like snout is the major source of morphological differentiation among them. This finding suggests that the radiation of this speciose genus might have been driven by adaptation to different food sources.

Adaptation, Biological↗

Craniofacial levels and the morphological maturation of the human skull.

It is well known that the human skull achieves adult size through a superior-inferior gradient of maturation. Because the basicranium matures in size before the face, it has been suggested that the form of the basicranium might have ontogenetic knock-on effects on that of the face. However, although sequential spatially organized maturation of size is well described in the cranium, the maturation of skull shape is not. Knowledge of the maturation of shape is important, nevertheless, because it is claimed that the early determination of the spatial configuration of basicranial components, where the facial skeleton attaches, is relevant in the spatio-temporal ontogenetic cascade from basicranium to face. This paper examines the ontogeny of various components of the human skull in 28 individuals from the longitudinal Denver Growth Study. Sixty-six landmarks and semilandmarks were digitized on 228 X-rays and analysed using geometric morphometric methods. Bootstrapped confidence intervals for centroid size support previous studies suggesting a supero-inferior gradient of growth maturation (size over time), while developmental maturation (shape over time) is more complex. A sequence of shape maturation is described, in which the earliest structure to mature in shape was the midline cranial base (7-8 years), followed by the lateral cranial floor (11-12), midline neurocranium (9-10) and facial and mandibular structures (15-16). The absolute ages of shape maturation of the latter three depended on the criterion of maturity used, which was not the case for the basicranial components. Additionally, ontogenetic dissociations were found between the maturation of size and shape of the midline cranial base and lateral floor, possibly underlining its role as structural 'interface' between brain and facial ontogeny. These findings imply potential for bidirectional developmental influences between the lateral cranial floor and the face until about 11-12 years. The findings are discussed with regard to their relevance for palaeoanthropology and especially the evolutionary and developmental bases of skull morphological variation.

Adolescent↗

Developmental regulation of skull morphology. I. Ontogenetic dynamics of variance.

In the absence of processes regulating morphogenesis and growth, phenotypic variance of a population experiencing no selective mortality should increase throughout ontogeny. To determine whether it does, we measure variance of skull shape using geometric morphometrics and examine its ontogenetic dynamics in the precocial cotton rat (Sigmodon fulviventer) and the altricial house mouse (Mus musculus domesticus). In both species, variance of shape halves between the two youngest samples measured (between 1 and 10 days postnatal and 10 and 15 days postnatal, respectively) and thereafter is nearly constant. The reduction in variance did not appear to result from a general regulation of skull size or developmental timing, although skull size may also be regulated and developmental timing is an important component of the variation in skull shape of young house mice. The ontogenetic dynamics of variance suggest two possible scenarios. First, variation generated during fetal or early postnatal growth is not immediately compensated and therefore accumulates, whereas later in growth, variation is continually generated and rapidly compensated. Second, variation generated during fetal and early postnatal growth is rapidly compensated, after which no new variance is produced. Based on a general model for bone growth, we hypothesize that variance is generated when bone grows under the direction of disorganized muscular movements and decreases with increasing neuromuscular control. Additionally, increasing coherence of signals transmitted by the growing brain and sensory organs, which exert tensile forces on bone, may also canalize skull shape.

Anatomy↗

Epigenetic interactions and the structure of phenotypic variation in the cranium.

Understanding the developmental and genetic basis for evolutionarily significant morphological variation in complex phenotypes such as the mammalian skull is a challenge because of the sheer complexity of the factors involved. We hypothesize that even in this complex system, the expression of phenotypic variation is structured by the interaction of a few key developmental processes. To test this hypothesis, we created a highly variable sample of crania using four mouse mutants and their wild-type controls from similar genetic backgrounds with developmental perturbations to particular cranial regions. Using geometric morphometric methods we compared patterns of size, shape, and integration in the sample within and between the basicranium, neurocranium, and face. The results highlight regular and predictable patterns of covariation among regions of the skull that presumably reflect the epigenetic influences of the genetic perturbations in the sample. Covariation between relative widths of adjoining regions is the most dominant factor, but there are other significant axes of covariation such as the relationship between neurocranial size and basicranial flexion. Although there are other sources of variation related to developmental perturbations not analyzed in this study, the patterns of covariation created by the epigenetic interactions evident in this sample may underlie larger scale evolutionary patterns in mammalian craniofacial form.

Animals↗

A comparative study of adult facial morphology and its ontogeny in the fossil macaque Macaca majori from Capo Figari, Sardinia, Italy.

This study examines the morphology of the face in the fossil macaque Macaca majori from Capo Figari (north-eastern Sardinia, Italy) in a comparative ontogenetic context. Thus, a fairly complete face from an adult representative of this fossil species is compared with 3 extant macaque species: Macaca sylvanus (of which species it is questioned whether it is a subspecies, M. sylvanus majori), Macaca mulatta and Macaca fascicularis. Additional incomplete subadult and adult specimens are also examined in order to compare their facial ontogeny with that of the same living species. The comparisons are based on facial landmark data and are undertaken using geometric morphometric methods. These studies indicate that the adult facial morphology and ontogeny of face size and shape in M. majori share much in common with extant macaque species. However, the adult M. majori face displays some unique morphological features, in particular with regard to lateral flaring and relative size of the zygomatic roots. From the study of a limited sample of fossils there is an indication that this flaring arises during postnatal growth, and in consequence the ontogeny of the face of this fossil species may be different from that of M. sylvanus and the other macaque species included in this analysis. From these studies, we conclude that M. majori shows differences in adult facial morphology and possibly in ontogeny from M. sylvanus compatible with a specific rather than subspecific distinction.

Animals↗

Facial shape and asymmetry by three-dimensional laser surface scanning covary with cognition in a sexually dimorphic manner.

The embryological unity over early fetal life of the anterior brain, neuroepithelium, neural crest, and facial ectoderm is responsible for facial dysmorphogenesis in disorders of early brain development, including schizophrenia. This study examined covariance of facial shape and asymmetry with cognition in a normal sample of 36 men and 51 women using geometric morphometrics. Facial shape and asymmetry covaried with verbal and visual spatial cognitive functions in a sexually dimorphic manner. Events over early fetal life may be an important determinant of sexually dimorphic covariance of anterior facial shape and asymmetry with aspects of cognition that involve the anterior brain.

Adult↗

Genetics and geometry of canalization and developmental stability in Drosophila subobscura.

BACKGROUND: Many properties of organisms show great robustness against genetic and environmental perturbations. The terms canalization and developmental stability were originally proposed to describe the ability of an organism to resist perturbations and to produce a predictable target phenotype regardless of random developmental noise. However, the extent to which canalization and developmental stability are controlled by the same set of genes and share underlying regulatory mechanisms is largely unresolved. RESULTS: We have analyzed the effects of clinical genetic variation (inversion polymorphism) on wing asymmetry by applying the methods of geometric morphometrics in the context of quantitative genetics using isochromosomal lines of Drosophila subobscura. For the analysis of overall size, developmental stability was positively correlated with levels of heterozygosity and development at the optimal temperature. For analyses of shape, the overall comparisons by matrix correlations indicate that inter- and intraindividual variation levels were poorly correlated, a result also supported when comparing the vectors describing patterns of variation of landmark position. The lack of similarity was basically due to the discrepancy between the genetic and environmental components of the interindividual variation. Finally, the analyses have also underscored the presence of genetic variation for directional asymmetry. CONCLUSIONS: The results strongly support the hypothesis that environmental canalization and developmental stability share underlying regulatory mechanisms, but environmental and genetic canalization are not functionally the same. A likely explanation for this lack of association is that natural wing shape variation in Drosophila populations is loosely related to individual fitness.

Analysis of Variance↗

Epidermal growth factor receptor and transforming growth factor-beta signaling contributes to variation for wing shape in Drosophila melanogaster.

Wing development in Drosophila is a common model system for the dissection of genetic networks and their roles during development. In particular, the RTK and TGF-beta regulatory networks appear to be involved with numerous aspects of wing development, including patterning, cell determination, growth, proliferation, and survival in the developing imaginal wing disc. However, little is known as to how subtle changes in the function of these genes may contribute to quantitative variation for wing shape, per se. In this study 50 insertional mutations, representing 43 loci in the RTK, Hedgehog, TGF-beta pathways, and their genetically interacting factors were used to study the role of these networks on wing shape. To concurrently examine how genetic background modulates the effects of the mutation, each insertion was introgressed into two wild-type genetic backgrounds. Using geometric morphometric methods, it is shown that the majority of these mutations have profound effects on shape but not size of the wing when measured as heterozygotes. To examine the relationships between how each mutation affects wing shape hierarchical clustering was used. Unlike previous observations of environmental canalization, these mutations did not generally increase within-line variation relative to their wild-type counterparts. These results provide an entry point into the genetics of wing shape and are discussed within the framework of the dissection of complex phenotypes.

Animals↗

Integration and modularity of quantitative trait locus effects on geometric shape in the mouse mandible.

The mouse mandible has long served as a model system for complex morphological structures. Here we use new methodology based on geometric morphometrics to test the hypothesis that the mandible consists of two main modules, the alveolar region and the ascending ramus, and that this modularity is reflected in the effects of quantitative trait loci (QTL). The shape of each mandible was analyzed by the positions of 16 morphological landmarks and these data were analyzed using Procrustes analysis. Interval mapping in the F(2) generation from intercrosses of the LG/J and SM/J strains revealed 33 QTL affecting mandible shape. The QTL effects corresponded to a variety of shape changes, but ordination or a parametric bootstrap test of clustering did not reveal any distinct groups of QTL that would affect primarily one module or the other. The correlations of landmark positions between the two modules tended to be lower than the correlations between arbitrary subsets of landmarks, indicating that the modules were relatively independent of each other and confirming the hypothesized location of the boundary between them. While these results are in agreement with the hypothesis of modularity, they also underscore that modularity is a question of the relative degrees to which QTL contribute to different traits, rather than a question of discrete sets of QTL contributing to discrete sets of traits.

Animals↗

Virtual anthropology: the digital evolution in anthropological sciences.

The discovery and explanation of differences among organisms is a major concern for evolutionary and systematic biologists. In physical anthropology, the discrimination of taxa and the qualitative and quantitative description of ontogenetic or evolutionary change require, of course, the analysis of morphological features. Since the 1960s, a remarkable amount of fossil material was excavated, some of it still awaiting a detailed first analysis, some of it requiring re-examination by more developed methods. While the fossil record grew continuously, a revolution in anthropological research took place with advances in computer technology in the 1980s: a handful of innovative researchers working in specialized anthropology laboratories or medical departments developed the methodological inventory needed to extract critical information from subjects in vivo and from fossilized remains. A considerable part of this information is preserved in the physically heretofore inaccessible interior of anatomical structures. Virtual Anthropology (VA) is a means of making them visible and measurable. Thus, VA also allows access to 'hidden' landmarks; in addition, the large number of semilandmarks accessible on the form enhances the power of Geometric Morphometrics analysis. Furthermore, the density information in volume data allows manipulations such as segmentation, impossible with the real, physical object. Moreover, metric body measurements generally, and cranial measurements specifically, are also an important source of information for the analysis of the ontogenetic development of the skeletal system, and--last but not least--for clinical use (e.g., operation planning, operation simulation, prosthetics). Thus, there developed a fruitful interdisciplinary cooperation between statistics, medicine, and physical anthropology.

Anthropology, Physical↗

New methods and techniques in anthropology.

Since the discovery of the Tyrolean Iceman in 1991, advanced imaging and post-processing techniques have been successfully applied to anthropological research. Among the specific techniques are spiral computed tomography and 3-dimensional reconstructions, which include stereolithographic and fused deposition modeling of volume data sets. The Iceman's skull was the first to be produced using stereolithography; subsequently, it has been successfully applied in preoperative planning. With the advent of high-end performance graphics workstations and biomedical image processing software packages, 3-dimensional reconstructions have become established as routine tools for analyzing volume data sets. These techniques enabled dramatically new insights to be gained in the field of physical anthropology. Computed tomography became the ideal research tool to access the internal structures of various precious fossils without even touching--let alone damaging--them. Among the most precious are specimens from the genus Australopithecus (1.8 Myr-3.5 Myr), as well as representatives of Homo heidelbergensis (200 kyr-600 kyr) and Homo neanderthalensis (40 kyr-100 kyr); such fossils have been CT-scanned during the last five years. The fossils often are filled with a stone matrix or other encrustations. During the post-processing routines, highly advanced algorithms were used to remove these encrustations virtually (the concrete fossils remain untouched). Thus it has been possible to visualize the morphological structures that are hidden by the matrix layer. Some specimens have been partially destroyed, but it has been possible for the missing parts were reconstructed on the computer screen in order to get estimations of brain volume and endocranial morphology, both major fields of interest in physical anthropology. Moreover, the data in computerized form allows new descriptions of morphological structures using geometric morphometrics. Some of the results may change aspects and interpretations in human evolution and approaches to long-standing questions in this field. We subsume the introduction of these new imaging and post-processing techniques into a new field of research: Virtual Anthropology.

Animals↗

On morphological clocks and paleophylogeography: towards a timescale for Sorex hybrid zones.

Phylogeography--the study of within-species phylogenetic and geographic divergence--has been primarily the domain of molecular evolutionists because molecular markers record population structure on smaller scales than do traditional morphological traits. But when geometric morphometrics are combined with distance-based phylogenetics molar shape divergence appears to record population-level phylogeny, a fact that allows extant and fossil populations to be combined in a single phylogeographic study. The European Sorex araneus complex--a genetically complicated group composed of multiple karyotypic races and species--illustrates the principle. The phylogeographic patterns revealed by molar shape broadly agree with scenarios based on molecular data and circumstantial evidence. Importantly, the inclusion of fossil samples of known age allows minimum divergence times to be inferred. Some races of S. araneus may have diverged more than 120,000 years ago, but others may have diverged less than 14,000. Supporting evidence that molar shape can be used to reconstruct phylogeographic relationships comes from strong correlations between molar shape distances and both phylogenetic divergence time and cytochrome b sequence divergence in datasets where these variables are known independently (fossil carnivorans from a well-constrained stratigraphic setting and shrew species of the genus Sorex, respectively). However, molar shape may have a 'saturation point' beyond which it is not applicable.

Animals↗

Population affinities of 19th Century Cuban crania: implications for identification criteria in South Florida Cuban Americans.

Identification criteria, specifically discriminant function formulae derived from traditional craniometrics, currently used in South Florida for Cuban Americans and other "Hispanic" groups, are unsuitable to provide adequate biological profiles due to complex biological histories as well as widely diverse geographic origins. Florida's total population is approximately 16 million (15,982,378) individuals. Of the total population 2.682,715, or 16.8%, are self-identified as "Hispanic". South Florida (herein defined as Miami-Dade, Broward and Collier Counties) is home to 60% of the total Hispanic population of Florida with 1,291,737 (48.15%) residing in Miami-Dade County. The Hispanic population of Miami-Dade County makes up 57.0% of the total population of 2,253,362. Each recognized sub-group of Hispanics (Mexican, Puerto Rican, and Cuban) includes its own geographic point-of-origin and population history. Cuban-Americans (arriving in the late 1950's and early 1960's) make up the largest sub-population of Florida's Hispanics in any county and in Miami-Dade number 650,601 or 51% of the total Latin population. Additionally, as in other agricultural states, Florida has a very large population of undocumented workers who primarily arrive from Texas and points south of the Straits of Florida. Thus the application of the available traditional craniometric and non-metric methods are not appropriate for South Florida's Latin population. To begin to address this issue in relation to South Florida's Cuban population, we present an analysis of cranio-facial shape variation in a 19th Century Cuban sample, 17th Century Spanish sample, a Precontact Cuban sample, and Terry Blacks using geometric morphometric methods. Significant biological shape differences and patterns of variation are observed among the groups. These results provide us with a context in which to begin to understand the biological variation of Cuban Americans, which will enable the development of identification criteria specific for this U.S. hybrid Hispanic community.

Anthropology, Physical↗

Discrimination of Aedes aegypti (Diptera: Culicidae) laboratory lines based on wing geometry.

Recent techniques of geometric morphometrics were applied to the study of four Thailand laboratory lines of the dengue vector Aedes aegypti. These lines differed by their geographic origin and the number of generations spent in the laboratory. Using a phase contrast microscope, a set of sixteen landmarks of the wings could be identified. A subsequent Procrustean superposition analysis produced the corresponding size and shape variables which were statistically analyzed. Size showed a significant decrease with the number of generations spent in the laboratory. Shape allowed an almost perfect reclassification of the specimens. This tool is able to accurately distinguish different laboratory lines and will be useful in field studies related to entomological surveillance and vector control.

Aedes↗