The use of otolith shape to determine stock structure of <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis> along the Tunisian coast

Engraulis encrasicolus is of great economic importance in the Mediterranean. However, little is known about its stock structure. Otolith shape analysis has been successfully used for fish stock identification. In this study, the stock structure of anchovy caught off the open sea and the coastal area of the Gulf of Tunis, lagoon of Bizerte and Lake of Ichkeul were investigated using otolith shape. Otolith shape was determined by Fourier analysis and then compared among specimens sampled from different areas with forward stepwise canonical discriminant analysis. Significant differences in otolith shape between the open sea and inshore anchovy groups were detected. Otolith shape of anchovy collected in the Lake of Ichkeul was distinct from the other groups. This finding suggests a clear discreteness of the open sea and the continental groups. The data highlighted the potential for using otolith shape analysis for anchovy stock identification, as well as the role of oceanographic features in determining stock separation. These findings will have major implications for anchovy fisheries management in Tunisia. By using a precautionary approach and considering the three areas as separate stocks, fisheries management strategies should be adjusted to achieve optimum sustainable production from each stock and to avoid decreases in genetic variety.     

Size and shape variation in Australopithecus afarensis proximal femora

The degree of size and shape variation in the A. afarensis fossil sample has been interpreted in a variety of ways. Size variation has been described as exceeding that of extant hominoids, similar to that of strongly sexually dimorphic hominoids, and best matched to modern humans. The degree of shape variation has been characterized both as great and negligible. Recent fieldwork has increased the proximal femoral sample, providing new data with which to examine variation. The proximal femur of A. afarensis is analyzed in a comparative framework in order to gauge the magnitude of size and shape variation in this element.Seven of the best-preserved A. afarensis proximal femora contribute to the analysis (A.L. 128-1, A.L. 152-2, A.L. 211-1, A.L. 288-1ap, A.L. 333-3, A.L. 333-123, A.L. 827-1). Comparative samples from Pan, Pongo, Gorilla, and Homo provide context for interpreting variation among the fossils. The coefficient of variation (CV) of linear measurements is used to estimate size variation. Bootstrap resampling of CVs from extant hominoids provides distributions for comparison to A. afarensis CVs. Ratios of linear measurements provide scale-free shape variables that are used in pairwise comparisons. The Euclidean distance between pairs of A. afarensis are compared to the Euclidean distances between extant hominoid pairs.As found in some earlier analyses, size variation in A. afarensis is accommodated best in gorillas and orangutans. The magnitude of difference in shape between A. afarensis pairs is exceeded by most taxa, indicating that shape variation is not extreme. These general findings are contradicted by a few instances of excessive size and shape variation. These are uncharacteristic results and could point to temporal bias, although other alternatives are explored. The signal from the proximal femur is that size variation in A. afarensis is like that of the strongly sexually dimorphic apes, and shape variation is well within the range of most hominoids irrespective of their degree of size dimorphism.     

Hypoxia drives plastic divergence in cichlid body shape

Organisms experience multiple selective agents that can influence phenotypes through heritable and/or plastic changes, often reflecting complex interactions between phenotype and environment. Environmental factors can directly influence phenotypes, but also indirectly affect phenotypic variation when genetic/plastic change in one trait results in correlated genetic/plastic change in another trait. In fishes, body shape is a trait that might be particularly prone to influence from environmental pressures that act on other morphological features. Variation in dissolved oxygen among aquatic environments has a large impact on the size of the gills and brains of fishes. It is likely that dissolved oxygen interacts with other environmental factors to both directly and indirectly influence patterns of body shape variation. We examined effects of dissolved oxygen on body shape variation among populations of an African cichlid fish (Pseudocrenilabrus multicolor) from multiple high- and low-oxygen sites within a single drainage in Uganda. A split-brood laboratory experiment was used to estimate plasticity of gill and brain size, and we used morphometric analyses to identify variation in body shape in F₁ offspring. Several analyses enabled us to identify genetic effects among populations, and effects of oxygen acting either directly on body shape or indirectly through its effects on gill and brain size. A large part of the variation in body shape was due to plastic variation in gill size associated with dissolved oxygen. Fish raised under low oxygen had deeper heads and shorter bodies, and this variation was driven by both direct effects of oxygen and indirect effects of gill size variation. Body shape variation in fishes should reflect interacting effects of multiple environmental factors that act directly or indirectly on morphology. Body shape might be particularly difficult to predict when phenotypes are plastic, because changes among populations would occur rapidly and be unrelated to genetic variation.     

Head shape evolution in monitor lizards (Varanus): interactions between extreme size disparity,phylogeny and ecology

Characterizing patterns of observed current variation, and testing hypotheses concerning the potential drivers of this variation, is fundamental to understanding how morphology evolves. Phylogenetic history, size and ecology are all central components driving the evolution of morphological variation, but only recently have methods become available to tease these aspects apart for particular body structures. Extant monitor lizards (Varanus) have radiated into an incredible range of habitats and display the largest body size range of any terrestrial vertebrate genus. Although their body morphology remains remarkably conservative, they have obvious head shape variation. We use two‐dimensional geometric morphometric techniques to characterize the patterns of dorsal head shape variation in 36 species (375 specimens) of varanid, and test how this variation relates to size, phylogenetic history and ecology as represented by habitat. Interspecific head shape disparity is strongly allometric. Once size effects are removed, principal component analysis shows that most shape variation relates to changes in the snout and head width. Size‐corrected head shape variation has strong phylogenetic signal at a broad level, but habitat use is predictive of shape disparity within phylogenetic lineages. Size often explains shape disparity among organisms; however, the ability to separate size and shape variation using geometric morphometrics has enabled the identification of phylogenetic history and habitat as additional key factors contributing to the evolution of head shape disparity among varanid lizards.     

The role of diet and temperature in shaping cranial diversification of South American human populations: an approach based on spatial regression and divergence rate tests

Aim Understanding the importance of ecological factors in the origin and maintenance of patterns of phenotypic variation among populations, in an explicit geographical context, is one of the main goals of human biology, ecology and evolutionary biology. Here we study the ecological factors responsible for craniofacial variation among human populations from South America. Location South America. Methods We studied a dataset of 718 males from 40 South American populations, coming from groups that inhabited different geographical and ecological regions. Cranial size and shape variation were studied using 30 cranial measurements. We first used spatial correlograms and interpolated maps to address spatial patterns. We then regressed the shape (principal component scores) and size variables against ecology (mean annual temperature and diet) using multiple and multivariate spatial regression. Finally, the expected magnitudes of shape and size divergence under the influence of genetic drift and mutations alone were evaluated using neutral expectation for the divergence rate. Results The spatial correlograms showed a cline affecting the entire South American distribution. Interpolated maps showed that size and allometric shape vary from south‐east to north‐west. Multiple and multivariate regression analyses suggested that diet has the largest and most significant effect on this pattern of size and allometric shape variation. Finally, the results of the divergence rate test suggested that random processes alone cannot account for the morphological divergence exhibited by cranial size and allometric shape scores among southernmost populations. Main conclusions Correlograms, spatial regression and divergence rate analyses showed that although local factors (neutral processes or local environmental conditions) are important to explain spatial interpopulation differentiation in cranial characteristics among these populations, there is significant correlation of cranial size and allometric shape variation with diet. Gene flow among human populations, or local environmental conditions, could explain spatial variation mainly at smaller spatial scales, whereas the large‐scale pattern of the South American dataset is mainly related to the high proportion of carbohydrates and low proportion of proteins consumed.     

Morphology informed by phylogeny reveals unexpected patterns of species differentiation in the aquatic moss Rhynchostegium riparioides s.l

Bryophyte floras typically exhibit extremely low levels of endemism. The interpretation, that this might reflect taxonomic shortcomings, is tested here for the Macaronesian flora, using the moss species complex of Rhynchostegium riparioides as a model. The deep polyphyly of R. riparioides across its distribution range reveals active differentiation that better corresponds to geographic than morphological differences. Morphometric analyses are, in fact, blurred by a size gradient that accounts for 80% of the variation observed among gametophytic traits. The lack of endemic diversification observed in R. riparioides in Macaronesia weakens the idea that the low rates of endemism observed in the Macaronesian bryophyte flora might solely be explained by taxonomic shortcomings. To the reverse, the striking polyphyly of North American and European lineages of R. riparioides suggests that the similarity between the floras of these continents has been over-emphasized. Discriminant analyses point to the existence of morphological discontinuities among the lineages resolved by the molecular phylogeny. The global rate of error associated to species identification based on morphology (0.23) indicates, however, that intergradation of shape and size characters among species in the group challenges their identification.     

Body shape and size divergence among populations of Poecilia vivipara in coastal lagoons of south-eastern Brazil

Geometric shape analyses were used to study body shape and size variation among populations of the livebearing fish Poecilia vivipara inhabiting the recently formed coastal lagoons of Grussaí and Iquipari in Northern Rio de Janeiro State, Brazil. The largest components of morphological variation among females were between different habitats in the same lagoon, whereas for males there were larger differences between lagoons than between habitats. The shape differences were mostly localized in the head region and midbody, which indicated different patterns of locomotion and foraging behaviour optimized for the habitat experienced by each population. The pattern of size variation was similar to that of size-independent shape variation.     

Elliptical fourier analysis of symphyseal shape in great ape mandibles

The midsagittal profile of the mandibular symphysis has served as both a taxonomic marker and a phylogenetically salient character in debates over hominoid evolution. Nevertheless, the utility of symphyseal shape as an informative attribute for paleobiological reconstructions is suspect. Quantification of shape variation has proven to be particularly problematic; it has long been recognized that conventional linear measurements (and the indices derived from them), while replicable, summarize aspects of shape very poorly because of the vast amount of contour information that is lost in the process.In this study, a type of Fourier analysis is applied to cross-sectional contours of ape mandibles in order to provide a mathematical accounting of shape variation in a "global" sense; that is, by applying the "totality" of contour information in a comparative analysis. Shape variation in the mandibular symphysis is explored through the decomposition of coordinate data into elliptical Fourier coefficients. These coefficients are used to compute average taxonomic distances (ATD) among individuals of chimpanzees, gorillas, and orang-utans. The resulting shape-based distances are summarized via clustering (UPGMA) and ordination (principal coordinates analysis-PCO). Principal coordinate scores are subjected to analysis of variance in univariate and multivariate designs; these data are also applied to discriminant function analyses.Species and sex effects on morphology are statistically significant; however, no significant interaction of these factors is indicated. This would seem to imply that patterns of sexual dimorphism are not distinct among great apes; to the contrary, within-species sex comparisons reveal that significant size and shape dimorphism is present only in Gorilla. Despite significant size dimorphism in Pan and Pongo, significant shape differences between males and females are not apparent in these taxa.These results suggest that it is theoretically possible to sort taxa by a symphyseal shape criterion, but the discriminant function results suggest that there still exists a large potential for error in assigning particular shapes to a given species or sex. Thus, despite real shape differences among these species, the use of symphyseal shape as a character in species identification or in systematic arguments remains limited and problematic.     

Geometric craniometric analysis of sexual dimorphism and ontogenetic variation: A case study based on two geographically disparate species, Aethomys ineptus from southern Africa and Arvicanthis niloticus from Sudan (Rodentia: Muridae)

Non-geographic morphometric variation, particularly at the level of sexual dimorphism and ontogenetic (age-related) variation, has been documented in rodents, and useful for establishing whether to analyse sexes separately or together, and for selecting adult specimens for subsequent data recording and analysis. However, such studies have largely been based on traditional morphometric analyses of linear measurements that mainly focus on overall size, rather than shape-related morphometric variation. Unit-free, landmark/outline-based geometric morphometric analyses are considered to offer a more appropriate tool for assessing shape-related morphometric variation. In this study, we used geometric cranial morphometric analysis to assess the nature and extent of sexual dimorphism and age variation within the Tete veld rat, Aethomys ineptus (Thomas and Wroughton, 1908) from southern Africa and the African Nile rat, Arvicanthis niloticus (Desmarest, 1822) from Sudan. The results obtained were in turn compared with previously published results based on independent geometric and traditional cranial morphometric data from the same sampled populations examined in the present study. While our geometric morphometric results detected statistically significant sexual dimorphism in cranial shape within Ar. niloticus only, previously published results based on traditional morphometric data failed to detect significant sexual dimorphism within this species. However, similar to previously published traditional morphometric data, our geometric morphometric results detected statistically significant age-related variation in cranial shape and size within both Ae. ineptus and Ar. niloticus, with individuals of age classes 5 and 6 being considered to represent adult specimens. Our results highlight the importance of carefully evaluating both size- and shape-related non-geographic morphometric variation prior to the analysis of geographic variation and the delineation of species. Erroneous conclusions of non-geographic variation may have implications in the interpretation of geographic and evolutionary processes that may be responsible for morphological differences at both the inter- and intra-specific levels.     

Genetic, geographic, and environmental correlates of human temporal bone variation

Temporal bone shape has been shown to reflect molecular phylogenetic relationships among hominoids and offers significant morphological detail for distinguishing taxa. Although it is generally accepted that temporal bone shape, like other aspects of morphology, has an underlying genetic component, the relative influence of genetic and environmental factors is unclear. To determine the impact of genetic differentiation and environmental variation on temporal bone morphology, we used three-dimensional geometric morphometric techniques to evaluate temporal bone variation in 11 modern human populations. Population differences were investigated by discriminant function analysis, and the strength of the relationships between morphology, neutral molecular distance, geographic distribution, and environmental variables were assessed by matrix correlation comparisons. Significant differences were found in temporal bone shape among all populations, and classification rates using cross-validation were relatively high. Comparisons of morphological distances to molecular distances based on short tandem repeats (STRs) revealed a significant correlation between temporal bone shape and neutral molecular distance among Old World populations, but not when Native Americans were included. Further analyses suggested a similar pattern for morphological variation and geographic distribution. No significant correlations were found between temporal bone shape and environmental variables: temperature, annual rainfall, latitude, or altitude. Significant correlations were found between temporal bone size and both temperature and latitude, presumably reflecting Bergmann's rule. Thus, temporal bone morphology appears to partially follow an isolation by distance model of evolution among human populations, although levels of correlation show that a substantial component of variation is unexplained by factors considered here.     

Inter- and intraspecific variation in grass phytolith shape and size: a geometric morphometrics perspective

Background and AimsThe relative contributions of inter- and intraspecific variation to phytolith shape and size have only been investigated in a limited number of studies. However, a detailed understanding of phytolith variation patterns among populations or even within a single plant specimen is of key importance for the correct taxonomic identification of grass taxa in fossil samples and for the reconstruction of vegetation and environmental conditions in the past. In this study, we used geometric morphometric analysis for the quantification of different sources of phytolith shape and size variation.MethodsWe used landmark-based geometric morphometric methods for the analysis of phytolith shapes in two extant grass species (Brachypodium pinnatum and B. sylvaticum). For each species, 1200 phytoliths were analysed from 12 leaves originating from six plants growing in three populations. Phytolith shape and size data were subjected to multivariate Procrustes analysis of variance (ANOVA), multivariate regression, principal component analysis and linear discriminant analysis.Key ResultsInterspecific variation largely outweighed intraspecific variation with respect to phytolith shape. Individual phytolith shapes were classified with 83 % accuracy into their respective species. Conversely, variation in phytolith shapes within species but among populations, possibly related to environmental heterogeneity, was comparatively low.ConclusionsOur results imply that phytolith shape relatively closely corresponds to the taxonomic identity of closely related grass species. Moreover, our methodological approach, applied here in phytolith analysis for the first time, enabled the quantification and separation of variation that is not related to species discrimination. Our findings strengthen the role of grass phytoliths in the reconstruction of past vegetation dynamics.     

A geometric morphometric appraisal of beak shape in Darwin's finches

Beak size and shape in Darwin's finches have traditionally been quantified using a few univariate measurements (length, depth, width). Here we show the improved inferential resolution of geometric morphometric methods, as applied to three hierarchical levels: (i) among seven species on Santa Cruz Island, (ii) among different sites on Santa Cruz for a single species (Geospiza fortis), and (iii) between large and small beak size morphs of G. fortis at one site (El Garrapatero). Our results support previous studies in finding an axis of shape variation (long/shallow/pointy vs. short/deep/blunt) that separates many of the species. We also detect additional differences among species in the relative sizes and positions of the upper and lower mandibles and in curvature of the mandibles. Small-scale, but potentially relevant, shape variation was also detected among G. fortis from different sites and between sympatric beak size morphs. These results suggest that adaptation to different resources might contribute to diversification on a single island.     

Consequences of ignoring spatial variation in population trend when conducting a power analysis

Long‐term, large‐scale monitoring programs are becoming increasingly common to document status and trends of wild populations. A successful program for monitoring population trend hinges on the ability to detect the trend of interest. Power analyses are useful for quantifying the sample size needed for trend detection, given expected variation in the population. Four components of variation (within‐year variation at a given site, interannual variation within a site, variation among sites in the interannual variation, and variation among sites in mean abundance or density) are commonly considered in power analyses for population trend, but a fifth is rarely considered: variation among sites in the local trend. Spatial variation in trend is expected to reduce statistical power, but the magnitude of this reduction has not been fully explored. We used computer simulations to evaluate the consequences of ignoring spatial variation in trend under a variety of sampling designs and wide ranges of other components of variation. The effect of spatial variation in trend on power was minor when other input parameters took extreme values that made the trend either very difficult or very easy to detect. However, at moderate values of the other parameters, spatial variation in trend had a strong effect, reducing statistical power by up to 60%. In some cases, ignoring spatial variation in trend resulted in an 80% probability of a type I error (falsely detecting a trend in a stable population). Spatial variation in trend is therefore an important consideration when designing a long‐term monitoring program for many species, especially those affected by local conditions at sites that are repeatedly surveyed. If variation in trend is ignored, as in many previous power analyses, the recommended sampling design will likely be insufficient to detect the trend of interest and lead to potentially false conclusions of a stable population.     

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.     

Adaptation and constraint in the evolution of Drosophila melanogaster wing shape

SUMMARY We have taken advantage of parallel instances of natural selection on body size in Drosophila melanogaster to investigate constraints and adaptation affecting wing shape. Using recently developed techniques for statistical shape analysis, we have examined variation in wing shape in similar body size clines on three continents. Gender-related shape differences were constant among all populations, suggesting that gender differences represent a developmental constraint on wing shape. In contrast, the underlying shape varied significantly between continents and shape change within each cline (i.e., between small and large body size populations) also varied between continents. Therefore, variation at these two levels presumably results from either drift or natural selection. Functional considerations suggest that shape variation between the continents is unlikely to be adaptive. However, cline-related shape change, which we show has a significant allometric component, may be adaptive. The overall range of wing shape variation, across a large range of wing size, is extremely small, and the possibility that wing shape is subject to stabilizing selection (or canalization) is discussed.     

Variation in Body Shape across Species and Populations in a Radiation of Diaptomid Copepods

Inter and intra-population variation in morphological traits, such as body size and shape, provides important insights into the ecological importance of individual natural populations. The radiation of Diaptomid species (~400 species) has apparently produced little morphological differentiation other than those in secondary sexual characteristics, suggesting sexual, rather than ecological, selection has driven speciation. This evolutionary history suggests that species, and conspecific populations, would be ecologically redundant but recent work found contrasting ecosystem effects among both species and populations. This study provides the first quantification of shape variation among species, populations, and/or sexes (beyond taxonomic illustrations and body size measurements) to gain insight into the ecological differentiation of Diaptomids. Here we quantify the shape of five Diaptomid species (family Diaptomidae) from four populations each, using morphometric landmarks on the prosome, urosome, and antennae. We partition morphological variation among species, populations, and sexes, and test for phenotype-by-environment correlations to reveal possible functional consequences of shape variation. We found that intraspecific variation was 18-35% as large as interspecific variation across all measured traits. Interspecific variation in body size and relative antennae length, the two traits showing significant sexual dimorphism, were correlated with lake size and geographic location suggesting some niche differentiation between species. Observed relationships between intraspecific morphological variation and the environment suggest that divergent selection in contrasting lakes might contribute to shape differences among local populations, but confirming this requires further analyses. Our results show that although Diaptomid species differ in their reproductive traits, they also differ in other morphological traits that might indicate ecological differences among species and populations.     

Phenotypic plasticity in leaf morphology of Crataegus monogyna (Rosaceae): an experimental study with taxonomic implications

Many leaf characters are considered in the taxonomy of Crataegus in Europe, and several have been used in studies of the extent of hybridization in populations of northwest Europe. In such analyses it is assumed that the environmental component of phenotypic variation in such characters is insignificant. We tested this assumption by analysing the variation in the size and shape of leaves borne on clone cuttings of Crataegus monogyna maintained under identical conditions apart from the availability of soil nutrients. The resulting variation among leaves from this single genotype was as great as that observed previously across populations. Furthermore, although most of this variation could not be explained, a part could be attributed to differences in nutrient availability; of nine leaf characters investigated, eight showed significant variation due to this source, and in seven, the variation covaried significantly with nutrient level. The systematic implications of this are briefly explored.     

Ecological and evolutionary factors in dental morphological diversification among modern human populations from southern South America

The knowledge of processes involved in morphological variation requires the integrated analysis of evolutionary and ecological factors. Here, we investigate the factors responsible for dental variation among human populations from southern South America. The aim of this work is to test the correspondence of dental size and shape variation with geographical, molecular (i.e. mtDNA) and ecological (i.e. climate, diet and food preparation) variables employing comparative phylogenetic methods, which have not previously been extensively applied at a within-species level. The results of the Procrustes analysis show a significant association of shape variables with molecular distance and geography, whereas dental size is not associated with molecular or geographical distances among groups. Phylogenetic generalized least-squares analysis, which takes into account the evolutionary autocorrelation among populations, shows a significant relationship between dental size variation and diet, while temperature and pottery do not correspond with dental size or shape. Specifically, groups with diets rich in carbohydrates, as well as the maritime hunter-gatherers, have the smallest teeth. In summary, our results support ecological factors as the dominant factor on dental size diversification in this region, while evolutionary relationships account for variation in dental shape.