Environment and space as drivers of variation in skull shape in two widely distributed South‐American Tayassuidae,Pecari tajacu and Tayassu pecari (Mammalia: Cetartiodactyla)

The influence of the environment on the geographical variation of morphological traits has been recognized in a number of taxa. Pecari tajacu and Tayassu pecari are ideal models to investigate intraspecific geographic variation in skull because of their wide and heterogeneous geographical distribution in South America. We used geometric morphometric procedures to examine the geographical variation in skull shape of 294 adult specimens of these species from 134 localities. We quantified to what extent skull shape variation was explained by environment, skull size and geographical space using variation partitioning analysis. We detected a strong pattern of geographic variation for P. tajacu skull shape, but not for T. pecari. The environment seems to be the major selective force that drives skull shape variation in both species. Nevertheless, other spatially structured processes (e.g. genetic drift, gene flow) might also have affected variation in the skull shape of the more widespread species P. tajacu. Allometric relationships might reflect the biomechanical constraints that are thought to be strong enough to limit size‐related changes in T. pecari skull shape.     

Intraspecific variation in the skull morphology of the black caiman Melanosuchus niger (Alligatoridae,Caimaninae)

Melanosuchus niger is a caimanine alligatorid widely distributed in the northern region of South America. This species has been the focus of several ecological, genetic and morphological studies. However, morphological studies have generally been limited to examination of interspecific variation among extant species of South American crocodylians. Here, we present the first study of intraspecific variation in the skull of M. niger using a two‐dimensional geometric morphometric approach. The crania of 52 sexed individuals varying in size were analysed to quantify shape variation and to assign observed shape changes to different types of intraspecific variation, that is, ontogenetic variation and sexual dimorphism. Most of the variation in this species is ontogenetic variation in snout length, skull depth, orbit size and the width of the postorbital region. These changes are correlated with bite force performance and probably dietary changes. However, a comparison with previous functional studies reveals that functional adaptations during ontogeny seem to be primarily restricted to the postrostral region, whereas rostral shape changes are more related to dietary shifts. Furthermore, the skulls of M. niger exhibit a sexual dimorphism, which is primarily size‐related. The presence of non‐size‐related sexual dimorphism has to be tested in future examinations.     

Geographic pattern of cranial differentiation in the Asian Midday Jird Meriones meridianus (Rodentia: Muridae: Gerbillinae) and its taxonomic implications

The existence of cryptic species in the midday jird (Meriones meridianus) has been suggested in literature, although based on little empirical data to support this hypothesis. In this study, a two‐dimensional landmark‐based geometric morphometric approach was used to investigate whether patterns in intraspecific variation in skull shape and size exist, using 110 skull specimens from more than 20 different localities along the distribution range of M. meridianus. This is the first study of morphological differences on such a big sample size and geographical range, and it tries to find whether skull shape variation in this species is best described as being clinal or rather reflecting cryptic diversity. The latter seems to be the case, as a dimorphic skull phenotype was found, reflecting a geographic disparity between the Middle East and the Far East specimens both in shape and in size. Distinct cranial differences were found in the overall cranial size and, also at the level of the inflation of the bulla, the elongation of the nasal, the length of the teeth row and the incisive foramen, as well as the distance in between the latter two. It thus seems that M. meridianus from Middle East is morphologically distinct from that of the Far East. Furthermore, our results also demonstrate that clinal variation could explain variation within Middle East populations, whereas a more heterogenous pattern is found for those of the Far East. The hypothesis that the observed phenotypic variation may reflect cryptic species is discussed, with the recommendation for a thorough taxonomical revision of the genus in the region.     

Are geometric morphometric analyses replicable? Evaluating landmark measurement error and its impact on extant and fossil Microtus classification

Geometric morphometric analyses are frequently employed to quantify biological shape and shape variation. Despite the popularity of this technique, quantification of measurement error in geometric morphometric datasets and its impact on statistical results is seldom assessed in the literature. Here, we evaluate error on 2D landmark coordinate configurations of the lower first molar of five North American Microtus (vole) species. We acquired data from the same specimens several times to quantify error from four data acquisition sources: specimen presentation, imaging devices, interobserver variation, and intraobserver variation. We then evaluated the impact of those errors on linear discriminant analysis‐based classifications of the five species using recent specimens of known species affinity and fossil specimens of unknown species affinity. Results indicate that data acquisition error can be substantial, sometimes explaining >30% of the total variation among datasets. Comparisons of datasets digitized by different individuals exhibit the greatest discrepancies in landmark precision, and comparison of datasets photographed from different presentation angles yields the greatest discrepancies in species classification results. All error sources impact statistical classification to some extent. For example, no two landmark dataset replicates exhibit the same predicted group memberships of recent or fossil specimens. Our findings emphasize the need to mitigate error as much as possible during geometric morphometric data collection. Though the impact of measurement error on statistical fidelity is likely analysis‐specific, we recommend that all geometric morphometric studies standardize specimen imaging equipment, specimen presentations (if analyses are 2D), and landmark digitizers to reduce error and subsequent analytical misinterpretations.     

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.     

The geographic structure of morphological variation in eight species of fiddler crabs (Ocypodidae: genus Uca) from the eastern United States and Mexico

Species with larger geographic distributions are more likely to encounter a greater variety of environmental conditions and barriers to gene flow than geographically‐restricted species. Thus, even closely‐related species with similar life‐history strategies might vary in degree and geographic structure of variation if they differ in geographic range size. In the present study, we investigated this using samples collected across the geographic ranges of eight species of fiddler crabs (Crustacea: Uca) from the Atlantic and Gulf coasts of North America. Morphological variation in the carapace was assessed using geometric morphometric analysis of 945 specimens. Although the eight Uca species exhibit different degrees of intraspecific variation, widespread species do not necessarily exhibit more intraspecific or geographic variation in carapace morphology. Instead, species with more intraspecific variation show stronger morphological divergence among populations. This morphological divergence is partly a result of allometric growth coupled with differences in maximum body size among populations. On average, 10% of total within‐species variation is attributable to allometry. Possible drivers of the remaining morphological differences among populations include gene flow mediated by ocean currents and plastic responses to various environmental stimuli, with isolation‐by‐distance playing a less important role. The results obtained indicate that morphological divergence among populations can occur over shorter distances than expected based on dispersal potential. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 248–270.     

Morphometric study of two species of wood miceApodemus sylvaticus andA. flavicollis (Rodentia: Muridae): traditional and geometric morphometric approach

Skull shape and size were investigated in 311 specimens ofApodemus sylvaticus (Linnaeus, 1758) andA. flavicollis (Melchior, 1834), using traditional and geometric morphometric approach. We searched for potential morphometric diagnostic criteria between these morphologically similar mice and we an alyzed the interpopulation morphometric variation. Specific status of all individuals was identified by scoring four diagnostic allozyme markers. Discriminant analysis based on 16 cranial measure ments as well as that based on land mark data for ventral side of the skull discriminated in dividuals with 99.7% accuracy. The classification functions useful for practical discrimination of unknown specimens are given. Our data showed neither clear intraspecific geographical pattern among studied populations nor evidence for character displacement in syntopic populations ofA. sylvaticus andA. flavicollis.     

Assessment of rampant genitalic variation in the spider genus Homalonychus (Araneae, Homalonychidae)

Abstract. Animal genitalia are often complex and thought to vary little within species but differ between closely related species making them useful as primary characters in species diagnosis. Spiders are no exception, with nearly all of the 40,462 (at the time of this writing) described species differentiated by genitalic characteristics. However, in some cases, the genitalia of putative species are not uniform, but rather vary within species. When intraspecific variation overlaps interspecific variation, it can be difficult (if not impossible) to place a name on a specimen. The quantification of shape variation in genitalia has not often been attempted, probably because until recently it was not a methodologically and computationally simple process. In the two currently recognized species of the spider genus Homalonychus , genitalic variation is rampant in both male and female structures, with some parts of the genitalia (e.g., the retrolateral tibial apophysis) differing in each specimen examined. In this study, geometric morphometric analysis employing landmark data is used to quantify both intra- and interspecific variation in this genus. The large amount of variation is condensed into two or three groups depending on the structures examined, and these groups correspond to either the two species or to previously established mitochondrial DNA clades within one of the species. The results also show that analyses of female structures do not separate the groups as readily as the analyses of the male structures. The large amount of variation present in some structures is not correlated with geography or population genetic structure.     

An input of heterochronies to the formation of interspecific differences in wood mice, Sylvaemus (Rodentia)

Difference in adult size and body shape among the closely related species is the result of diversification in morphogenetic processes. We examined age variation in various external and craniometric characters of four Sylvaemus species from Ukrane fauna, S. uralensis, S. arianus, S. sylvaticus, and S. tauricus, using cross-sectional techniques. The intensive growth period occurred to be different in various species. Rapid growth of S. tauricus continues during all life span, while in other species growth rate becomes slower at different stages. This peculiarity of S. tauricus may be the result of prolonged and more intensive production of somatotropin by adenohypophysis. Side effects of adenohypophysis activation are resulted from changes in S. tauricus skull shape and more intensive pigmentation in comparison with three other species. Prolongation of growth could be developmental mechanism for size increasing in evolution of S. tauricus. The independence of body size and pigmentation is principally the same at interspecific and intraspecific level.     

A phylogenetic view on skull size and shape variation in the smooth newt (Lissotriton vulgaris,Caudata, Salamandridae)

In this study, we explore skull size and shape variation in the smooth newt, a taxon with substantial morphological differentiation and complex phylogeographic relations. By projecting phylogenies into the morphospace of the skull shape, we explore the variation in and differentiation of this complex morphological structure within a phylogenetic framework. For these analyses, we used a dataset that covers the most southern part of the species’ distribution range, including all conventionally recognized subspecies. The study revealed different patterns of divergence in skull shape between sexes, which is paralleled by intraspecific differentiation. The divergence in dorsal skull shape is concordant with the phylogenetic divergence, as the most diverged clades of the smooth newt (Lissotriton vulgaris kosswigi and Lissotriton vulgaris lantzi) exhibit a skull shape that significantly diverges from the smooth newt’s mean shape configuration. The results of this study also indicate that ventral skull portion, which is more directly related to feeding and foraging, shows higher variation between populations than dorsal skull portion, which appears to be less variable and phylogenetically informative.     

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.     

A morphometric and genetic framework for the genus Gazella de Blainville, 1816 (Ruminantia: Bovidae) with special focus on Arabian and Levantine mountain gazelles

Gazella is one of the most species‐rich genera within horned ruminants. Despite overall similarity in body size and morphology, gazelles show variability in coloration and horn morphology. Unfortunately, however, species differentiation based on these characters, or on discrete skull characters, is very difficult due to high intraspecific variability. Furthermore, most species have fragmented and allopatric distributions, so that species boundaries were hard to define in the past. Mitochondrial DNA sequences have proven useful for investigating gazelle taxonomy in recent years, but especially for old museum material, i.e. type specimens, destructive sampling is often impossible. We provide a comprehensive morphometric framework for the genus Gazella based on linear skull measurements reconciled with results from molecular phylogenetic analysis based on the largest dataset available so far. In particular for males, the skull morphology shows interspecific differences concurrent with DNA data and provides a reliable tool for species identification. Based on morphometric data we synonymize G. karamii with G. marica, and confirm the identification of the G. arabica and G. a. rueppelli type skulls from analyses of mitochondrial DNA sequences. © 2013 The Linnean Society of London     

Microevolutionary patterns in the common caiman predict macroevolutionary trends across extant crocodilians

Both extinct and extant crocodilians have repeatedly diversified in skull shape along a continuum, from narrow‐snouted to broad‐snouted phenotypes. These patterns occur with striking regularity, although it is currently unknown whether these trends also apply to microevolutionary divergence during population differentiation or the early stages of speciation. Assessing patterns of intraspecific variation within a single taxon can potentially provide insight into the processes of macroevolutionary differentiation. For example, high levels of intraspecific variation along a narrow‐broad axis would be consistent with the view that cranial shapes can show predictable patterns of differentiation on relatively short timescales, and potentially scale up to explain broader macroevolutionary patterns. In the present study, we use geometric morphometric methods to characterize intraspecific cranial shape variation among groups within a single, widely distributed clade, Caiman crocodilus. We show that C. crocodilus skulls vary along a narrow/broad‐snouted continuum, with different subspecies strongly clustered at distinct ends of the continuum. We quantitatively compare these microevolutionary trends with patterns of diversity at macroevolutionary scales (among all extant crocodilians). We find that morphological differences among the subspecies of C. crocodilus parallel the patterns of morphological differentiation across extant crocodilians, with the primary axes of morphological diversity being highly correlated across the two scales. We find intraspecific cranial shape variation within C. crocodilus to span variation characterized by more than half of living species. We show the main axis of intraspecific phenotypic variation to align with the principal direction of macroevolutionary diversification in crocodilian cranial shape, suggesting that mechanisms of microevolutionary divergence within species may also explain broader patterns of diversification at higher taxonomic levels.     

Divergence in size,but not in shape: variation in skull size and shape within Ommatotriton newts

Using a geometric morphometric approach, we explored the variation in skull size and skull shape in banded newts (genus Ommatotriton). The genus Ommatotriton is represented by two allopatric, genetically well‐defined species: Ommatotriton ophryticus and O. vittatus. Within each species, two subspecies have been recognised. The samples used in this study cover the geographical and genetic variation within each species. We found statistically significant variation in skull size between species and among populations within species. When corrected for size, there was no significant variation in shape between species. Our results indicate that the variation in skull shape within the genus Ommatotriton is almost entirely due to size‐dependent, allometric shape changes. The exception is the shape of the ventral skull in males. Males of O. ophryticus and O. vittatus significantly diverge in the shape of the ventral cranium. The ventral skull, more precisely the upper jaw and palate, is directly functionally related to feeding. In general, our results indicate that allometry is a significant factor in the morphological variation of banded newts. However, the divergence in the ventral skull shape of males indicates that sexual selection and niche partitioning may have influenced the evolution of skull shape in these newts.     

Geographic variation in the greater Japanese shrew-mole,Urotrichus talpoides: Combining morphological and chromosomal patterns

Although much information on the species dynamics of small mammals may be gleaned from the integration of morphological and molecular data sets, the two are not routinely combined when species boundaries and definitions are investigated. The greater Japanese shrew-mole (Urotrichus talpoides) presents a rare example of intraspecific and geographical euchromatic chromosomal variation. In this study a combination of 2D landmark-based and outline-based geometric morphometric methods were used to provide the first quantitative examination of variation in skull morphology occurring between populations of U. talpoides. Geographic variation was found to be most conspicuous in ventral and dorsal cranial morphology, and less evident for outline-based analyses of the dentary, thereby indicating differing magnitudes of skeletal plasticity associated with geographic variation in each system. Both ventral and dorsal cranial morphology differed significantly between shrew-moles from western and eastern Honshu, in agreement with previously identified chromosomal variation boundaries. Inclusion of other island populations revealed shape differences between shrew-moles in Kyushu, Tsushima and North Honshu. These results lend general support for the unification of morphological and chromosomal data when assessing species boundaries.     

Multivariate discrimination between East African cape hares (Lepus capensis) and savanna hares (L. victoriae) based on occipital bone shape

In East Africa, cape hares (Lepus capensis) and savanna hares (L. victoriae) look much alike where their ranges overlap. Earlier studies suggested discrimination between the two species by several skull traits, but did not present morphometric statisitics. Our present discriminant analysis based on seven metric variables of the occipital bone related to skull length (condylobasal length) resulted in a high (95.1%) overall probability of correct separation of the two species. While all cape hares were classified correctly, correct classification was a bit lower in savanna hares (88.2%). A principal components analysis of the same variables confirmed the shape difference for the two species. Both region and sex-specific variation in the shape of the studied occipital bone complex was found in savanna hares but not in cape hares. The somewhat reduced level of correct identification of savanna hares might be due to a tendency for higher shape variation. Application of our discriminant analysis to other regions in East Africa is discussed.     

Intraspecific shape variation in horseshoe crabs: The importance of sexual and natural selection for local adaptation

A morphometric analysis of the body shape of three species of horseshoe crabs was undertaken in order to infer the importance of natural and sexual selection. It was expected that natural selection would be most intense, leading to highest regional differentiation, in the American species Limulus polyphemus, which has the largest climatic differences between different populations. Local adaptation driven by sexual selection was expected in males but not females because horseshoe crab mating behaviour leads to competition between males, but not between females. Three hundred fifty-nine horseshoe crabs from nine populations, representing three species, were analyzed using a digitizer to position sixty morphometric landmarks in a three-dimensional space. Discriminant analysis revealed strong regional structuring within a species, which suggests strong philopatry, and showed the existence of geographically-based intraspecific variation. An admixture analysis showed regional intraspecific differentiation for males and females of L. polyphemus and males of the Asian horseshoe crab Carcinoscorpius rotundicauda, but not for females of C. rotundicauda and another Asian horseshoe crab, Tachypleus gigas. Differences in shape variation between sexes were tested with F-tests, which showed lower intrapopulation morphometric variation in males than females. These results indicate a lower degree of local adaptation on body shape in C. rotundicauda and T. gigas than in L. polyphemus and a lower degree of local adaptation in females than in males.