A morphometric analysis of extant and early miocene fossil hominoid maxillo-dental specimens

It is demonstrated in this paper that before we can hope to formulate phylogenetic relationships between and amongst fossil hominoid material it is first necessary to sex the material accurately. In order to determine whether the morphological and morphometrical variability seen in fossil specimens is due to sexual or inter species dimorphism, it is necessary to calibrate fossil specimens against extant hominoid species' morphologies. Only after fossil specimens have been sexed is it possible to differentiate between morphologies that are related to sex and those that are species specific. This will help reduce fossil misallocation. A morphometric analysis of extant and fossilProconsul hominoid material is presented. Each fossil specimen has been sexed according to symplesiomorphic sex morphologies as defined in this paper. After the fossil specimens have been sexed they are analyzed using multivariate statistics. The identification of differing sex patterns within the specimens examined here suggests that a new species ofProconsul may have to be considered.     

Finite element modeling of shell shape in the freshwater turtle Pseudemys concinna reveals a trade‐off between mechanical strength and hydrodynamic efficiency

Aquatic species can experience different selective pressures on morphology in different flow regimes. Species inhabiting lotic regimes often adapt to these conditions by evolving low‐drag (i.e., streamlined) morphologies that reduce the likelihood of dislodgment or displacement. However, hydrodynamic factors are not the only selective pressures influencing organismal morphology and shapes well suited to flow conditions may compromise performance in other roles. We investigated the possibility of morphological trade‐offs in the turtle Pseudemys concinna. Individuals living in lotic environments have flatter, more streamlined shells than those living in lentic environments; however, this flatter shape may also make the shells less capable of resisting predator‐induced loads. We tested the idea that “lotic” shell shapes are weaker than “lentic” shell shapes, concomitantly examining effects of sex. Geometric morphometric data were used to transform an existing finite element shell model into a series of models corresponding to the shapes of individual turtles. Models were assigned identical material properties and loaded under identical conditions, and the stresses produced by a series of eight loads were extracted to describe the strength of the shells. “Lotic” shell shapes produced significantly higher stresses than “lentic” shell shapes, indicating that the former is weaker than the latter. Females had significantly stronger shell shapes than males, although these differences were less consistent than differences between flow regimes. We conclude that, despite the potential for many‐to‐one mapping of shell shape onto strength, P. concinna experiences a trade‐off in shell shape between hydrodynamic and mechanical performance. This trade‐off may be evident in many other turtle species or any other aquatic species that also depend on a shell for defense. However, evolution of body size may provide an avenue of escape from this trade‐off in some cases, as changes in size can drastically affect mechanical performance while having little effect on hydrodynamic performance. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

The comparative morphology of the larynx in snakes

Young, B.A. 2000. The comparative morphology of the larynx in snakes. —Acta Zoologica (Stockholm) 81 : 177–193 The larynx and glottal tube were examined in 10 specimens each of 22 snake species. Qualitative analysis through dissection and clearing and staining revealed distinct morphological variation in the cartilage and intrinsic musculature of the larynx. Quantitative analysis of nine morphometric characters revealed laryngeal sexual dimorphism in three species, significant interspecific differences in the coefficients for every morphometric feature regressed on body size, no significant intraspecific differences in the log transformed means, and poor discrimination of species using principal component analysis. The observed variations in laryngeal morphology did not correlate with the phylogeny of the species, or with habitat preferences.     

Geometric morphometric analysis reveals that the shells of male and female siphon whelks Penion chathamensis are the same size and shape

Secondary sexual dimorphism can make the discrimination of intra and interspecific variation difficult, causing the identification of evolutionary lineages and classification of species to be challenging, particularly in palaeontology. Yet sexual dimorphism is an understudied research topic in dioecious marine snails. We use landmark-based geometric morphometric analysis to investigate whether there is sexual dimorphism in the shell morphology of the siphon whelk Penion chathamensis. In contrast to studies of other snails, results strongly indicate that there is no difference in the shape or size of shells between the sexes. A comparison of P. chathamensis and a related species demonstrates that this result is unlikely to reflect a limitation of the method. The possibility that sexual dimorphism is not exhibited by at least some species of Penion is advantageous from a palaeontological perspective as there is a rich fossil record for the genus across the Southern Hemisphere.     

Thecamoebians from Late Permian Gondwana sediments of peninsular India

The evolutionary history of thecamoebians (testate amoebae) extends back to the Neoproterozoic Era. However, until now, these have had a restricted, discontinuous and modest record across the world. The studied sediment of Raniganj Formation (Godavari Graben), Andhra Pradesh, India has been assigned as Late Permian on the basis of co-occurring age-diagnostic Late Permian palynomorphs. About sixteen thecamoebian species and one taxon incertae sedis have been recorded here in the palynological slides on the basis of shell morphology and morphometry. Out of these, five belong to the family Arcellidae, seven to Centropyxidae, two to Trigonopyxidae, one to Difflugiidae, one to Plagiopyxidae, and one is regarded incertae sedis. The morphometric characteristics of fossil forms resemble their corresponding extant species studied from ecologically diverse fresh water wetlands in India. In general, the ratio of shell diameter and aperture diameter of Late Permian fossil and extant specimens show significant correlation in all the studied species. Except that, the ratio of shell length and breadth is the distinguishing feature between Centropyxis aerophila and C. aerophila ‘sylvatica’, rather than the ratio of shell length and longest diameter of the shell aperture in both fossil and extant forms. The study elucidates the minimal morphological evolution in thecamoebians and their survival during mass extinction periods and stressful environmental conditions over the geological timescale.     

Sex Differences in Discrimination of Shoal Size in the Guppy (Poecilia reticulata)

For a diversity of species, differences in sexual and parental roles, along with differences in body morphology, often result in males and females having different diets, distinct predators and even different patterns of habitat use. As a consequence, the two sexes often face different environmental challenges and selection may favour the evolution of sex differences in cognition. We tested this prediction in the guppy (Poecilia reticulata). Under perceived hazard, individual guppies join the larger available social group, a behaviour that is thought to minimise predation risk. In this species, females are more frequently exposed to predation and more averse to predation risk; we therefore expected greater accuracy in shoal size discrimination in females. We compared the accuracy of male and female guppies in discriminating shoals of 4 and 6 conspecifics, which represents the upper limit of discrimination for this species. Overall, we found no sex differences in the accuracy of discriminating the two shoals. However, while females showed this ability at the beginning of the test, males began to select the larger group only after several minutes. In three control experiments, we found indications that this sex difference cannot be accounted for by differences in motivation or antipredator strategies between the two sexes, suggesting female guppies are more efficient at rapidly estimating shoal size.     

Identification of ecophenotypic trends within three European freshwater mussel species (Bivalvia: Unionoida) using traditional and modern morphometric techniques

Most species of freshwater mussels (Unionoida) show a wide variability in shell form and size but an understanding of which factors determine unionoid morphology is poor. We identified ecophenotypic trends in shell and internal characters within three unionoid species from two habitat types (marinas and river) of the River Thames, UK, using traditional and modern morphometric techniques. In marinas, all species grew to larger maximum sizes than in the river, which might be a result of higher temperatures and phytoplankton densities in marinas. Unio pictorum in marinas was more elongated than in the river and Fourier shape analysis revealed a trend from dorsally arched river specimens to straight dorsal and pointed posterior margins in marina individuals. The degree of shell elongation and shape of dorso‐posterior margin were not associated with sediment composition, but were associated with the different hydrological characters of the two habitat types. Relative shell width was a poor indicator of collection site and influenced by allometric growth. Unlike U. pictorum, a difference in shell elongation of marina and river mussels could not be detected in Unio tumidus and Anodonta anatina. However, all three species showed the same trends regarding the shape of the dorso‐posterior shell margin. This shell character may thus have broad ecological significance and could have considerable utility to palaeontologists, taxonomists, and conservation biologists. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 814–825.     

Evolution of opercle bone shape along a macrohabitat gradient: species identification using mtDNA and geometric morphometric analyses in neotropical sea catfishes (Ariidae)

Transitions between the marine and freshwater macrohabitat have occurred repeatedly in the evolution of teleost fishes. For example, ariid catfishes have moved from freshwater to marine environments, and vice versa. Opercles, a skeletal feature that has been shown to change during such transitions, were subjected to 2D geometric morphometric analyses in order to investigate evolutionary shape changes during habitat transition in ariid catfishes and to test the influence of habitat on shape changes. A mtDNA marker, which proved useful in previous studies, was used to verify species identities. It greatly improved the assignment of specimens to a species, which are difficult to assign by morphology alone. The application of a mtDNA marker confirmed the occurrence of Notarius biffi in Central America, South of El Salvador. Molecular identification together with principal component analysis (PCA) and further morphological inspection of neurocrania indicated the existence of a cryptic species within Bagre pinnimaculatus. Principal component (PC) scores of individual specimens clustered in morphospace by genus rather than by habitat. Strong phylogenetic structure was detected using a permutation test of PC scores of species means on a phylogenetic tree. Calculation of Pagel's λ suggested that opercle shape evolved according to a Brownian model of evolution. Yet canonical variate analysis (CVA) conducted on the habitat groups showed significant differences in opercle shapes among freshwater and marine species. Overall, opercle shape in tropical American Ariidae appears to be phylogenetically constrained. This verifies the application of opercle shape as a taxonomic tool for species identification in fossil ariid catfishes. At the same time, adaptation to freshwater habitats shows characteristic opercle shape trajectories in ariid catfishes, which might be used to detect habitat preferences in fossils.     

INVESTIGATIONS OF ANGIOSPERMS FROM THE EOCENE OF NORTH AMERICA: STIPULATE LEAVES OF THE RUBIACEAE INCLUDING A PROBABLE POLYPLOID POPULATION

A comparative study was made of the gross morphology, fine venation and cuticular features of Leitneria fioridana Chapman, the single living representative of the order Leitneriales and Leitneria eocenica (Berry) Brown, presumbaly a related fossil species. In addition to the type material, newly collected fossil specimens were investigated from clay pits in the Middle Eocene, Claiborne Formation, of western Tennessee and Kentucky. Foliate stipules attached to the petioles of several specimens suggest the assignment of this fossil leaf type to the genus Leitneria is incorrect. The nature of the gross morphology, fine venation and cuticular features confirms the misidentification. Previously, various specimens of this fossil leaf type have been placed in eight species of seven genera in seven families of six angiosperm orders, none of which are correct systematically. The gross morphology, venation and cuticular characters of the fossil leaf are distributed among a few extant South American genera of arborescent Rubiaceae. The fossil is an extinct rubiaceous leaf type which cannot be placed within a single modern subfamily, tribe or genus of the family. The organ genus, Paleorubiaceophyllum is proposed for these leaves. Three varieties of a single fossil species, P. eocenicum, are recognized. One variety with epidermal cells nearly twice the size of the others may represent a polyploid population.     

Sexual dimorphism in canine shape among extant great apes

There have been numerous attempts to sex fossil specimens using the canine dentition. Whether focused on canine size or canine shape, most of these efforts share two deficiencies: lack of quantification of male-female differences in the adopted criteria and a failure to adequately explore among extant species the discriminatory power of these criteria. Here, canine shape indices relating to relative canine height, upper canine root/crown proportionality, and relative length of the lower canine mesial ridge were calculated for males and females of all species and subspecies of extant great apes and two species of gibbons. The accuracy of these indices for identifying the sex of the extant ape specimens was investigated through discriminant analysis and the use of bivariate plots of the two upper and two lower canine indices. The indices were found to be highly accurate in identifying the sex of great ape individuals, not only in single-species and subspecies samples but in mixed-species samples as well; assignment error rates were mostly between 0 and 4%. Accuracy was lowest in Pan (error rates as high as 15%) and highest in Pongo (one error). In most cases, error rates were lower in the upper canines. The effectiveness of these shape indices for sexing might be related to the degree of absolute canine size dimorphism; the indices did not effectively segregate males and females among minimally canine-dimorphic gibbons. The mixed-species results reveal that same-sex index values are remarkably concordant across great ape species, as are the patterns of spatial segregation of males and females in the bivariate plots. Results suggest that, while the indices can be used with some confidence to sex individual fossil specimens, their greatest utility will be for identifying the sex of groups of canines united by size and morphology. © 1995 Wiley-Liss, Inc.     

Computer-assisted paleoanthropology

Paleoanthropologists are confronted by a steadily growing number of fossil specimens exhibiting diversity in both apparent morphology and state of preservation. Studying this material to answer phylogenetic and functional questions requires extensive qualitative assessment accompanied by quantitative evaluation of large volumes of data. Over the past decade, major new developments in both respects have been made possible through the advent of medical imaging technologies, most notably computer tomography (CT), and through concomitant progress in computer graphics technology. In paleoanthropology, these techniques offer noninvasive tools for visualization of inaccessible regions of the skeleton, computer-assisted reconstruction of fragmentary fossil specimens, and morphometric or biomechanical analysis of data derived from CT images. A series of CT based studies has already yielded new insights into character differences between fossil hominid species. © 1998 Wiley-Liss, Inc.     

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.     

Sexual,habitat‐constrained and parasite‐induced dimorphism in the shell of a freshwater mussel (Anodonta anatina,Unionidae)

Intraspecific trends in freshwater mussel (unionoid) shells that are consistently associated with differences in the mussels' sex and/or parasitic infestation can potentially be used to reconstruct sex ratios or parasitic levels of modern and ancient unionoid populations. In contrast to morphological patterns within mammal species, such dimorphic trends within unionoid species are, however, poorly understood. This study investigates, for the first time, to what extent sex, trematode infection and indirect habitat effects determine shell morphology in the freshwater mussel Anodonta anatina. Three of the five study populations displayed significant sexual shell width dimorphism. Here, shells of females were significantly wider than males, probably as a result of altered shell growth to accommodate marsupial gills. In two of these populations, female shells were additionally significantly thinner than those of males, which could be a result of resource depletion by offspring production. Two other A. anatina populations showed no significant dimorphic patterns, and our results indicate that this interpopulational difference in the degree of sexual dimorphism may reflect the overarching effect of habitat on morphology. Thus, populations in the most favourable habitats exhibit faster growth rates, attain larger maximum sizes and produce more offspring, which results in more swollen gills and consequently more inflated shells of gravid females compared to less fecund populations. None of the populations showed any evidence for sexual dimorphism in overall size, growth rate, sagittal shape and density of shells. In addition to sexual dimorphisms, infestation by bucephalid trematode parasites (Rhipidocotyle sp.) significantly altered sagittal and lateral shell shape of A. anatina in one of the populations, with infected specimens growing wider and more elongated. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

MODERN AND HISTORICAL EVIDENCE FOR NATURAL HYBRIDIZATION BETWEEN SYMPATRIC SPECIES IN MANDARINA (PULMONATA: CAMAENIDAE)

Geographical and temporal variation in gene exchange between two endemic land snail species, Mandarina aureola and Mandarina ponderosa, was studied on Hahajima Island of the Bonin Islands. Allozyme variation in modern samples, and variation in the color and shell morphology of modern and fossil samples, suggest a complex geographical and historical pattern of hybridization. These two species occur in sympatry, and their shell morphologies and protein genotypes are markedly divergent. However, many specimens of M. aureola, collected from the middle region of the island, exhibit intermediate shell morphologies and possess marker alleles of M. ponderosa. Fossil samples of the two species strongly suggest that these intermediates were hybrids with M. ponderosa that were produced since the end of the Pleistocene. Each of these species, in addition, is subdivided into two genetically and morphologically divergent parapatric races. Interspecific hybridization appears to have produced genetical and morphological admixture among these four distinctive groups of populations. The past distribution and geographic variation of M. ponderosa can be traced in the distribution of M. ponderosa-derived genotypes in current populations of M. aureola. Temporal changes of the color pattern in the fossil populations of hybrids suggest that the traits introduced from M. ponderosa to M. aureola have been affected by natural selection and could replace traits of living species when advantageous. Moreover, these introgressed genes appeared to provide novel properties that enabled M. aureola to advance into a new environment. Relatively independent change in shell color and morphology further suggests mosaic evolution following the hybridization events. Connectively, these data reveal how hybridization events may be an important source of evolutionary novelties and make it clear that the phenomenon of reticulate evolution cannot be ignored.     

Monster or multiplacophoran: A teratological specimen of the chiton Acanthopleura granulata (Mollusca: Polyplacophora) with a valve split into independent and symmetrical halves

Teratological specimens deviate from the conserved form of their species. In doing so, they serve as natural experiments that refine our knowledge of developmental mechanisms and the natural limits of phenotypes. Here, we describe a specimen of the West Indian Fuzzy Chiton Acanthopleura granulata (Gmelin, 1791) with a fifth valve split into two halves. Using micro-CT to non-invasively visualize the external and internal morphology of this specimen, we find that the half valves are symmetrical and independent from each other and from any of the other valves. The presence of girdle-like tissue between the split valves suggests that this shell abnormality arose in early development and was not the product of damage to the adult animal. While the present specimen of A. granulata is clearly abnormal for its species, its split valve may provide some insights into the developmental pathways that would underlie macroevolutionary transitions to multi-plated chiton forms known from the fossil record.     

Geometric morphometric analysis of shell shape variation in Conus (Gastropoda: Conidae)

Geometric morphometric techniques allow for the direct quantification and analysis of variation in biological shape and have been used in studies in systematic biology. However, these techniques have not been used for species discrimination in the gastropod genus Conus, a major taxon of significant tropical reef predators recognized for their peptide‐based toxins. Here, we used landmark digitization and analysis to show that five species commonly studied for their conotoxins –Conus consors, Conus miles, Conus stercusmuscarum, Conus striatus, and Conus textile – can be effectively distinguished from each other by their shape, as manifested in the results of a principal components analysis (PCA) and the generated thin‐plate splines. Two piscivorous species, C. stercusmuscarum and C. striatus, show clear overlaps in the PCA plot, although each taxon clusters within itself, as does each of the others. The loadings on the first two principal components show that the forms of the shells' aperture and spire are particularly important for discrimination. Phylogenetic analysis using neighbour‐joining methods shows that group separations are comparable with published phylogenetic schemes based on molecular data and feeding mode (i.e. piscivory, vermivory, molluscivory). The results of this study establish the utility of geometric morphometric methods in capturing the interspecific differences in shell form in the genus Conus. This may lead to the utilization of these methods on other gastropod taxa and the creation of species‐recognition programs based on shell shape. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 296–310.     

Use of shape to distinguish Chamelea gallina and Chamelea striatula (Bivalvia: Veneridae): Linear and geometric morphometric methods

The commercially fished striped venus clams Chamelea gallina and C. striatula (Bivalvia: Veneridae) are difficult to distinguish by inexperienced observers and the taxonomy of these species is still an issue of discussion. The differences in shape between C. gallina and C. striatula from Portuguese coastal waters were studied through conventional linear and geometric morphometric analysis, using both contour (elliptic Fourier analysis) and landmark-based methods. The relationships shell length vs. height, width, and total weight were significantly different between species. However, because there was a considerable overlap in the data sets, the species could not be distinguished using any combination of those linear measurements. Geometric morphometric methods provided shape variables that led to 0-6% misclassification rates between species; linear morphometric measures led to 16.8% error. Contour analysis revealed differences primarily in the shell umbo and lunular area. The umbo was more "sharp" and the lunula less pronounced in C. striatula than in C. gallina. Generalized procrustes superimposition (landmark analysis) showed that the main differences between species reside in the length of the pallial sinus. Thus, an index was developed (PI: Pallial Index = pallial sinus length/shell length), which was successfully used to separate the species (with 100% correct classification), i.e., specimens with PI lower than 0.119 belonged to C. gallina, whereas greater PI values were attributed to C. striatula. The use of these geometric morphometric methods allowed the detection of differences in shape between these two species and to develop an easy-to-use identification index. We encourage the development of analogous indices that apply the methods of geometric morphometrics to distinguish between other species whose identification is complicated.     

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.