Getting a grip on the evolution of grasping in musteloid carnivorans: a three‐dimensional analysis of forelimb shape

The ability to grasp and manipulate is often considered a hallmark of hominins and associated with the evolution of their bipedal locomotion and tool use. Yet, many other mammals use their forelimbs to grasp and manipulate objects. Previous investigations have suggested that grasping may be derived from digging behaviour, arboreal locomotion or hunting behaviour. Here, we test the arboreal origin of grasping and investigate whether an arboreal lifestyle could confer a greater grasping ability in musteloid carnivorans. Moreover, we investigate the morphological adaptations related to grasping and the differences between arboreal species with different grasping abilities. We predict that if grasping is derived from an arboreal lifestyle, then the anatomical specializations of the forelimb for arboreality must be similar to those involved in grasping. We further predict that arboreal species with a well‐developed manipulation ability will have articulations that facilitate radio‐ulnar rotation. We use ancestral character state reconstructions of lifestyle and grasping ability to understand the evolution of both traits. Finally, we use a surface sliding semi‐landmark approach capable of quantifying the articulations in their full complexity. Our results largely confirm our predictions, demonstrating that musteloids with greater grasping skills differ markedly from others in the shape of their forelimb bones. These analyses further suggest that the evolution of an arboreal lifestyle likely preceded the development of enhanced grasping ability.     

Mode of miniaturisation influences body shape evolution in New World anchovies (Engraulidae)

We explored the macroevolutionary dynamics of miniaturisation in New World anchovies by integrating a time-calibrated phylogeny, geometric morphometrics and phylogenetic comparative methods. We found that the paedomorphic species Amazonsprattus scintilla occupies a novel region of shape space, while the dwarf species Anchoviella manamensis has an overall shape consistent with other anchovies. We found that miniaturisation did not increase overall clade disparity in size or shape beyond the expectations of Brownian motion, nor were there differences in rates of size or shape evolution among clades. Overall, our study shows that while the mode of miniaturisation influences shape evolution, the phenotypic novelty produced by the evolution of miniaturisation did not seem to alter macroevolutionary dynamics.     

Postmetamorphic ontogenetic allometry and the evolution of skull shape in Nest‐building frogs Leptodactylus (Anura: Leptodactylidae)

Allometry constitutes an important source of morphological variation. However, its influence in head development in anurans has been poorly explored. By using geometric morphometrics followed by statistical and comparative methods we analyzed patterns of allometric change during cranial postmetamorphic ontogeny in species of Nest‐building frogs Leptodactylus (Leptodactylidae). We found that the anuran skull is not a static structure, and allometry plays an important role in defining its shape in this group. Similar to other groups with biphasic life‐cycle, and following a general trend in vertebrates, ontogenetic changes mostly involve rearrangement in rostral, otoccipital, and suspensorium regions. Ontogenetic transformations are paralleled by shape changes associated with evolutionary change in size, such that the skulls of species of different intrageneric groups are scaled to each other, and small and large species show patterns of paedomorphic/peramorphic features, respectively. Allometric trajectories producing those phenotypes are highly evolvable though, with shape change direction and magnitude varying widely among clades, and irrespective of changes in absolute body size. These results reinforce the importance of large‐scale comparisons of growth patterns to understand the plasticity, evolution, and polarity of morphological changes in different clades.     

The influence of size on body shape diversification across Indo‐Pacific shore fishes*

Understanding the causes of body shape variability across the tree of life is one of the central issues surrounding the origins of biodiversity. One potential mechanism driving observed patterns of shape disparity is a strongly conserved relationship between size and shape. Conserved allometry has been shown to account for as much as 80% of shape variation in some vertebrate groups. Here, we quantify the amount of body shape disparity attributable to changes in body size across nearly 800 species of Indo‐Pacific shore fishes using a phylogenetic framework to analyze 17 geometric landmarks positioned to capture general body shape and functionally significant features. In marked contrast to other vertebrate lineages, we find that changes in body size only explain 2.9% of the body shape variation across fishes, ranging from 3% to 50% within our 11 sampled families. We also find a slight but significant trend of decreasing rates of shape evolution with increasing size. Our results suggest that the influence of size on fish shape has largely been overwhelmed by lineage‐specific patterns of diversification that have produced the modern landscape of highly diverse forms that we currently observe in nature.     

The effects of latitude,body size,and sexual selection on wing shape in a damselfly

Under natural selection, wing shape is expected to evolve to optimize flight performance. However, other selective factors besides flight performance may influence wing shape. One such factor could be sexual selection in wing sexual ornaments, which may lead to alternative variations in wing shape that are not necessarily related to flight performance. In the present study, we investigated wing shape variations in a calopterygid damselfly along a latitudinal gradient using geometric morphometrics. Both sexes show wing pigmentation, which is a known signal trait at intra‐ and interspecific levels. Wing shape differed between sexes and, within the same sex, the shape of the hind wing differed from the front wing. Latitude and body size explained a high percentage of the variation in wing shape for female front and hind wings, and male front wings. In male hind wings, wing pigmentation explained a high amount of the variation in wing shape. On the other hand, the variation in shape explained by pigmentation was very low in females. We suggest that the conservative morphology of front wings is maintained by natural selection operating on flight performance, whereas the sex‐specific differences in hind wings most likely could be explained by sexual selection. The observed sexual dimorphism in wing shape is likely a result of different sex‐specific behaviours. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 263–274.     

Postcranial element shape and function: assessing locomotor mode in extant and extinct mustelid carnivorans

Assessment of locomotor modes in fossil taxa must often be made on the basis of heavily fragmented postcranial material. Previous authors have used quantitative methods to determine locomotor function from whole postcranial elements. The goals of this project were to assess the ability of element shape to discern between locomotor modes through landmark analysis, and to apply the results to assessment of fossils. Results suggest that element shape is a good predictor of function, but that different elements have different predictive capacities for each locomotor mode. Additionally, a relationship between size and shape exists that appears to drive morphological differentiation in the group. Finally, data from the extant sample were applied to fossil material of the extinct Plio-Pleistocene taxon Trigonictis . The results suggest that the locomotor mode of Trigonictis was generalized and probably an intermediate between the half-bound locomotion found in weasels and ferrets and the scansorial locomotion of martens and fishers.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 895–914.     

Covariation between forelimb muscle anatomy and bone shape in an Australian scratch-digging marsupial: Comparison of morphometric methods

The close association between muscle and bone is broadly intuitive; however, details of the covariation between the two has not been comprehensively studied. Without quantitative understanding of how muscle anatomy influences bone shape, it is difficult to draw conclusions of the significance of many morphological traits of the skeleton. In this study, we investigated these relationships in the Quenda (Isoodon fusciventer), a scratch-digging marsupial. We quantified the relationships between forelimb muscle anatomy and bone shape for animals representing a range of body masses (124–1,952 g) using two-block partial least square analyses. Muscle anatomy was quantified as muscle mass and physiological cross-sectional area (PCSA), and we used two morphometric methods to characterize bone shape: seven indices of linear bone proportions, and landmarks analysis. Bone shape was significantly correlated with body mass, reflecting allometric bone growth. Of the seven bone indices, only shoulder moment index (SMI) and ulna robustness index (URI) showed a significant covariation with muscle anatomy. Stronger relationships between muscle anatomy and forelimb bone shape were found using the landmark coordinates: muscle mass and PCSA were correlated with the geometric shape of the scapula, humerus, and third metacarpal, but to a lesser extent with shape of the ulna. Overall, our data show that landmark coordinates are more sensitive than bone indices to capturing shape changes evident throughout ontogeny, and is therefore a more appropriate method to investigate covariation with forelimb muscle anatomy. Single-species studies investigating ontogeny require refined methods to accurately develop understanding of the important relationships between muscle force generation and bone shape remodeling. Landmark analyses provide such a method.     

Habitat‐specific trends in ontogeny of body shape in stickleback from coastal archipelago: Potential for rapid shifts in colonizing populations

We investigated ontogenetic trends in body shape of 54 freshwater (48 lake, seven stream) and six anadromous populations of threespine stickleback (Gasterosteus aculeatus L.) from the Haida Gwaii archipelago off the west coast of Canada. Multivariate analysis of covariance on the partial warp scores generated from 12 homologous landmarks on 1,958 digital images of subadult and adult male stickleback indicated that there was considerable variability of population ontogenetic slopes. We used discriminant function analysis to quantify body shape and determined that anadromous stickleback, which are ancestral to the freshwater populations, have a strongly negative ontogenetic slope (?5.62; increased streamlining with increased size). All freshwater populations exhibit a more positive slope (91% differed significantly from the marine slope), with the differences being most accentuated in populations from ponds and streams. In pristine lakes, ontogenetic slope could be predicted by lake volume as well as multivariate measures of habitat. Evidence from field transplant experiments of one of the intact populations indicates a rapid change (5 years) from allometric to isometric growth, equivalent to about half of the total slope variation among intact populations on the archipelago. We interpret this shift as developmental plasticity and suggest this may comprise the precursor for selection of optimal body shapes in these stickleback populations. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Diversification of body shape in Sebastes rockfishes of the north‐east Pacific

Body shape variation is integrally related to many aspects of fish ecology, including locomotion and foraging, and can indicate the functional diversity of fish assemblages. Few studies have thoroughly characterized body shape in a diverse marine fish clade, or investigated both temporal and spatial patterns of variation in body shape disparity. Here, I use digital photographs to measure geometric body shape in 66 species of north‐east Pacific rockfish (Sebastes spp.), including a correction for error introduced by arching of specimens. Different components of interspecific shape variation show associations with fish size, depth habitat, trophic niche and phylogenetic relationships. Overall, the accumulation of body shape disparity appears to have been near‐constant over time, and shows little variation across the latitudinal range of rockfish.     

Testing convergent and parallel adaptations in talpids humeral mechanical performance by means of geometric morphometrics and finite element analysis

The shape and mechanical performance in Talpidae humeri were studied by means of Geometric Morphometrics and Finite Element Analysis, including both extinct and extant taxa. The aim of this study was to test whether the ability to dig, quantified by humerus mechanical performance, was characterized by convergent or parallel adaptations in different clades of complex tunnel digger within Talpidae, that is, Talpinae+Condylura (monophyletic) and some complex tunnel diggers not belonging to this clade. Our results suggest that the pattern underlying Talpidae humerus evolution is evolutionary parallelism. However, this insight changed to true convergence when we tested an alternative phylogeny based on molecular data, with Condylura moved to a more basal phylogenetic position. Shape and performance analyses, as well as specific comparative methods, provided strong evidence that the ability to dig complex tunnels reached a functional optimum in distantly related taxa. This was also confirmed by the lower phenotypic variance in complex tunnel digger taxa, compared to non‐complex tunnel diggers. Evolutionary rates of phenotypic change showed a smooth deceleration in correspondence with the most recent common ancestor of the Talpinae+Condylura clade. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Evolutionary convergence of body shape and trophic morphology in cichlids from Lake Tanganyika

A recent phylogenetic analysis of mitochondrial DNA sequences from eretmodine cichlids from Lake Tanganyika indicated independent origins of strikingly similar trophic specializations, such as dentition characters. Because genetic lineages with similar trophic morphologies were not monophyletic, but instead were grouped with lineages with different trophic phenotypes, raises the question of whether trophic morphology covaries with additional morphological characters. Here, we quantified morphological variation in body shape and trophically associated traits among eretmodine cichlids using linear measurements, meristic counts and landmark‐based geometric morphometrics. A canonical variates analysis (CVA) delineated groups consistent with dentition characters. Multivariate regression and partial least squares analyses indicated that body shape was significantly associated with trophic morphology. When the phylogenetic relationships among taxa were taken into account using comparative methods, the covariation of body shape and trophic morphology persisted, indicating that phylogenetic relationships were not wholly responsible for the observed pattern. We hypothesize that trophic ecology may be a key factor promoting morphological differentiation, and postulate that similar body shape and feeding structures have evolved multiple times in independent lineages, enabling taxa to invade similar adaptive zones.     

Geometric morphometrics reveals body shape differences between sympatric redfish Sebastes mentella, Sebastes fasdatus and their hybrids in the Gulf of St Lawrence

Significant body shape differences were observed between two commercially important redfish species Sebastes fasdatus and Sebastes mentella , in the Gulf of St Lawrence, first classified according to the variability at the MDH-A* locus, a diagnostic criterion for the two species in allopatry. The observed sexual dimorphism was probably related to the presence of gravid females in the sample. Sebastes mentella appeared to be more fusiform than S. fasdatus , a characteristic that may reflect the more pelagic behaviour of S. mentella . The body shape of their hybrids was closer to that of S. mentella . Similarities between hybrids and S. mentella were also observed for meristic characteristics and depth distribution. The implications of these similarities for introgression are discussed. Body shape variability of specimens of hybrid origin was greater than that of either parental species, with features ranging from S. mentella-like to S. fasciatus -like. The role of introgression in generating morphological variability is emphasized.     

Indices of Anseriform body shape based on the relative size of major skeletal elements and their relationship to reproductive effort

To assess whether relative clutch mass (RCM) in Anseriformes (wildfowl or waterfowl) is constrained by body shape, principal components (PCs) of size‐adjusted measurements of five major skeletal elements of adult males and females of 60 species of Anseriformes provided indices of body shape. PC1 accounted for 69.8% of the variance and contrasted anterior elements (cranium and sternum) to posterior elements (synsacrum, femur and tibiotarsus). PC1 scores were high in species with a ‘duck‐like’ body shape and low in those with a ‘goose‐like’ body shape. Over the phylogeny of Anseriformes, decreased PC1 scores were associated with feeding on land. PC2 accounted for 18.6% of the variance and contrasted core elements (sternum and synsacrum) with peripheral elements (cranium, femur and tibiotarsus). High PC2 scores were associated with dependence on animal food, particularly in diving species. PC3 accounted for 7.7% of the variance and reflected mainly the relative size of the femur, which was low in diving species. Controlling statistically for phylogenetically independent contrasts in female body shape, there was a significant positive partial correlation between RCM and PC1, suggesting that independent of body size, body shape imposes constraints on reproductive effort in Anseriformes. The results suggest that models of the evolution of reproductive effort in this order, and perhaps in other orders, of birds should control for the effects of body shape.     

Body shape evolution among ploidy levels of the Squalius alburnoides hybrid complex (Teleostei,Cyprinidae)

Hybridization, ploidy level and genomic constitution may be important to respond to different environments, by producing different phenotypes and thus reducing competitive interaction. Through geometric morphometrics, we examined variation in body size and shape among biotypes of the Squalius alburnoides hybrid complex and their sperm donor (Squalius carolitertii). Results showed that S. carolitertii is significantly larger in size than the biotypes of the complex. No significant relationship was observed between ploidy and body size among S. alburnoides biotypes. Significant variation in body shape was found between S. carolitertii and S. alburnoides, and between tetraploids and the other biotypes. These differences in biotypes may reduce resource competition, highlighting the potential importance of resource availability favouring one biotype over another. In S. alburnoides, the adaptation to different trophic niches through modification of trophic morphology, body shapes, and feeding behaviour, may result from an increase in ploidy and genomic constitution. This adaptation may account also for the formation and maintenance of this nonsexual complex.