Sexual differences in locomotor performance in Tropidurus catalanensis lizards (Squamata: Tropiduridae) – body shape,size and limb musculature explain variation between males and females

Sexual dimorphism (SD) is the evolutionary outcome of selection acting differently on males and females. Several studies describe sexual differences in body size, although other morphological traits might be allometric between sexes and imply functional consequences. Here we test whether morphological differences between sexes in size and shape in the lizard Tropidurus catalanensis explain variation in performance of four locomotor traits. Our results show that males are larger than females and also exhibit longer limbs, longer muscles and larger muscle cross‐sectional areas, while females have longer trunks and more sharped anterior claws; males outperform females in all locomotor performances measured. Sexual differences in sprinting and climbing is related with body size, and climbing performance is also explained by limb lengths, by differences in lengths and cross‐sectional areas of specific muscles, and by interlimb distances. Between‐sex differences in exertion are also related to SD, despite associations with sharper posterior claws that are independent of sex. Grasping performance, however, is associated with some muscle and morphological parameters that are not sexually dimorphic. Together our results suggest that morphology might be under sexual selection in T. catalanensis, given that better locomotor performance likely favours male lizards in typical activities of this polygenic species, such as territory defence and female acquisition. Moreover, the longer trunks that characterize females may confer more space to accommodate eggs. On the other hand, territory defence by males probably increases their exposure to predators, resulting in a synergistic effect of sexual and natural selection in the evolution of SD in T. catalanensis.     

Sexual dimorphism in traits related to locomotion: ontogenetic patterns of variation in Podarcis wall lizards

Sexual dimorphism in body size and shape in animals is normally linked to sexual selection mechanisms that modify the morphological properties of each sex. However, sexual dimorphism of ecologically relevant traits may be amplified by natural selection and result in the ecological segregation of both sexes. In the present study, we investigated patterns of sexual dimorphism of morphological traits relevant for locomotion in two lacertid lizards, Podarcis bocagei and Podarcis carbonelli, aiming to identify ontogenetic sources of variation. We analysed trunk and limb variation in relation to total body size, as well as the covariation of different traits, aiming to shed light on the proximate causation of adult sexual dimorphism. We find that, although immatures are generally monomorphic, adult females have a longer trunk, and adult males have longer fore and hind limbs. Both sexes differ substantially with respect to their growth trajectories and relationships between traits, whereas, in some cases, there are signs of morphological constraints delimiting the observed patterns. Because of the direct connection between limb size/shape and locomotor performance, which is relevant both for habitat use and escape from predators, the observed patterns of sexual dimorphism are expected to translate into ecological differences between both sexes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 530–543.     

Skeletal indicators of locomotor adaptations in living and extinct rodents

Living rodents show great diversity in their locomotor habits, including semiaquatic, arboreal, fossorial, ricochetal, and gliding species from multiple families. To assess the association between limb morphology and locomotor habits, the appendicular skeletons of 65 rodent genera from 16 families were measured. Ecomorphological analyses of various locomotor types revealed consistent differences in postcranial skeletal morphology that relate to functionally important traits. Behaviorally similar taxa showed convergent morphological characters, despite distinct evolutionary histories. Semiaquatic rodents displayed relatively robust bones, enlarged muscular attachments, short femora, and elongate hind feet. Arboreal rodents had relatively elongate humeri and digits, short olecranon processes of the ulnae, and equally proportioned fore and hind limbs. Fossorial rodents showed relatively robust bones, enlarged muscular attachments, short antebrachii and digits, elongate manual claws, and reduced hind limb elements. Ricochetal rodents displayed relatively proximal insertion of muscles, disproportionate limbs, elongate tibiae, and elongate hind feet. Gliding rodents had relatively elongate and gracile bones, short olecranon processes of the ulnae, and equally proportioned fore and hind limbs. The morphological differences observed here can readily be used to discriminate extant rodents with different locomotor strategies. This suggests that the method could be applied to extinct rodents, regardless of ancestry, to accurately infer their locomotor ecologies. When applied to an extinct group of rodents, we found two distinct ecomorphs represented in the beaver family (Castoridae), semiaquatic and semifossorial. There was also a progressive trend toward increased body size and increased aquatic specialization in the giant beaver lineage (Castoroidinae). J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Morphological variation does not influence locomotor performance within a cohort of hatchling lizards (Amphibolurus muricatus, Agamidae)

A causal link between morphology and performance is a central tenet of ecomorphological analyses, but there are few detailed analyses of exactly how morphological variation within a hatchling cohort maps onto locomotor performance, and especially whether or not different tasks favour different morphologies (or vice versa). We measured morphological traits (including body length, mass, head size, limb proportions and fluctuating asymmetry [FA]) on a large sample of laboratory-incubated hatchling lizards ( Amphibolurus muricatus , Agamidae), and used principal component analysis to reduce this data set to four major axes of variation (size, shape and two FA axes). Running speeds of each lizard were measured on raceways at four inclines, from level (0°) through to steep (45°). Unsurprisingly, steeper inclines reduced locomotor speeds. Absolute body size was the only morphological trait that was consistently related to sprinting performance, and the relationships were similar at each incline. Within-cohort variation in body shape and FA among this large sample was unrelated to locomotor speeds, thus challenging the common assumption of a causal link between these variables. The only exception was a weak trend for greater hind limb length to enhance locomotor performance more at steep inclines than at shallower angles. In general, our data suggest that different morphological traits do not differentially maximize locomotor performance up variable inclines. Overall, our data provide a cautionary note about the generality of causal connections between within-cohort morphological variation and locomotor performance under different environmental contexts.     

Morphological and functional differentiation in the lumbar spine of lorisids and galagids

The striking contrast in positional behavior exhibited by lorisids (slow quadrupedalism/suspension) and galagids (leaping/quadrupedalism) is well reflected in their postcranial morphology, particularly in the limbs. Although they exhibit very different spinal postures and movements, vertebral adaptations have been less well explored in these taxa. This study addressed morphological and functional differentiation in the lumbar vertebrae of four species of lorisids and five species of galagids. Linear and angular measurements of lumbar vertebrae were compared among taxa using canonical variates analysis (CVA) in conjunction with pairwise comparisons among selected variables. The results were interpreted in the context of a broader comparative sample, including the addition of indriids to the CVA. Compared to galagids, lorisids have relatively shorter lumbar spinous processes that are more perpendicularly (to caudally) oriented relative to a coronal plane. Lorisids also have relatively wider laminae and more transversely oriented prezygapophyses. These features promote lumbar stability and reflect antipronogrady, multiplane spinal movements, and upside-down suspension. Within lorisids, vertebral body length and height vary with body size, reflecting the additional resistance to bending that is required for larger body sizes. Galagid lumbar shape is influenced by body size, but does not show strong variation in accordance with positional behavior differences as defined here. Galagids, indriids, and lorisids are distinct in lumbar morphology and function, but their similarities in lumbar length reduction are suggestive of antipronograde postures in the common ancestor of the galagids, including those who have shifted to a more quadrupedal locomotor repertoire.     

Climatic variability and body size variation in the muskrats (Ondatra zibethicus) of North America

Summary Patterns of geographic variation in nine morphological characters of adult muskrats (Ondatra zibethicus) are investigated by multiple regression and canonical correlation analysis. Three variables describing the annual precipitation regimen account for 16 to 33% of the variance in each of the skeletal measurements. Highly variable precipitation patterns result in low rates of germination and survivorship for preferred aquatic food plants which in turn reduces average total food availability for muskrats. In large individuals nutritive demands may exceed supply, thus selection favors smaller body size in areas of low food availability.After removing the variation attributable to three precipitation variables, the residual variation in the morphological variables is exposed to canonical correlation analysis with a set of 10 environmental and geographic variables. A canonical variate loaded for climatic seasonality accounts for 60% of the variance in a canonical variate of the morphological residuals which load as a general body size variate. It is argued that seasonality is a major factor selecting for large body size in muskrats and other organisms. During seasonal periods of resource abundance natural selection favors individuals with rapid growth to a large size, while concurrently enhancing survivorship through oncoming periods of resource shortage.     

Echocardiographic changes in the development of the athlete' s heart in 9 to 20-year-old male subjects

The purpose of this cross-sectional investigation was to estimate the age at which specific traits of the "athlete's heart" first appear and how they evolve from the beginning of regular physical training until young adulthood in healthy active males. Male athletes (n=389) and non-athletes (n=55) aged between 9 and 20 years were examined by two-dimensionally guided M-mode and Doppler echocardiography. Intragroup differences were examined by t-tests for independent samples between age groups of two years each. Morphologic variables were related to body size by using ratio indices in which the power terms of numerator and denominator were matched. Relative left ventricular muscle mass (LVMM) was significantly larger in the athletic males at age of 11-12, and this significant difference was maintained with advancing age. Most of this increase of LVMM could be attributed to the increase in wall thickness that became significantly manifest first in the 13- to 14-year-old athletic subjects but was demonstrable in all the other groups. A significantly larger left ventricular internal diameter was only found in the age-group of 15-16. Fractional shortening percentage (FS%) did not show any change, while resting heart rate was decreased in our athletic groups.     

Modular diversification of the locomotor system in damselfishes (Pomacentridae)

As fish move and interact with their aquatic environment by swimming, small morphological variations of the locomotor system can have profound implications on fitness. Damselfishes (Pomacentridae) have inhabited coral reef ecosystems for more than 50 million years. As such, habitat preferences and behavior could significantly constrain the morphology and evolvability of the locomotor system. To test this hypothesis, we used phylogenetic comparative methods on morphometric, ecological and behavioral data. While body elongation represented the primary source of variation in the locomotor system of damselfishes, results also showed a diverse suite of morphological combinations between extreme morphologies. Results show clear associations between behavior, habitat preferences, and morphology, suggesting ecological constraints on shape diversification of the locomotor system. In addition, results indicate that the three modules of the locomotor system are weakly correlated, resulting in versatile and independent characters. These results suggest that Pomacentridae is shape may result from the interaction between (1) integrated parts of morphological variation that maintain overall swimming ability and (2) relatively independent parts of the morphology that facilitate adaptation and diversification. J. Morphol. 277:603–614, 2016. © 2016 Wiley Periodicals, Inc.     

Variation in morphology and functional performance across distinct evolutionary lineages of the Moorish gecko (Tarentola mauritanica) from the Iberian Peninsula

Deciphering the mechanisms that underlie morphological and functional diversity is essential for understanding how organisms adapt to their environment. Interestingly, phenotypic divergence does not necessarily correspond to the geographic and genetic separation between populations. Here, we explored the morphological and functional divergence among populations of two genetically differentiated clades of the Moorish gecko, Tarentola mauritanica. We used linear and geometric morphometrics to quantify morphological variation and investigated how it translates into biting and CLIMBING PERFORMANCE, to better understand the mechanisms potentially underlying population and lineage divergence. We found marked morphological differences between clades, both in body size and head shape. However, much of this differentiation is more strongly related to local variation between populations of the same clade, suggesting that recent ecological events may be more influential than deep evolutionary history in shaping diversity patterns in this group. Despite a lack of association between morphology and functional diversification in the locomotor system of the Moorish gecko, straightforward links are observed between head morphology and biting performance, providing more hints on the possible underlying causes. Indeed, variation in bite force is mostly determined by size variation and sexual dimorphism, and differences between the two clades concern how sexual variation is expressed, reinforcing the idea that both social and ecological factors contribute in shaping differentiation. Interestingly, the individuals from the islets off the coast of Murcia exhibit particular morphological and functional traits, which suggests that the ecological conditions related to insularity may drive the phenotypic differentiation of this population.     

Leaping behavior of Pithecia pithecia and Chiropotes satanas in eastern Venezuela

I observed leaping behavior in the white-faced saki (Pithecia pithecia) and the black-bearded saki (Chiropotes satanas satanas) for 15 and 10 months, respectively, as part of a larger study of positional behavior in the tribe Pitheciini. I used focal animal instantaneous sampling to observe the two species on separate islands in their natural habitat at Guri Lake, Venezuela. Leaping behavior correlates with patterns of forest use and body size, and differences between the species relate more to habitat preferences than to habitat differences per se. Pithecia usually chose vertical or highly angled supports of lower tree portions for take-off and landing, and took off from a stationary posture. Chiropotes took off from the main crown or terminal branches, gaining momentum from locomotor movement before performing a leaping take-off. Pithecia's vertical body orientation and longer leap distance allowed it to assume a mid-flight tuck to prepare for a hindlimb-first landing onto a solid support, and to absorb landing forces with its relatively longer hindlimbs. Chiropotes remained more pronograde throughout its leaps, and minimized landing forces by landing on all four limbs onto numerous flexible supports in the terminal branches. The smaller-bodied P. pithecia is specialized for vertical clinging and leaping, and exhibits behavioral and morphological parallels with other vertical clingers and leapers. The larger C. satanas is a generalized leaper that lacks morphological specializations for leaping. Pithecia's use of solid supports in the lower tree portions allows it to move quietly through the forest-one of a suite of behaviors related to predator avoidance. This example of variation within one behavioral category has implications for devising locomotor classifications and interpreting fossil remains.     

Evolutionary relationships between morphology, performance and habitat openness in the lizard genus Niveoscincus (Scincidae: Lygosominae)

Correlations between an animal's morphology and ecological parameters such as habitat characteristics emphasize the intimate link between phenotype and the environment, but are often difficult to interpret because the functional consequences of morphological variation are frequently unknown. We provide one of the few studies relating limb morphology, functional capabilities, and habitat in reptiles. We tested the hypothesis that species occupying open microhabitats would possess relatively longer limbs and faster sprint speeds than those occurring in more closed microhabitats. A number of morphological characteristics relevant to locomotion were quantified, including the length of the bones of the fore- and hindlimbs and body size. A phylogenetic analysis was then used to examine the evolutionary relationships between morphology, locomotor performance and microhabitat openness in seven species of Niveoscincus and one species of the closely related genus Pseudemoia. A significant evolutionary relationship was established between sprinting ability, morphology, and the openness of the microhabitat occupied by a species. The phylogenetic analysis demonstrated an evolutionary trend in Niveoscincus of species occupying open microhabitats (e.g. N. greeni, N ocellatus) being large with long limbs and high sprinting ability, while those occupying closed microhabitats (e.g. N coventryi, P. entrecasteauxii) art smaller with short limbs and much slower maximum sprint speeds.     

The relevance of morphology for habitat use and locomotion in two species of wall lizards

Understanding if morphological differences between organisms that occupy different environments are associated to differences in functional performance can suggest a functional link between environmental and morphological variation. In this study we examined three components of the ecomorphological paradigm – morphology, locomotor performance and habitat use – using two syntopic wall lizards endemic to the Iberian Peninsula as a case study to establish whether morphological variation is associated with habitat use and determine the potential relevance of locomotor performance for such an association. Differences in habitat use between both lizards matched patterns of morphological variation. Indeed, individuals of Podarcis guadarramae lusitanicus, which are more flattened, used more rocky environments, whereas Podarcis bocagei, which have higher heads, used more vegetation than rocks. These patterns translated into a significant association between morphology and habitat use. Nevertheless, the two species were only differentiated in some of the functional traits quantified, and locomotor performance did not exhibit an association with morphological traits. Our results suggest that the link between morphology and habitat use is mediated by refuge use, rather than locomotor performance, in this system, and advise caution when extrapolating morphology-performance-environment associations across organisms.     

Ecological and developmental context of natural selection: maternal effects and thermally induced plasticity in the frog Bombina orientalis

Variation in fitness generated by differences in functional performance can often be traced to morphological variation among individuals within natural populations. However, morphological variation itself is strongly influenced by environmental factors (e.g., temperature) and maternal effects (e.g., variation in egg size). Understanding the full ecological context of individual variation and natural selection therefore requires an integrated view of how the interaction between the environment and development structures differences in morphology, performance, and fitness. Here we use naturally occurring environmental and maternal variation in the frog Bombina orientalis in South Korea to show that ovum size, average temperature, and variance in temperature during the early developmental period affect body sizes, shapes, locomotor performance, and ultimately the probability of an individual surviving interspecific predation in predictable but nonadditive ways. Specifically, environmental variability can significantly change the relationship between maternal investment in offspring and offspring fitness so that increased maternal investment can actually negatively affect offspring over a broad range of environments. Integrating environmental variation and developmental processes into traditional approaches of studying phenotypic variation and natural selection is likely to provide a more complete picture of the ecological context of evolutionary change.     

Evolution of sexual dimorphism in the digit ratio 2D:4D--relationships with body size and microhabitat use in iguanian lizards

The ratio between lengths of digit II and IV (digit ratio 2D:4D) is a morphological feature that likely affects tetrapod locomotor performances in different microhabitats. Modifications of this trait may be triggered by changes in steroids concentrations during embryo development, which might reflect direct selection acting on digit ratio or be solely a consequence of hormonal differences related for example to body size. Here we apply both conventional and phylogenetic analyses on morphological data from 25 lizard species of 3 families of Iguania (Iguanidae, Polychrotidae, and Tropiduridae), in order to verify whether selective pressures related to locomotion in different microhabitats could override the prenatal developmental cues imposed on the digit ratio 2D:4D by differences in body size between males and females. Data suggest that this trait evolved in association with ecological divergence in the species studied, despite the clear effect of body size on the digit ratio 2D:4D. The ecological associations of size-corrected digit ratios were restricted to one sex, and females of species that often use perches exhibited small digit ratios in the front limbs, which translated into larger sexual dimorphism indexes of arboreal species. The results, together with the subsequent discussion, provide outlines for further investigation about possible developmental mechanisms related to the evolution of adaptive changes in digit lengths that may have occurred during the evolution of ecological divergence in squamates.     

Transitions in functional morphology from “large branchiopods” to Cladocera: Video and confocal microscopic studies of Cyclestheria hislopi (Cyclestherida) and Sida crystallina (Cladocera: Ctenopoda)

Great diversity is found in morphology and functionality of arthropod appendages, both along the body axis of individual animals and between different life-cycle stages. Despite many branchiopod crustaceans being well known for displaying a relatively simple arrangement of many serially post-maxillary appendages (trunk limbs), this taxon also shows an often unappreciated large variation in appendage morphology. Diplostracan branchiopods exhibit generally a division of labor into locomotory antennae and feeding/filtratory post-maxillary appendages (trunk limbs). We here study the functionality and morphology of the swimming antennae and feeding appendages in clam shrimps and cladocerans and analyze the findings in an evolutionary context (e.g., possible progenetic origin of Cladocera). We focus on Cyclestheria hislopi (Cyclestherida), sister species to Cladocera and exhibiting many “large” branchiopod characters (e.g., many serially similar appendages), and Sida crystallina (Cladocera, Ctenopoda), which likely exhibits plesiomorphic cladoceran traits (e.g., six pairs of serially similar appendages). We combine (semi-)high-speed recordings of behavior with confocal laser scanning microscopy analyses of musculature to infer functionality and homologies of locomotory and filtratory appendages in the two groups. Our morphological study shows that the musculature in all trunk limbs (irrespective of limb size) of both C. hislopi and S. crystallina comprises overall similar muscle groups in largely corresponding arrangements. Some differences between C. hislopi and S. crystallina, such as fewer trunk limbs and antennal segments in the latter, may reflect a progenetic origin of Cladocera. Other differences seem related to the appearance of a specialized type of swimming and feeding in Cladocera, where the anterior locomotory system (antennae) and the posterior feeding system (trunk limbs) have become fully separated functionally from each other. This separation is likely one explanation for the omnipresence of cladocerans, which have conquered both freshwater and marine free water masses and a number of other habitats.     

Size dimorphism in Rankinia [Tympanocryptis] diemensis (Family Agamidae): sex-specific patterns and geographic variation

Sexual dimorphism has implications for a range of biological and ecological factors, and intersexual morphological differences within a species provide an ideal opportunity for investigating evolutionary influences on phenotypic variation. We investigated sexual size dimorphism (SSD) in an agamid species, Rankinia [Tympanocryptis] diemensis , to determine whether overall size and/or relative morphological trait size differences exist and whether geographic variation in size dimorphism occurs in this species. Relative morphological trait proportions included a range of head, limb, and inter-limb measurements. We found significant overall intersexual adult size differences; females were the larger sex across all sites but the degree of dimorphism between the sexes did not differ between sites. This female-biased size difference is atypical for agamid lizards, which are usually characterized by large male body size. In this species, large female-biased SSD appears to have evolved as a result of fecundity advantages. The size of relative morphological trait also differed significantly between the sexes, but in the opposite direction: relative head, tail, and limb sizes were significantly larger in males than females. This corresponds to patterns in trait size usually found in this taxonomic group, where male head and limb size is important in contest success such as male–male rivalry. There were site-specific morphological differences in hatchlings, including overall body size, tail, inter-limb, thigh, and hindlimb lengths; however, there were no sex-specific differences indicating the body size differences present in the adult form occur during ontogeny.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 699–709.     

Body size and shape evolution in host races of the tick Ixodes uriae

The tick Ixodes uriae is a common ectoparasite of seabirds, and is widely distributed across the circumpolar regions of both hemispheres. Previous work demonstrated the existence of genetically distinct host races of this ectoparasite, occurring across its current range. The objective of the present study was to examine whether these host races have evolved measurable morphological differences. We measured a set of morphological variables on 255 non‐engorged ticks (nymphs and adults) collected from three sympatrically occurring host species in the North Atlantic. Genotyping at eight microsatellite markers enabled us to analyse the relationship between patterns of morphological and neutral genetic variation. Multivariate analyses showed that most morphological variation was associated with size differences among tick individuals. Body size differed among races, but only in adult life stages. A linear discriminant analysis based on shape variation revealed three distinct morphological clusters corresponding to the three tick host races. These results, along with correlated patterns of host‐related genetic variation, suggest that differences among host‐related groups are not simply the result of phenotypic plasticity or drift, but rather reflect host‐associated adaptations. Experimental work and observations across the range of I. uriae will now be required to test the genetic basis and adaptive nature of morphological differences.     

Sports anthropological investigations on somatotypology of elite karateka

The goal of this work was to find differences between the somatotypes of elite karateka. For this purpose the somatotypes of Conrad, Heath & Carter were used. This sports anthropological study was carried out on the assumption that constant karate training causes changes in body composition. It should clarify whether different body types, including sexual dimorphism, are to be found in people practicing the karate disciplines kata and kumite. 80 male and female elite karateka were tested. As a comparison group, 62 leisure sports persons and 66 hobby karateka were used as control groups. The measurements were taken under standardised conditions and the results were examined statistically (ANOVA). The body types of Conrad, which were confirmed by the somatocharts of Heath & Carter, showed differences between the elite karateka in comparison to the control group. According to this, the typical elite karateka is more athletic and smaller. He weighs less than the sports persons of the comparison fitness group. Moreover, within the karate disciplines, the kata practisers are more endomorph than their colleagues. The kumite athletes take more ektomorph positions in the somatocharts (Heath & Carter 1990). On the basis of these results the assumption of a differentiated somatotype can be confirmed in the two groups and also within karate between the disciplines kata and kumite. This study concludes that there is both a kata and a kumite somatotype among karate practisers. Further research is required regarding longitudinal measurements of the change in body composition of young karateka in the course of their competitive careers. The career-affecting factors of training frequency, training age, and frequency of injury, related to the level of achievement, leave room for further investigation. This also applies to possible ethnic body-build differences and the athletes' level of achievement.     

Ontogeny of sexual dimorphism and phenotypic integration in heritable morphs

In this study we investigated the developmental basis of adult phenotypes in a non-model organism, a polymorphic damselfly (Ischnura elegans) with three female colour morphs. This polymorphic species presents an ideal opportunity to study intraspecific variation in growth trajectories, morphological variation in size and shape during the course of ontogeny, and to relate these juvenile differences to the phenotypic differences of the discrete adult phenotypes; the two sexes and the three female morphs. We raised larvae of different families in individual enclosures in the laboratory, and traced morphological changes during the course of ontogeny. We used principal components analysis to examine the effects of Sex, Maternal morph, and Own morph on body size and body shape. We also investigated the larval fitness consequences of variation in size and shape by relating these factors to emergence success. Females grew faster than males and were larger as adults, and there was sexual dimorphism in body shape in both larval and adult stages. There were also significant effects of both maternal morph and own morph on growth rate and body shape in the larval stage. There were significant differences in body shape, but not body size, between the adult female morphs, indicating phenotypic integration between colour, melanin patterning, and body shape. Individuals that emerged successfully grew faster and had different body shape in the larval stage, indicating internal (non-ecological) selection on larval morphology. Overall, morphological differences between individuals at the larval stage carried over to the adult stage. Thus, selection in the larval stage can potentially result in correlated responses in adult phenotypes and vice versa.