Morphological correlates of locomotor performance in hatchling Amblyrhynchus cristatus

Previous studies of locomotor performance from a variety of perspectives often assumed that speed and limb length were strongly correlated. Despite support of this assumption from biomechanical models, few empirical studies have demonstrated a significant relationship between measures of locomotor capacity, such as maximum velocity, and length of the hindlimb at either the inter- or intra-specific level. We examined whether one measure of locomotor performance, maximum velocity, correlates with body size and elements of the hindlimb in hatchling marine iguanas (Amblyrhynchus cristatus). Larger hatchlings ran faster. Removing the effects of body size revealed that relative lengths of the tibia and hindfoot correlated with size-adjusted maximum velocity. Individuals with relatively long tibia and short pes were relatively faster than individuals with short tibia and long pes. Functional morphological analyses predict that femur length should correlate with maximum velocity. However, our analyses failed to support this prediction. Because hatchling marine iguanas exploit relatively open habitats, the relationship between maximum velocity and limb morphology may be interpreted as an adaptation enhancing escape from predators.     

Locomotor Performance Varies With Adult Phenotype in Ornamented/Non‐Ornamented Wolf Spiders

Locomotor performance constitutes a major component of whole‐animal performance and is involved in several fitness‐related behaviors. Locomotor capabilities may also correspond positively or negatively to sexually selected traits (e.g., male ornamentation and/or courtship displays). Negative correlations are predicted if secondary sexual traits take the form of morphological modifications that impose physical or energetic limitations. We tested this cost of secondary sexual traits by comparing locomotor performance in male Schizocosa wolf spiders that exhibit two distinct phenotypes. These phenotypes vary in the presence/absence of a morphological feature assumed to function as sexual ornamentation—foreleg brushes. Given the conspicuously large brushes of hair on the brush‐legged phenotype, we expected these males to suffer in locomotor performance. We tested this cost by comparing locomotor performance among male phenotypes (brush‐legged and non‐ornamented) and females at immature and adult life stages. We did not find strong support for costs of brushes on locomotion. First, brush‐legged males showed similar average speeds and endurance as both non‐ornamented males and females. Second, while brush‐legged males were slower at maximum speeds than non‐ornamented males as matures (but not as immatures), they were no slower than mature females. Further, we found no variation in endurance among phenotypes or life stages. Finally, brush size did not correspond to speed. Patterns of morphological variation in traits other than ornamentation may explain these patterns: when morphological variation in leg lengths was accounted for, differences in maximum speed among groups disappeared. We suggest that the faster speeds achieved by non‐ornamented males arise as a by‐product of selection on morphology and musculature potentially necessary for vigorous courtship.     

The quick and the fast: the evolution of acceleration capacity in Anolis lizards

Although of prime ecological relevance, acceleration capacity is a poorly understood locomotor performance trait in terrestrial vertebrates. No empirical data exist on which design characteristics determine acceleration capacity among species and whether these design traits influence other aspects of locomotor performance. In this study we explore how acceleration capacity and sprint speed have evolved in Anolis lizards. We investigate whether the same or different morphological traits (i.e., limb dimensions and muscle mass) correlate with both locomotor traits. Within our sample of Anolis lizards, relative sprint speed and acceleration capacity coevolved. However, whereas the variation in relative acceleration capacity is primarily explained by the variation in relative knee extensor muscle mass, the variation in relative sprint speed is correlated to the variation in relative femur, tibia, and metatarsus length as well as knee extensor muscle mass. The fact that the design features required to excel in either performance trait partly overlap might explain the positive correlation between the variation in relative sprint speed and acceleration capacity. Furthermore, our data show how similar levels of sprint performance can be achieved through different morphological traits (limb segment lengths and muscle mass) suggesting that redundant mapping has potentially played a role in mitigating trade-offs.     

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.     

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.     

Potential targets for selection during the evolution of viviparity in cold-climate reptiles

Viviparity (live-bearing) has evolved from oviparity (egg-laying) in more than 100 lineages of squamate reptiles (lizards and snakes). This transition generally has occurred in cool climates, where thermal differentials between eggs in the (cool) nest versus the (warm) maternal oviduct influence embryonic development, in ways that may enhance offspring fitness. To identify specific traits potentially under selection, we incubated eggs of a montane scincid lizard at conditions simulating natural nests, maternal body temperatures, and an intermediate stage (2-week uterine retention of eggs prior to laying). Incubation at maternal temperatures throughout incubation affected the hatchling lizard’s activity level and boldness, as well as its developmental rate, morphology, and locomotor ability. A treatment that mimicked the initial stages of the transition toward viviparity had a major effect on some hatchling traits (locomotor speeds), a minor effect on others (tail length, total incubation period) and no effect on yet others (offspring behaviors). More generally, different aspects of the phenotype are sensitive to incubation conditions at different stages of development; thus, the evolution of reptilian viviparity may have been driven by a succession of advantages that accrued at different stages of embryogenesis.     

中华鳖新孵幼体的热耐受性、体温昼夜变化和运动能力的热依赖性

研究中华鳖新孵幼体的热耐受性、体温及温度对运动能力的影响 .结果表明 ,在干燥和潮湿环境下 ,选择体温分别为 2 8.0℃和 30 .3℃ ;潮湿环境下 ,临界高温和低温分别为 40 .9℃和 7.8℃ .在缺乏温度梯度的热环境中 ,水温对幼鳖体温的影响比气温更直接 ,体温和环境温度的昼夜变化相一致 ,说明幼鳖生理调温能力很弱 .在有温度梯度的热环境中 ,幼鳖能通过行为调温将体温维持到较高且较恒定的水平 ,导致体温昼夜变化不明显 .幼鳖运动能力有显著的热依赖性 ,在一定温度范围内随体温升高而增强 .体温31.5℃时 ,幼鳖的运动表现最好 ,最大续跑距离、单位时间跑动距离和单位时间停顿次数分别为 1.87m、4 92m·min-1和 6 .2次·min-1.体温过高时 ,运动能力下降 .当体温为 33 .0℃时 ,最大续跑距离、单位时间跑动距离和单位时间停顿次数分别为 1.30m、4.2 8m·min-1和 7.7次·min-1.     

Effect of mutations on developmental stability and canalization in morphological traits in Drosophila ananassae

The present study was designed to determine the effects of visible mutations of large effect on developmental stability and canalization in different morphological traits, namely, sternopleural bristle number, wing length, wing to thorax ratio, ovariole number, and sex comb tooth number (SCTN) in Drosophila ananassae. We have compared the mean trait size, fluctuating asymmetry (FA) (as an index of developmental stability), and morphological variation (as an index of canalization) of different mutant strains (yellow body color, y; claret eye color, ca; plexus wing, px; spread wing, spr; ebony body and sepia eye color, e se; yellow body and claret eye color, y ca; and cardinal eye color, curled wing, and ebony body color, cd cu e) with wild-type strain. The mean trait size of all morphological traits differs significantly among the wild-type and mutant strains. The wild-type and mutant strains vary significantly for the morphological variation and also for the levels of the FA in different morphological traits. However, we have found no increase in either the variance or in the degree of FA with the increase of the mutations (except in SCTN in y mutant). The plausible reasons for the variation in wild-type and mutant strains with particular reference to developmental stability and canalization have been discussed.     

四种利用不同生境蜥蜴运动能力的形态特征相关性

动物体态特征、功能表现和生境利用之间是否存在相关性是当前生态形态学领域的一个研究焦点。在实验室条件下测定分别利用开阔地面、草丛、岩石、树丛生境的 4种蜥蜴 (中国石龙子、北草蜥、山地麻蜥和变色树蜥 )的形态特征和运动能力 ,着重探讨蜥蜴运动能力与形态特征之间的相关性。 4种蜥蜴的头体长大小依次为 :中国石龙子 >变色树蜥 >北草蜥 >山地麻蜥。就相对体长而言 ,中国石龙子 >山地麻蜥和北草蜥 >变色树蜥 ,而头大小、附肢长度和尾长的种间差异趋势则相反 ;体高的种间差异为北草蜥 >中国石龙子和变色树蜥 >山地麻蜥。在平面上 ,山地麻蜥和北草蜥的速度显著大于中国石龙子和变色树蜥 ;在斜面上 ,变色树蜥和山地麻蜥的速度显著高于中国石龙子。变色树蜥斜面附着能力最强 ,中国石龙子最弱。生境利用不同的蜥蜴形态迥异 ,运动能力亦因此有显著的差异。本研究结果支持动物形态特征与其功能表现相关的观点。     

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.     

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.     

Effects of pregnancy on locomotor performance: an experimental study on lizards

Pregnancy is associated with reduced locomotor performance in many types of animals, but we do not know to what degree this correlation is caused by simple physical burdening, versus physiological changes associated with pregnancy, or to confounding variables (such as season or female body size) that simultaneously influence both reproductive investment and locomotor speeds. To identify causal effects of burden on locomotion, we need to experimentally manipulate the size of the load being carried. Injection of sterile fluid into the peritoneal cavities of 84 garden skinks (Lampropholis guichenoti) showed that speeds decreased with increasing burdens. Lizards with a burden equivalent to 25% of their body mass ran about 15% slower, mirroring the situation seen in gravid lizards of this population. Thus, simple physical burdening appears to be the primary causal component of the locomotor cost of reproduction within these animals. A lizard's sex, body size and shape had little effect on its running ability either before or after treatment, but faster lizards showed a greater performance decrement after burdening than did their slower conspecifics.     

Geographic variation in lizard phenotypes: importance of the incubation environment

Geographic variation in phenotypes can result from proximate environmental effects as well as from underlying genetic factors. Reciprocal transplant experiments, in which organisms are moved from one area to another, offer a powerful technique to partition the effects of these two factors. However, many studies that have utilized this technique have focused on the post-hatching organism only and ignored potential effects of environmental influences acting during embryonic development. We examined the phenotypic responses of hatchling scincid lizards ( Lampropholis guichenoti ) incubated in the laboratory under thermal regimes characteristic of natural nests in two study areas in southeastern Australia. Although the sites were less than 120 km apart, lizards from these two areas differed in thermal regimes of natural nests, and in hatchling phenotypes (morphology, locomotor performance). We incubated eggs from each area under the thermal regimes typical of both sites. Some of the traits we measured (e.g. hatchling mass and snout-vent length) showed little or no phenotypic plasticity in response to differences in incubation conditions, whereas other traits (e.g. incubation period, tail length, inter-limb length, body shape, locomotor performance) were strongly influenced by the thermal regime experienced by the embryo. Thus, a significant proportion of the geographic variation in morphology and locomotor performance of hatchling lizards may be directly induced by differences in nest temperatures rather than by genetic divergence. We suggest that future studies using the reciprocal transplant design should consider environmental influences on all stages of the life-history, including embryonic development as well as post-hatching life.     

Quantitative-morphometric and functional-anatomical studies on the locomotor apparatus of athletes. IV. Discussion, resume and literature

The locomotor system of 155 athletes from 3 athletic disciplines was examined, and the data of 20 anthropometrical measurements and the results for the corresponding disciplines were recorded subjected to a multivariate morphometrical analysis with the following aim: 1. To evaluate the morphological and functional-anatomical criteria of the locomotor system which quantified the variation differences in the body composition of the athletic groups, and which indicated a quantifiable functional-anatomical relationship between the variation in the body composition and the result in the corresponding disciplines. 2. To show the bivariate relationship between the morphological parameter and the results in the corresponding disciplines, and furthermore to evaluate the amount of result-increasing or result-decreasing components in a morphological variable. 3. To analyse the morphological variation of the 20 body composition variables and the corresponding results in the disciplines by means of factor analysis. This is achieved by the use of multivariate methods to analyse the variation in the athlete's body composition and the result-influencing-components. The following results were obtained: 1. There are partly pronounced morphological differences between the 3 athletic groups, quantified by discriminant functions. The differences in the locomotor system were reduced to 2 independent axes of variation, with one axis being identified as the general size variation between the 3 groups, and the other as the difference in the variation of the degree of strength of the musculature of the trunk and the limbs by considering the length and breadth measurements. For both canonical axes, the shotputters have above-average high canonical numbers compared to the other 2 groups. The sprinters and long jumpers show conformity in the general size variation of the locomotor system, but they differ in the second axis. Whilst in both groups the lower limbs are similarly constituted in form, shape and size, the canonical values of the trunk and the upper limbs of the sprinters are higher. In this respect, they tend more towards the shot-putter group. The morphological differences obtained for the athlete's locomotor system are functional-anatomically related to the corresponding athletic disciplines. 2. In almost every case, the regression functions for the shot-putters show an approximately linear relationship between the morphological variables and the result of the shot-putt. The quadratic and cubic regression functions only deviate slightly from the linearity, i.e. the result will be altered depending on the height and size of the body composition (within the possible range of biological variation). On the other hand, the sprinting result, either increased or decreased, is determined by a limited morphological variation of the parameter of the locomotor system...     

Geographic variation in 'costs of reproduction' in the scincid lizard Lampropholis guichenoti

1. A major focus of life-history research has been the analysis of reproductive effort (RE). However, while clearly defined in theory, RE has proved very difficult to measure. Consequently, researchers have looked for indices that estimate the components of RE. Uncritical use of indices of reproductive investment, such as measures of costs of reproduction, without verification of the underlying assumptions of the link between these traits, may lead to spurious conclusions. In this paper, the common assumption that the physical burden of the clutch impairs locomotor ability in gravid reptiles is examined.
2. Two neighbouring populations of Australian scincid lizards ( Lampropholis guichenoti) are similar in adult body sizes, body shapes and reproductive output (egg sizes, clutch sizes, relative clutch masses).
3. Despite morphological and reproductive similarities, the effects of pregnancy on maternal locomotor ability (running speeds, as measured in a laboratory raceway) differed dramatically between the two populations.
4. Lizards from the two populations ran at similar speeds when nongravid (i.e. after egg-laying), but pregnancy significantly reduced running speeds in one population and increased them in the other. Thus, superficial similarities in body size and reproductive output masked a strong divergence in the locomotor 'costs' of reproduction.
5. Caution is advised if using simple measures of reproductive output (e.g. relative clutch mass) as indices of reproductive effort, or to generalize results even among conspecific populations.     

The relationship between limb morphology, kinematics, and force during running: the evolution of locomotor dynamics in lizards

Terrestrial locomotion occurs via the hierarchical links between morphology, kinematics, force, and center-of-mass mechanics. In a phylogenetically broad sample of seven lizard species, we show that morphological variation drives kinematic variation, which, in turn, drives force variation. Species with short limbs use a short stride–high frequency strategy when running at steady-speed and to change speeds. This link between morphology and kinematics results in relatively small vertical forces during the support phase of the stride cycle. Conversely, species with long limbs use a long stride–low frequency strategy, resulting in large vertical forces during the support phase. In view of these findings, we suggest that limb length may predict locomotor energetics in lizards because energetics are largely determined by vertical forces and stride frequency. Additionally, we propose an energetic trade-off with both long- and short-limbed species paying the most energy to move, whereas intermediate-limbed species move using less energy. Finally, when these traits are mapped onto a lizard phylogeny, we show that locomotor functional morphology exhibits both deep phylogenetic effects and contemporary patterns of evolutionary convergence. Overall, the present study provides a foundation for testing hypotheses regarding the integration and evolution of functional traits in lizards and animals in general.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 634–651.     

Behavioural variation in natural populations. V. Morphological correlates of locomotion in the garter snake (Thamnophis radix)

A series of morphologieal and locomotor performance variables was measured in a population of newborn garter snakes to determine whether performance capacity has a significant morphological basis in these animals. Morphological traits measured were body length and mass, number of body and tail vertebrae and numbers of vertebral abnormalities. Locomotor performances included burst and mid-distance speed and distance and time crawled before anti-predator displays were assumed. All performance variables were repeatable in daily replicate trials ( P < 0.001). Individual burst speed, mid-distance speed, and distance crawled were significantly correlated pairwise ( P < 0.01). Most morphological and performance variables had a significant mass dependence (static allometry), although the effects were rather weak ( r ² < 0.1, except for body length): larger animals performed better and had fewer abnormalities. There were significant associations between some morphological traits and locomotor performance. Morphological factors accounted for 19% of the variation in mid-distance speed and 14% of the variation in antipredator behavior by multiple regression analysis. Canonical correlation of all performance and morphological variables simultaneously accounted for 24% of the observed variation in performance. Numbers of body and tail vertebrae (assayed by scale counts) had an interactive effect on speed of locomotion.     

Effects of constant and fluctuating temperatures on egg survival and hatchling traits in the northern grass lizard (Takydromus septentrionalis, Lacertidae)

To understand how nest temperatures influence phenotypic traits of reptilian hatchlings, the effects of fluctuating temperature on hatchling traits must be known. Most investigations, however, have only considered the effects of constant temperatures. We incubated eggs of Takydromus septentrionalis (Lacertidae) at constant (24 degrees C, 27 degrees C, 30 degrees C and 33 degrees C) and fluctuating temperatures to determine the effects of these thermal regimes on incubation duration, hatching success and hatchling traits (morphology and locomotor performance). Hatching success at 24 degrees C and 27 degrees C was higher, and hatchlings derived from these two temperatures were larger and performed better than their counterparts from 30 degrees C and 33 degrees C. Eggs incubated at fluctuating temperatures exhibited surprisingly high hatching success and also produced large and well-performed hatchlings in spite of the extremely wide range of temperatures (11.6-36.2 degrees C) they experienced. This means that exposure of eggs to adversely low or high temperatures for short periods does not increase embryonic mortality. The variance of fluctuating temperatures affected hatchling morphology and locomotor performance more evidently than did the mean of the temperatures in this case. The head size and sprint speed of the hatchlings increased with increasing variances of fluctuating temperatures. These results suggest that thermal variances significantly affect embryonic development and phenotypic traits of hatchling reptiles and are therefore ecologically meaningful.     

Morphological correlates of burst speed and field movement patterns: the behavioural adjustment of locomotion in wall lizards (Podarcis muralis)

Locomotion of lizards has clear morphological determinants and is important for developing activities such as feeding, social interaction and predator avoidance. Thus, morphological variation is believed to have fitness consequences through affecting locomotor performance. This paper firstly evaluates the dependence of burst speed on morphology, and secondly examines the movement patterns of free-ranging undisturbed wall lizards ( Podarcis muralis ) engaged in several kinds of activity. Body size was the most important correlate of burst speed as performed at the optimal temperature for running in the laboratory. After removing size effects from performance and morphological traits, the length of some particular limb segments had positive influence on burst speed, but these effects were weak, each trait explaining less than 16% of variance in burst speed. Free-ranging P. muralis exhibited intermittent locomotion, with movement sequences interrupted by frequent short pauses. Field movement patterns greatly differed depending upon the kind of activity and were in most aspects independent of the size and sex of the animal. P. muralis involved in thermoregulation performed short and low-speed displacements; exploratory activities were characterized by frequent, slow and short movements. On the contrary, lizards involved in intraspecific pursuits and predator escape developed comparatively high speeds, although only exceptionally did they attain the size-specific burst speed predicted from the laboratory trials. Speed of escape increased with distance to the refuge and the animals are able to assess predation risks to modulate approach distance, speed and pauses, so maximum exertion is seldom required. The evolution of locomotor capacities exceeding routine needs is discussed in the context of the principle of 'excessive construction'.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 135–146.