Limb and tail lengths in relation to substrate usage in Tropidurus lizards

A close relationship between morphology and habitat is well documented for anoline lizards. To test the generality of this relationship in lizards, snout-vent, tail, and limb lengths of 18 species of Tropidurus (Tropiduridae) were measured and comparisons made between body proportions and substrate usage. Phylogenetic analysis of covariance by computer simulation suggests that the three species inhabiting sandy soils have relatively longer feet than do other species. Phylogenetic ANCOVA also demonstrates that the three species inhabiting tree canopies and locomoting on small branches have short tails and hind limbs. These three species constitute a single subclade within the overall Tropidurus phylogeny and analyses with independent contrasts indicate that divergence in relative tail and hind limb length has been rapid since they split from their sister clade. Being restricted to a single subclade, the difference in body proportions could logically be interpreted as either an adaptation to the clade's lifestyle or simply a nonadaptive synapomorphy for this lineage. Nevertheless, previous comparative studies of another clade of lizards (Anolis) as well as experimental studies of Sceloporus lizards sprinting on rods of different diameters support the adaptive interpretation.     

Limb proportions in climbing and ground-dwelling geckos (Lepidosauria, Gekkonidae): a phylogenetically informed analysis

Biomechanical considerations predict that limb proportions should differ between animals with climbing and ground-dwelling lifestyles. Ground-dwellers should have relatively long, parasagittal hind limbs, with high tibia:femur ratios, and relatively short fore limbs. Climbers should have relatively short limbs, with low tibia:femur ratios, and equally long hind and fore limbs. We tested these predictions using gecko species with different locomotion habits (climbing versus ground-dwelling). We measured snout-vent length and lengths of limb segments in 29 species of geckos and analysed them using both non-phylogenetic statistics (nested analysis of variance and principal component analysis) and phylogenetic statistics (analysis of covariance). Neither approach allowed us to find any consistent relationship between habitat use and the morphometric variables. We conclude that either relative limb lengths and limb proportions in geckos have not evolved in response to the physical demands of the microhabitat, or our understanding of those demands is insufficient. Accepted: 22 February 2001     

Ontogenetic and individual variation in size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis

The present study investigates relationships among size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis . Maximal running speed, body mass, snout-vent length, tail length, fore- and hind limb spans and thigh muscle mass were measured in 68 field-fresh individuals spanning the entire ontogenetic size range (1.3 48 g). Relative lengths of both foreand hind limbs decrease with increasing body mass (= negative allometry), whereas relative tail length and thigh muscle mass increase with body mass (= positive allometry). Repeatable and significant differences in maximal running speed exist among individuals. Maximal running speed scales as (body mass)^0.161, and 59% of the variation in maximal speed was related to body mass. Based on the results of the present and previous studies, data on scaling of body proportions alone appear inadequate to infer scaling relationships of functional characters such as top speed.
Surprisingly, individual variation in maximal speed is not related to individual variation in shape (relative limb, tail and body lengths). These components of overall shape are not independent; individuals tended to have either relatively long or relatively short limbs, tails and bodies for their body mass. Even the significant difference in multivariate shape between adult males and females has no measurable consequences for maximal speed. Speeds of field-fresh animals did not vary on a seasonal basis, and eight weeks of captivity had no effect on maximal running speeds. Gravid females and long-term (obese) captive lizards were both approximately 12% slower than field-fresh lizards.     

Prey-handling and the evolutionary ecology of sand-swimming lizards (Lerista : Scincidae)

Fossorial lizards differ in morphology from their surface-dwelling relatives. The Australian sphenomorphine skink genus Ctenotus consists of surface-dwelling species, and is closely related to the genus Lerista, which includes both surface-dwelling and fossorial species. Sand-swimming represents the derived condition and has evolved independently in several lineages of Lerista. The heads of lizards in the two genera differ in shape (blunt snout for Ctenotus versus wedge-shaped for Lerista) and in length relative to the body (approximately 20% of snout-vent length for Ctenotus versus 12% for sand-swimming Lerista). Do these specializations affect the sizes or types of prey that can be consumed by Lerista? We compared prey-handling by Ctenotus and Lerista to correlate morphological differences with differences in prey-handling ability, and to distinguish the effects of snout shape and head length. Feeding trials included three categories of insect prey that the lizards normally eat: soft-bodied larvae (Lepidoptera), hard-bodied larvae (Coleoptera), and roaches (Blatoidea). In comparisons based on the mass of a prey item relative to the mass of a lizard, Lerista had longer handling times for all prey categories and were limited to smaller prey than were Ctenotus. However, when comparisons were based on the length of prey relative to the length of a lizard's head, Lerista ate some elongate prey as fast or faster than did Ctenotus, and both genera successfully swallowed prey more than twice the length of their own head. Thus, the differences in prey-handling performance of Ctenotus and Lerista probably result from the fact that Lerista have a relatively shorter head than Ctenotus. All Lerista species, surface-dwelling and fossorial, have short heads compared to primitive sphenomorphine lizards. Fossorial species of Lerista have elongate trunks, and consequently their heads are shorter in proportion to trunk length than those of surface-dwelling Lerista. However, most fossorial species of Lerista are longer and heavier than any of their surface-dwelling congeners, and the heads of these fossorial species are large relative to the prey they encounter. As a consequence, the diets of large fossorial species of Lerista do not appear to be limited by their morphological specialization for sand-swimming.     

Shared extremes by ectotherms and endotherms: Body elongation in mustelids is associated with small size and reduced limbs

An elongate body with reduced or absent limbs has evolved independently in many ectothermic vertebrate lineages. While much effort has been spent examining the morphological pathways to elongation in these clades, quantitative investigations into the evolution of elongation in endothermic clades are lacking. We quantified body shape in 61 musteloid mammals (red panda, skunks, raccoons, and weasels) using the head‐body elongation ratio. We also examined the morphological changes that may underlie the evolution toward more extreme body plans. We found that a mustelid clade comprised of the subfamilies Helictidinae, Guloninae, Ictonychinae, Mustelinae, and Lutrinae exhibited an evolutionary transition toward more elongate bodies. Furthermore, we discovered that elongation of the body is associated with the evolution of other key traits such as a reduction in body size and a reduction in forelimb length but not hindlimb length. This relationship between body elongation and forelimb length has not previously been quantitatively established for mammals but is consistent with trends exhibited by ectothermic vertebrates and suggests a common pattern of trait covariance associated with body shape evolution. This study provides the framework for documenting body shapes across a wider range of mammalian clades to better understand the morphological changes influencing shape disparity across all vertebrates.     

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.     

Limb segment contributions to the evolution of hind limb length in phrynosomatid lizards

Longer hind limbs are often associated with faster maximum sprint speeds measured in the laboratory and sometimes with increased Darwinian fitness in studies of individual variation in natural populations. Limb length may be altered by changing the length of one or all segments, with different functional consequences. Segment length evolution can be influenced by both natural and sexual selection, and lineage‐specific effects (multiple solutions) may also occur. We examined the evolution of total hind limb length, as well as thigh, crus, pes, and toe length, among 46 species of phrynosomatids and also investigated the role of habitat use and shared evolutionary history in shaping limb morphology. Because sexes are usually behaviourally and morphologically dimorphic, we examined them separately. In females, habitat was only an important predictor of crus (lower leg) length. In males, habitat was not an important predictor of any variable. Overall, clade‐level differences were more important than habitat as predictors of segment or total hind limb length. Not all limb segments scaled isometrically with the combined length of other segments, and both sex and clade affected the scaling of some segments. These results suggest that clade‐level differences are more important than habitat use for explaining differences in limb length and proportions, and sexual dimorphism may be an important consideration in morphology–performance–behaviour–fitness relationships.     

Are there general laws for digit evolution in squamates? The loss and re‐evolution of digits in a clade of fossorial lizards (Brachymeles,Scincinae)

Evolutionary simplification of autopodial structures is a major theme in studies of body‐form evolution. Previous studies on amniotes have supported Morse's law, that is, that the first digit reduced is Digit I, followed by Digit V. Furthermore, the question of reversibility for evolutionary digit loss and its implications for “Dollo's law” remains controversial. Here, we provide an analysis of limb and digit evolution for the skink genus Brachymeles. Employing phylogenetic, morphological, osteological, and myological data, we (a) test the hypothesis that digits have re‐evolved, (b) describe patterns of morphological evolution, and (c) investigate whether patterns of digit loss are generalizable across taxa. We found strong statistical support for digit, but not limb re‐evolution. The feet of pentadactyl species of Brachymeles are very similar to those of outgroup species, while the hands of these lineages are modified (2‐3‐3‐3‐2) and a have a reduced set of intrinsic hand muscles. Digit number variation suggests a more labile Digit V than Digit I, contrary to Morse's law. The observed pattern of digit variation is different from that of other scincid lizards (Lerista, Hemiergis, Carlia). Our results present the first evidence of clade‐specific modes of digit reduction.     

Understanding hind limb weight support in chimpanzees with implications for the evolution of primate locomotion

Most quadrupedal mammals support a larger amount of body weight on their forelimbs compared with their hind limbs during locomotion, whereas most primates support more of their body weight on their hind limbs. Increased hind limb weight support is generally interpreted as an adaptation that reduces stress on primates' highly mobile forelimb joints. Thus, increased hind limb weight support was likely vital for the evolution of primate arboreality. Despite its evolutionary importance, the mechanism used by primates to achieve this important kinetic pattern remains unclear. Here, we examine weight support patterns in a sample of chimpanzees (Pan troglodytes) to test the hypothesis that limb position, combined with whole body center of mass position (COM), explains increased hind limb weight support in this taxon. Chimpanzees have a COM midway between their shoulders and hips and walk with a relatively protracted hind limb and a relatively vertical forelimb, averaged over a step. Thus, the limb kinematics of chimpanzees brings their feet closer to the COM than their hands, generating greater hind limb weight support. Comparative data suggest that these same factors likely explain weight support patterns for a broader sample of primates. It remains unclear whether primates use these limb kinematics to increase hind limb weight support, or whether they are byproducts of other gait characteristics. The latter hypothesis raises the intriguing possibility that primate weight support patterns actually evolved as byproducts of other traits, or spandrels, rather than as adaptations to increase forelimb mobility. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

ALTERNATE PATHWAYS OF BODY SHAPE EVOLUTION TRANSLATE INTO COMMON PATTERNS OF LOCOMOTOR EVOLUTION IN TWO CLADES OF LIZARDS

Body shape has a fundamental impact on organismal function, but it is unknown how functional morphology and locomotor performance and kinematics relate across a diverse array of body shapes. We showed that although patterns of body shape evolution differed considerably between lizards of the Phrynosomatinae and Lerista, patterns of locomotor evolution coincided between clades. Specifically, we found that the phrynosomatines evolved a stocky phenotype through body widening and limb shortening, whereas Lerista evolved elongation through body lengthening and limb shortening. In both clades, relative limb length played a key role in locomotor evolution and kinematic strategies, with long‐limbed species moving faster and taking longer strides. In Lerista, the body axis also influenced locomotor evolution. Similar patterns of locomotor evolution were likely due to constraints on how the body can move. However, these common patterns of locomotor evolution between the two clades resulted in different kinematic strategies and levels of performance among species because of their morphological differences. Furthermore, we found no evidence that distinct body shapes are adaptations to different substrates, as locomotor kinematics did not change on loose or solid substrates. Our findings illustrate the importance of studying kinematics to understand the mechanisms of locomotor evolution and phenotype‐function relationships.     

Comparison of diaphyseal growth between the Libben population and the Hamann-Todd chimpanzee sample

The differences in limb lengths and proportions between humans and chimpanzees are widely known. Humans have relatively shorter forelimbs and longer hind limbs than chimpanzees. Humans have a longer period of long bone formation than chimpanzees. Recent advances in estimating age-at-death in chimpanzees from their dentition have allowed us to reexamine long bone growth in chimpanzees using their skeletal remains and compare it with similar data for humans. A chronological normalization procedure allowing direct interspecific comparison of long bone growth is presented. The preadult chimpanzee sample (n = 43) is from the Hamann-Todd Osteological Collection from the Cleveland Museum of Natural History. All human specimens (n = 202) are from the late Woodland Libben Population currently housed at Kent State University. Relying on these cross-sectional data, we conclude that both species elongate their femora at similar absolute (length per unit time) but different relative (length relative to normalized dental age) rates. The species differ in the absolute growth rate of the humerus but share a common normalized rate of growth. Forelimb segment proportion differences between species are due to differential elongation rates of the segments. Hind limb diaphyseal proportions are the same in both species, which suggests that changes in segment length are proportional. Therefore, alternative developmental mechanisms exist in these closely related species which can produce changes in limb length. © 1996 Wiley-Liss, Inc.     

HOW LIZARDS TURN INTO SNAKES: A PHYLOGENETIC ANALYSIS OF BODY-FORM EVOLUTION IN ANGUID LIZARDS

Abstract One of the most striking morphological transformations in vertebrate evolution is the transition from a lizardlike body form to an elongate, limbless (snakelike) body form. Despite its dramatic nature, this transition has occurred repeatedly among closely related species (especially in squamate reptiles), making it an excellent system for studying macroevolutionary transformations in body plan. In this paper, we examine the evolution of body form in the lizard family Anguidae, a clade in which multiple independent losses of limbs have occurred. We combine a molecular phylogeny for 27 species, our morphometric data, and phylogenetic comparative methods to provide the first statistical phylogenetic tests of several long‐standing hypotheses for the evolution of snakelike body form. Our results confirm the hypothesized relationships between body elongation and limb reduction and between limb reduction and digit reduction. However, we find no support for the hypothesized sequence going from body elongation to limb reduction to digit loss, and we show that a burrowing lifestyle is not a necessary correlate of limb loss. We also show that similar degrees of overall body elongation are achieved in two different ways in anguids, that these different modes of elongation are associated with different habitat preferences, and that this dichotomy in body plan and ecology is widespread in limb‐reduced squamates. Finally, a recent developmental study has proposed that the transition from lizardlike to snakelike body form involves changes in the expression domains of midbody Hox genes, changes that would link elongation and limb loss and might cause sudden transformations in body form. Our results reject this developmental model and suggest that this transition involves gradual changes occurring over relatively long time scales.     

Mechanics of the limbs of the walrus (Odobenus rosmarus) and the california sea lion (Zalophus californianus)

Moment arms and cross-sectional areas of muscles from the forelimbs and hind limbs of Odobenus and Zalophus were used to estimate relative torques associated with several locomotor movements. Odobenus uses predominantly the hind limbs for aquatic propulsion, while Zalophus only uses the forelimbs. Torques contributing to locomotor movements used during the power and recovery strokes were substantially greater for the hind limbs of Odobenus and for the forelimbs of Zalophus. Apparently, the lineages of these two genera shared an aquatic ancestor during the Miocene; therefore, they also shared a common method of aquatic propulsion. This common method of aquatic propulsion could have been either forelimb, hind limb, or both. Since the Miocene then, the method of aquatic propulsion and the mechanical characteristics of the limbs have diverged for at least one of these two lineages. A model is advanced to explain key mechanical differences between Odobenus and Zalophus that enable Odobenus to maneuver effectively in a head-down posture on the ocean floor.     

新疆敏麻蜥3个地理种群趾长及趾长比的两性异形

趾长比是指动物不同趾长的比值,常被认为是早期雄激素或雌激素暴露的标记物之一.本文通过测量新疆敏麻蜥Eremias arguta 3个地理种群的雌雄成体各趾长并计算趾长比,探究不同地理种群趾长及趾长比的两性差异.结果表明,伊犁地区雄性后肢的第2趾显著大于雌性(P<0.05);富蕴县雄性前肢、后肢的第2、3、4、5趾长均显...     

Relationship between innervation and forelimb regenerative capacity in the postmetamorphic pond frog Rana brevipoda porosa.

The limb regenerative capacity and the quantity of innervation (the percentage of a cross-sectional area of amputation forelimb stump occupied by nerves) in the pond frog, Rana brevipoda porosa, was investigated in postmetamorphic froglets and adults of various sizes by means of amputating forelimbs through the zeugopodium. Nearly all the amputated limbs of newly metamorphosed froglets, 18-19 mm in snout-vent length, showed heteromorphic regeneration. However, the larger the body size, the lower the presence of limb regeneration. Limb regenerative capacity was completely lost in froglets and adults with snout-vents larger than 35 mm. The quantity of innervation of limbs was highest in newly metamorphosed froglets, gradually decreasing with growth. The nerve quantity in adults with a snout-vent length between 60-67 mm was approximately half that of the froglets. When the nerve supply was augmented by deviating ipsilateral sciatic nerve bundles to the forelimb stump, almost all limbs, which were usually non-regenerative with normal innervation, regenerated heteromorphically. These results show that the decline in limb regenerative capacity during postmetamorphic growth is in part attributable to the reduction in innervation levels to below the threshold level required for regeneration.     

DIFFERENCES IN THE TIMING OF PRECHONDROGENIC LIMB DEVELOPMENT IN MAMMALS: THE MARSUPIAL–PLACENTAL DICHOTOMY RESOLVED

In contrast to placentals, marsupials are born with forelimbs that are greatly developmentally advanced relative to their hind limbs. Despite significant interest, we still do not know why this is the case, or how this difference is achieved developmentally. Studies of prechondrogenic and chondrogenic limbs have supported the traditional hypothesis that marsupial forelimb development is accelerated in response to the functional requirements of the newborn's crawl to the teat. However, limb ossification studies have concluded that, rather than the forelimb being accelerated, hind limb development is delayed. By increasing the taxonomic coverage and number of prechondrogenic events relative to previous studies, and combining traditional phylogenetic analyses of event sequences with novel analyses of relative developmental rates, this study demonstrates that the timing of limb development in marsupials is more complex than commonly thought. The marsupial phenotype was derived through two independent evolutionary changes in developmental rate: (1) an acceleration of the forelimb's first appearance and (2) a delay of hind limb development from the bud stage onward. Surprisingly, this study also provides some support for an evolutionary acceleration of the marsupial hind limb's first appearance. Further study is needed on the developmental and genetic mechanisms driving these major evolutionary transitions.     

秦岭山区隆肛蛙胚后发育的初步观察

初步观察记录了秦岭山区隆肛蛙Feirana quadrana胚后发育过程.隆肛蛙蝌蚪的头长、体长、尾长和后肢长度随发育时间的增加而显著增长.将隆肛蛙的胚后发育过程划分为20期.其蝌蚪在发育中体色、荚膜(皮肤褶)、侧线以及唇齿式等结构都发生了相应的变化.唇齿式由以II∶ 5-5/II∶ 1-1型为主演变为以II∶ 6-6/II∶ 1-1、I∶ 6-6/II∶ 1-1和II∶ 5-5/II∶ 1-1型为主.     

Variation of Reproductive Traits and Female Body Size in the Most Widely-Ranging Terrestrial Reptile: Testing the Effects of Reproductive Mode, Lineage, and Climate

The European common lizard, Zootoca vivipara, is the most widespread terrestrial reptile in the world. It occupies almost the entire Northern Eurasia and includes four viviparous and two oviparous lineages. We analysed how female snout-vent length (SVL), clutch size (CS), hatchling mass (HM), and relative clutch mass (RCM) is associated with the reproductive mode and climate throughout the species range and across the evolutionary lineages within Z. vivipara. The studied variables were scored for 1,280 females and over 3,000 hatchlings from 44 geographically distinct study samples. Across the species range, SVL of reproductive females tends to decrease in less continental climates, whereas CS corrected for female SVL and RCM tend to decrease in climates with cool summer. Both relationships are likely to indicate direct phenotypic responses to climate. For viviparous lineages, the pattern of co-variation between female SVL, CS and HM among populations is similar to that between individual females within populations. Consistent with the hypothesis that female reproductive output is constrained by her body volume, the oviparous clade with shortest retention of eggs in utero showed highest HM, the oviparous clade with longer egg retention showed lower HM, and clades with the longest egg retention (viviparous forms) had lowest HM. Viviparous populations exhibited distinctly lower HM than the other European lacertids of similar female SVL, many of them also displaying unusually high RCM. This pattern is consistent with Winkler and Wallin’s model predicting a negative evolutionary link between the total reproductive investment and allocation to individual offspring.     

Intraspecific ecomorphological variation: linear and geometric morphometrics reveal habitat‐related patterns within Podarcis bocagei wall lizards

Morphological variation in relation to habitat is known to occur in several lizard groups. Comparative studies have linked morphology and habitat use, showing that locomotion is the principal mediator of this evolutionary relationship. Here, we investigate intraspecific ecomorphological variation in Podarcis bocagei by examining three habitat types, representing a variety between saxicolous and ground‐dwelling habits. Our results indicate variation in absolute and relative limb length, but patterns are only partially concordant to biomechanical predictions. Whereas the femur and hind foot are longer in ground‐dwelling lizards, confirming previous observations, the tibia and hind limb are relatively shorter, contradicting expectations. Additionally, head shape varies substantially between habitats, in line with a hypothesis of mechanical restrictions related to microhabitat and refuge use. Finally, we detect male‐specific variation between habitats in total body size and head size, providing evidence for interactions between natural and sexual selection. Although performance and behaviour studies are necessary to definitely confirm the functional and evolutionary significance of the observed patterns, our study indicates that ecomorphological adaptations can arise in a very short evolutionary time in this group of lizards.