Running in cold weather: morphology, thermal biology, and performance in the southernmost lizard clade in the world (Liolaemus lineomaculatus section: Liolaemini: Iguania)

The integration or coadaptation of morphological, physiological, and behavioral traits is represented by whole-organism performance traits such as locomotion or bite force. Additionally, maximum sprint speed is a good indicator of whole-organism performance capacity as variation in sprinting ability can affect survival. We studied thermal biology, morphology, and locomotor performance in a clade of Liolaemus lizards that occurs in the Patagonian steppe and plateaus, a type of habitat characterized by its harsh cold climate. Liolaemus of the lineomaculatus section display a complex mixture of conservative and flexible traits. The phylogenetically informed analyses of these ten Liolaemus species show little coevolution of their thermal traits (only preferred and optimum temperatures were correlated). With regard to performance, maximum speed was positively correlated with optimum temperature. Body size and morphology influenced locomotor performance. Hindlimbs are key for maximal speed, but forelimb length was a better predictor for sustained speed (i.e. average speed over a total distance of 1.2?m). Finally, sustained speed differed among species with different diets, with herbivores running on average faster over a long distance than omnivores.     

SPEED AND STAMINA TRADE-OFF IN LACERTID LIZARDS

Abstract.— Morphological and physiological considerations suggest that sprinting ability and endurance capacity put conflicting demands on the design of an animal's locomotor apparatus and therefore cannot be maximized simultaneously. To test this hypothesis, we correlated size‐corrected maximal sprint speed and stamina of 12 species of lacertid lizards. Phylogenetically independent contrasts of sprint speed and stamina showed a significant negative relationship, giving support to the idea of an evolutionary trade‐off between the two performance measures. To test the hypothesis that the trade‐off is mediated by a conflict in morphological requirements, we correlated both performance traits with snout‐vent length, size‐corrected estimates of body mass and limb length, and relative hindlimb length (the residuals of the relationship between hind‐ and forelimb length). Fast‐running species had hindlimbs that were long compared to their forelimbs. None of the other size or shape variables showed a significant relationship with speed or endurance. We conclude that the evolution of sprint capacity may be constrained by the need for endurance capacity and vice versa, but the design conflict underlying this trade‐off has yet to be identified.     

黑眶蟾蜍的两性异形与选型配对模式

本研究以黑眶蟾蜍(Duttaphrynus melanostictus)为研究对象,通过对比黑眶蟾蜍抱对个体的体长、头长、头宽、眼间距、鼓膜径、耳后腺长、眼径、前臂及手长、前肢长以及后肢长等形态特征,分析雌性黑眶蟾蜍繁殖输出与其体型的关系,探究黑眶蟾蜍两性异形模式及其与雌性生育力的关系;同时通过对配对个体形态学特征的相关性分析探究了黑眶蟾蜍的配对模式。结果表明,黑眶蟾蜍雌性体长和体重显著大于雄体;两性的所有局部形态特征均与体长成正相关;去除体长因素影响后,雄性头长以及后肢长均明显大于雌性,其余局部形态特征两性间则皆无显著差异。雌体的窝卵重、窝卵数均与其体长和体重成正相关关系。雌性成体的前肢长与抱对雄性成体的前肢长之间呈显著正相关,其余形态特征两性间均无相关性。研究表明,生育力选择是导致黑眶蟾蜍两性异形的重要驱动力;黑眶蟾蜍的选型配对模式未表现在个体大小上,而是体现在局部特征(前肢长),这不仅为揭示两栖类配对模式的普遍性提供了参考,还表明对两栖类选型配对的研究应以多个性状为对象。     

Potential ‘costs of reproduction’ in a skink: Inter‐ and intrapopulational variation

Reproductive costs are important determinants of reproductive effort in squamate reptiles. Consequently, differences in costs of reproduction between populations of geographically or climatically widespread species are likely to result in different patterns of reproductive effort. In the present study, the effect of pregnancy on sprint speed was examined in a small viviparous skink, Niveoscincus ocellatus (Gray 1845), from two populations at the climatic extremes of its distribution. Decreased sprint speed has the potential to be an important cost of reproduction in this species, through a reduced ability to avoid predation and/or decreased foraging efficiency. Lizards inhabiting the colder site were larger than those from the warmer site and, contrary to predictions from life history theory, had a higher reproductive effort. In both populations, sprint speed was lower in pregnant lizards than in either the same individuals after birth or non‐pregnant control lizards. Within each population, sprint speed was unrelated to the level of reproductive effort of the female in terms of either absolute mass of the reproductive burden or the burden relative to her post‐partum body mass. However, within each population, the mass of the clutch that an individual female was carrying relative to snout–vent length was an important determinant of her sprint speed while pregnant. Thus, within each population, a relatively high reproductive burden may potentially increase costs of reproduction in this species. Despite this relationship and predictions from life history theory suggesting that annual reproductive effort will be lower in populations with a large body size and delayed maturity, it is suggested that a higher reproductive effort at the cold site is possible because they have a higher absolute sprint speed because of their larger size and a relatively higher abundance of cover at the cold site, and differences in predation pressure may alter selective pressures on reproductive investment.     

Faster lizards sire more offspring: sexual selection on whole-animal performance

Sexual selection operates by acting on variation in mating success. However, since selection acts on whole-organism manifestations (i.e., performance) of underlying morphological traits, tests for phenotypic effects of sexual selection should consider whole-animal performance as a substrate for sexual selection. Previous studies have revealed positive relationships between performance and survival, that is, natural selection, but none have explicitly tested whether performance may influence reproductive success (through more matings), that is, sexual selection. Performance predicts dominance in some species, implying the effects of sexual selection, but how it does so has not been established, nor is it certain whether performance might be a by-product of selection for something else, for example, elevated circulating testosterone levels. We investigated the potential for sexual selection on sprint speed performance in collared lizards (Crotaphytus collaris), considering the potential mediating effects of circulating hormone levels. Among territorial, adult male collared lizards, only sprint speed significantly predicted territory area and number of offspring sired as determined by genetic paternity analysis. Body size, head size, and hind limb length had no effect. Neither plasma testosterone levels nor corticosterone levels correlated with sprint speed, territory area, or number of offspring sired. Thus, our results provide a direct link between whole-animal performance and reproductive success, suggesting that intrasexual selection can act directly on sprint speed performance and drive the evolution of underlying morphological traits.     

青海沙蜥的两性异型和雌性繁殖

作者研究了青海沙蜥(Phrynocephalus vlangalii)形态特征的两性异形和雌体繁殖特征。蜥蜴于2005年5月初捕自西宁以西约150km的倒淌河,被检形态特征包括体色、体长、腹长、尾长、头长和头宽,新排卵雌体维持在实验室梯度热环境中直至产仔。成体两性异形显著,而性未成熟个体缺乏两性异形。最大的成年雄体和雌体分别为70.2mmSVL(snout-vent length)和82.8mmSVL。雄性成体具有相对较大的头长、头宽和尾长,雌性成体SVL大于雄体且具有相对较大的腹长。对4个形态特征进行主成分分析(特征值≥0.5)区分出2个主成分,共解释83.9%的两性相关形态特征的变异。去除SVL差异的影响后,尾长、头长和头宽在第一主成分有较高的正负载系数(解释57.8%的变异),腹长在第二主成分有较高的负负载系数(解释26.1%的变异)。实验室梯度热环境下的雌体于6月下旬至7月中旬产单窝、2-6个后代。窝仔数和窝仔重与母体SVL呈正相关,幼仔重与母体SVL无关。未在青海沙蜥中检测到后代数量与大小之间的权衡。     

Functional consequences of morphological differentiation between populations of the Cape Dwarf Chameleon (Bradypodion pumilum)

Variation in phenotype between species or populations of the same species living in different habitats is often explained in an adaptive context with local habitat differences driving selection on morphological traits relevant in a given ecological context. Previous studies have demonstrated significant differences in limb and tail morphology between populations of the Cape Dwarf Chameleon (Bradypodion pumilum) living in closed vs. open habitats. However, the adaptive nature of the observed differences remains unclear. Here, we quantify the structural habitat use in two different populations, test whether the random habitat differs between the two sites and whether or not chameleons select perches randomly. Next, we test whether morphology is correlated with structural habitat use and test for differences in performance between populations. Our results demonstrate that habitats are structurally different, that chameleons in the two populations use perches of different diameters and that, in one of the populations, chameleons select relatively wider perches than available at random. Performance traits (hand and tail grip performance and sprint speed) are correlated with morphology (hand size, tail length and tibia length) and differ between sexes and populations. Moreover, performance is dependent on dowel size. These results suggest that differences in performance between populations are indeed adaptive and indicate the existence of true ecomorphs in chameleons of the genus Bradypodion. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 692–700.     

SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

Haematocrit correlates with tail ornament size in three populations of the Barn Swallow (Hirundo rustica)

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity.
2. The change of haematocrit of Barn Swallows ( Hirundo rustica ) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed.
3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males.
4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length.
5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males.     

Female-biased natal and breeding dispersal in an alpine lizard, Niveoscincus microlepidotus

We measured two aspects of dispersal in the alpine Australian scincid lizard, Niveoscincus micolepidotus : (1) natal dispersal, i.e. shift in home range over the lizard's first year of life, and (2) breeding dispersal, i.e. shifts of home ranges between breeding attempts as adults. On average, displacements were surprisingly small. Female neonates dispersed about twice as far as did males in the same cohort (means of 12 m vs. 6 m). A female's natal dispersal distance was not correlated with her body size or our estimate of physiological performance (sprint speed). However, larger, faster-running male neonates dispersed further than did smaller, slower males. As was the case for neonates, adult females moved significantly further between breeding seasons than did adult males (14.2 m vs. 9.6 m). Because of a female's long gestation period (more than 1 year), two groups of females occur simultaneously in the population, non-ovulated (i.e. with yolking folicles) and pregnant females (i.e. approaching parturition). Females that were not yet ovulated showed a markedly stronger dispersal in response to high reproductive effort (i.e. clutch size in relation to body condition) than did pregnant females. In adult males, body size was negatively correlated with dispersal distance, suggesting that although males have overlapping territories, they exhibit an increasing level of site tenacity with age and/or size. Thus, selection for the relatively more pronounced site tenacity in adult males may have resulted in the more marked philopatric behaviour compared to females also as neonates.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 277–283.     

Partial tail loss has no severe effects on energy stores and locomotor performance in a lacertid lizard, <Emphasis Type="Italic">Takydromus septentrionalis</Emphasis>

Many species of lizards use caudal autotomy as a defense strategy to avoid predation, but tail loss entails costs. These topics were studied experimentally in the northern grass lizard, Takydromus septentrionalis. We measured lipids in the three-tail segments removed from each of the 20 experimental lizards (adult females) initially having intact tails to evaluate the effect of tail loss on energy stores; we obtained data on locomotor performance (sprint speed, the maximal length traveled without stopping and the number of stops in the racetrack) for these lizards before and after the tail-removing treatments to evaluate the effect of tail loss on locomotor performance. An independent sample of 20 adult females that retained intact tails was measured for locomotor performance to serve as controls for successive measurements taken for the experimental lizards. The lipids stored in the removed tail was positively correlated with tailbase width when holding the tail length constant, indicating that thicker tails contained more lipids than did thinner tails of the same overall length. Most of the lipids stored in the tail were concentrated in the proximal portion of the tail. Locomotor performance was almost unaffected by tail loss until at least more than 71% of the tail (in length) was lost. Our data show that partial tail loss due to predatory encounters or other factors may not severely affect energy stores and locomotor performance in T. septentrionalis.     

THE EVOLUTION OF FORM AND FUNCTION: MORPHOLOGY AND LOCOMOTOR PERFORMANCE IN WEST INDIAN ANOLIS LIZARDS

I tested biomechanical predictions that morphological proportions (snout–vent length, forelimb length, hindlimb length, tail length, and mass) and maximal sprinting and jumping ability have evolved concordantly among 15 species of Anolis lizards from Jamaica and Puerto Rico. Based on a phylogenetic hypothesis for these species, the ancestor reconstruction and contrast approaches were used to test hypotheses that variables coevolved. Evolutionary change in all morphological and performance variables scales positively with evolution of body size (represented by snout–vent length); size evolution accounts for greater than 50% of the variance in sprinting and jumping evolution. With the effect of the evolution of body size removed, increases in hindlimb length are associated with increases in sprinting and jumping capability. When further variables are removed, evolution in forelimb and tail length exhibits a negative relationship with evolution of both performance measures. The success of the biomechanical predictions indicates that the assumption that evolution in other variables (e.g., muscle mass and composition) did not affect performance evolution is probably correct; evolution of the morphological variables accounts for approximately 80% of the evolutionary change in performance ability. In this case, however, such assumptions are clade-specific; extrapolation to taxa outside the clade is thus unwarranted. The results have implications concerning ecomorphological evolution. The observed relationship between forelimb and tail length and ecology probably is a spurious result of the correlation between these variables and hindlimb length. Further, because the evolution of jumping and sprinting ability are closely linked, the ability to adapt to certain microhabitats may be limited.     

Egg size, offspring quality and food level in planktonic rotifers

1. For two species of planktonic rotifers, Brachionus calyciflorus and Synchaeta pectinata , neonates from larger eggs were more resistant to starvation than neonates from smaller eggs, so egg size was positively correlated with offspring quality.
2. Theories of optimal offspring size in zooplankton predict either a decrease in offspring size with increasing food level, or maximum offspring size at intermediate food levels. In experiments using acclimated, even-aged individuals, neither rotifer species changed its egg size as predicted by theory. B . calyciflorus egg size increased with increasing food concentration, while S . pectinata egg size did not change.
3. The lack of fit to theory may have occurred because of constraints on minimum or maximum egg size, constraints on total reproductive effort, or because offspring fitness is influenced by traits in addition to neonate starvation time.     

SEXUAL SELECTION IN THE BARN SWALLOW HIRUNDO RUSTICA. III. FEMALE TAIL ORNAMENTS

Male secondary sexual characters are often expressed in females, and the maintenance of the character in females can be due to either direct selection on females favoring the maintenance of the trait, or a correlated response to selection in males. Here I report on determinants of and phenotypic selection on tail length of female barn swallows Hirundo rustica. The homologous trait in males is under strong directional sexual selection. Female tail length was positively associated with several reproductive parameters including total seasonal reproductive success, even when controlling for year and age effects. A change in female tail length from one year to another was often associated with a change in the reproductive parameters correlated with absolute tail length. There was little evidence for an association between female tail length and the duration of the incubation period (only females incubate) and absolute and relative female provisioning rates of offspring, and subsequent size of offspring. Tail length of female barn swallows was positively correlated with that of their mates. Female tail length was a heritable trait as determined from regression of the tail trait of offspring on that of their mother and their father, and there was a positive genetic correlation between the tail trait in males and females. In conclusion, female tail length reliably reflects female reproductive potential, and the trait appears to be under directional selection, which may result in an evolutionary response to selection because of the heritability of the tail trait.     

Locomotor performance of hatchling fence lizards (Sceloporus occidentalis): Quantitative genetics and morphometric correlates

Summary We examined heritabilities and correlations among measures of locomotor performance (speed, stamina) and among possible morphometric determinants of performance (hindlimb span, tail length) in families of hatchling lizards (Sceloporus occidentalis). We were particularly interested in determining whether these traits were heritable and thus might potentially respond genetically to selection. Moreover, we wished to determine whether speed and stamina are negatively genetically correlated, as suggested bya priori physiological and empirical considerations. All four traits appeared to be significantly heritable. Broadsense heritabilities were 0.33–0.36 for speed, 0.35–0.36 for stamina, 0.45–0.51 for hindlimb span, and 0.46–0.47 for tail length. Contrary to expectations, speed and stamina were not negatively genetically correlated. Hindlimb span and tail length, however, were negatively genetically correlated (but not phenotypically correlated). Hindlimb span and stamina were positively phenotypically correlated. Thus, for example, selection for longer hindlimb span could potentially result in shorter tails, contrary to evolutionary predictions based only on phenotypic correlations.     

Patterns of caudal-autotomy evolution in lizards

Using comparative techniques to account for phylogenetic effects, I examined patterns of evolution of caudal autotomy and foraging in 39 lizard species to test the hypothesis that caudal autotomy has co-evolved with morphology, locomotor performance, and foraging behaviour. There were significant positive associations between evolution of the point on the tail (distance from cloaca) at which tail loss occurs (an indirect measure of caudal autotomy) and evolution of each of the following: tail length, caudifemoralis longus (CFL) muscle length, and jump distance. The correlation with the evolution of sprint speed approached significance. These relationships primarily were due to the influence of tail-length evolution on autotomy-point evolution. With the effect of tail-length evolution removed, autotomy-point evolution was negatively correlated with the evolution of tail-loss frequency. The CFL restricts tail loss to portions of the tail posterior to the most distal point of its insertion in the tail. In addition, with the effect of tail-length evolution removed, CFL length co-evolved with sprint speed. These results indicate that tail morphology has co-evolved with caudal autotomy such that the evolution of the CFL has reduced caudal autotomy in certain groups of lizards.
Ambush foraging, the ability to lose the tail, intermediate CFL length, and low locomotor performance (i.e. slow sprint speed and short jump distance) are hypothesized to be the ancestral conditions in lizards using outgroup rooting. The diversification of lizard taxa has resulted in some lineages moving away from ancestral character states (i.e. family Teiidae, superfamily Varanoidea), while others are very similar or identical to their ancestors (i.e. superfamily Iguania).     

X chromosome influences sperm length in the stalk-eyed fly Cyrtodiopsis dalmanni

Whether sexually selected traits are sex linked can have profound effects on their evolution. In the diopsid stalk-eyed fly, Cyrtodiopsis dalmanni, sperm length and female reproductive tract morphology have coevolved across species, postcopulatory sexual selection is known to occur, and X-linked genes affect female ventral sperm receptacle size. Here, we estimate the location of quantitative trait loci (QTL) for spermatocyst tail length by using F2 progeny segregating for an X-linked factor that causes sex-ratio meiotic drive. We found two QTL, including a major X-linked QTL responsible for 25% of the variation in spermatocyst tail length 2.1 cM from the sex-ratio element and 0.8 cM from a major eye span QTL. Sex-ratio males produce shorter spermatocyst tails and shorter eye spans. Thus, X-linked factors affect both pre- and postcopulatory traits, and linkage between the alleles for short eye span and short spermatocyst tail length allow pre- and postcopulatory sexual selection to potentially act in concert against the transmission bias caused by the sex-ratio chromosome.     

Evolution of viviparity in cold-climate lizards: testing the maternal manipulation hypothesis

We used a cold-climate viviparous lacertid lizard (Eremias prezwalskii) as a model system to test the maternal manipulation hypothesis. Twenty-four gravid females collected from a population in Inner Mongolia (northern China) were maintained in the laboratory for the whole gestation length, with 12 allowed to bask for 14-h daily and the other 12 for 10-h daily. Females selected lower body temperatures but did not thermoregulate more precisely when gravid. The mean gestation length was shorter in females provided with longer basking opportunity. Neonates in the two treatments differed in tail length and the number of ventral scales but not in other examined morphological traits, with offspring born in the 14-h treatment having longer tails but fewer ventral scales. Offspring were sexually dimorphic at birth, with females being smaller in tail length, head length and fore-limb length but having more ventral scales than males of the same size. Offspring born in the 14-h treatment were not only faster runners but also grew faster than did offspring born in the 10-h treatment. Our data validate the main predictions of the maternal manipulation hypothesis that females should shift selected body temperatures during gestation to provide optimal thermal conditions for developing embryos and that phenotypic traits determined by maternal thermoregulation should enhance offspring fitness. Our study is the first to demonstrate that the maternal manipulation hypothesis applies to cold-climate viviparous reptiles.     

The function of long tails in female barn swallows (Hirundo rustica): an experimental study

The outermost tail feathers in male barn swallows (Hirundo rustica)are the target of a strong directional female mate preference.The tail ornament is also expressed in females, since femaleshave considerably longer tails than juveniles, either due to(1) a strong genetic correlation between the characters in thetwo sexes, or (2) direct sexual selection on females. To discriminatebetween these two hypotheses, we manipulated the length of theoutermost tail feathers in female barn swallows shortly afterarrival by either shortening or elongating the outermost tailfeathers, or maintaining their length among control individuals.Start of laying of the first clutch, reproductive performance,or provisioning of offspring did not show any significant differencesamong treatments. Original female tail length before manipulationwas unrelated to reproductive performance, while male tail lengthexplained some variation in the number of clutches and, to someextent, the total number of eggs laid per year. Females withlonger tails arrived earlier at the breeding grounds. Manipulatedfemale tail length was positively correlated to the tail lengthof their mates. Our results support the correlated responsehypothesis but do not support the sexual selection explanationfor the existence of exaggerated tail feathers in female barnswallows.     

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.