Condition-dependent spermatophore size is correlated with male's age in a bushcricket (Orthoptera: Phaneropteridae)

During mating, male bushcrickets transfer a spermatophore that consists of a sperm-containing ampulla and a product of the accessory glands, the spermatophylax, which females directly ingest. In the present study, we demonstrate male spermatophore allocation in the bushcricket Poecilimon zimmeri . Males of this species show condition-dependent spermatophore investment. This investment depended upon the age at first mating of males, with older individuals transferring larger spermatophores than younger ones of the same body mass. Independently of age, heavier males transfer larger spermatophores, but the size of males (as measured by femur length) was not a good predictor. Heavier males allocate a lower proportion of their mass to spermatophores and reach their maximal investment point earlier than less heavy males. Spermatophylax production levelled off to a species specific maximum earlier than that of sperm investment (measured as ampulla mass), suggesting that males face high levels of sperm competition.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 354–360.     

Evolution of jumping capacity in Tropidurinae lizards: does habitat complexity influence obstacle‐crossing ability?

Jumping performance is relevant for lizards in many ecological contexts and might be favoured during the colonization of structurally complex habitats. Although ground-dwelling lizards use jumps to overcome small obstacles in their natural environments, jumping capacity has been mostly studied in arboreal species. Here, we analysed the evolution of jumping behaviour and performance in lizards from eight ground-dwelling species of Tropidurinae attempting to cross obstacles of different heights in a jumping track, both when undisturbed and under continuous stimulation. To establish ecological correlates with habitat complexity, individuals from two contrasting Brazilian habitats, the arid Caatingas (sand species) and the savannah-like Cerrados (rock species), were compared. Rock species jumped more often and crossed higher obstacles than sand ones in both tests, and performed more vertical than horizontal jumps. Although sand species performed less jumps, they were more successful at crossing the obstacles presented in comparison with rock species. Phylogenetic analyses confirmed these findings and demonstrated a large divergence in jumping capacity between sister-species from different habitats. Therefore, the differences in propensity and endurance for jumping activity appear to be independent of phylogenetic relationships in Tropidurinae and likely reflect an adaptation to the contrasting environments inhabited. The ecological implications of these findings are discussed.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 393–402.     

Bias in interspecific allometry: examples from morphological scaling in varanid lizards

The traditional approach to allometric analysis entails the fitting of a straight line to logarithmic transformations of the data, after which parameters in a two-parameter allometric equation are estimated by back-transformation to the original scale. We re-examined published data for dimensions of the limbs in 22 species of varanid lizards to illustrate the biases that can be introduced into allometric analyses by applying the aforementioned protocol. Statistical models fit to the original data by linear and nonlinear regression conformed better with underlying assumptions than did models obtained by back-transformation from logarithms, and the former generally were better than the latter for describing limb dimensions over the full range in body size. Allometric exponents estimated by the traditional method therefore were based on inappropriate and inaccurate statistical models and, consequently, were biased and misleading. Investigators can avoid problems such as these by performing preliminary graphical and statistical analyses on data in their original scale and by validating the fitted model. Logarithmic transformations should be used sparingly and only for cause.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 296–305.     

Relations between microhabitat use and limb shape in phrynosomatid lizards

With the exception of the well-documented case for anoline lizards, recent studies have found few evolutionary relationships between morphology and habitat use in lizards despite clear-cut biomechanical predictions. One of the factors typically hampering these analyses is the clustering of habitat use within evolutionary lineages. In the present study, body shape was quantified for male and female lizards of 30 species of phrynosomatid lizards. This group was selected as little clustering of ecological variables seemed to be present. The results of traditional analyses indicate that evolutionary correlates of habitat use were prominent in the hindlimbs of both sexes. Species living in open habitats are characterized by longer femurs, and longer hindlimbs relative to the forelimb. Moreover, males from ground-dwelling species utilizing open habitats have longer toes on the hind foot than males from climbing species. Phylogenetic analyses indicated strong evolutionary associations between habitat use and the relative length of front and hindlimbs, with species from open terrestrial habitats having significantly shorter frontlimbs relative to their hindlimb than rock or tree climbing species. Evolutionary associations between morphology and habitat use were generally stronger for male lizards, indicating a potentially important contribution of sexual selection to the evolution of differences in limb proportions.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 149–163.     

Sexual dimorphism of heads and abdomens: Different approaches to ‘being large’ in female and male lizards

Sexual‐size dimorphism (SSD) is widespread in animals. Body length is the most common trait used in the study of SSD in reptiles. However, body length combines lengths of different body parts, notably heads and abdomens. Focusing on body length ignores possible differential selection pressures on such body parts. We collected the head and abdomen lengths of 610 lizard species (Reptilia: Squamata: Sauria). Across species, males have relatively larger heads, whereas females have relatively larger abdomens. This consistent difference points to body length being an imperfect measure of lizard SSD because it comprises both abdomen and head lengths, which often differ between the sexes. We infer that female lizards of many species are under fecundity selection to increase abdomen size, consequently enhancing their reproductive output (enlarging either clutch or offspring size). In support of this, abdomens of lizards laying large clutches are longer than those of lizards with small clutches. In some analyses, viviparous lizards have longer abdomens than oviparous lizards with similar head lengths. Our data also suggest that male lizards are under sexual selection to increase head size, which is positively related to winning male–male combats and to faster grasping of females. Thus, larger heads could translate into higher probability to mate. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 665–673.     

Ontogenetically stable dimorphism in a lacertid lizard (Acanthodactylus boskianus) with tests of methodology and comments on life-history

Recent arguments in the literature prompted us to compare methods for assessing sexual dimorphism in body proportions of lacertid lizards, using Acanthodactylus boskianus . Although expressing body-part measurements as proportional to head length was the most effective method, we recommend using trunk length for the baseline as a general method for lizards. We also argue that, when aiming to assess sexual dimorphism in body proportions of lizards, if the context is ecological, all available adults should be included. However, for morphology and taxonomy, small sub-samples of the largest individuals that maximally express their genetic morphological potential should be used. In A. boskianus , the sexual dimorphism of mensural characters in adults was typical: males were larger, with relatively larger head and appendages. However, the ontogeny of this dimorphism was unusual in that the differences existed already in youth and thereafter persisted isometrically. The sexual dimorphism of meristic characters was male-biased in numbers of femoral pores and of caudal vertebrae, and female-biased in numbers of ventral plates along the trunk and of precaudal vertebrae. Size dimorphism may conceivably play a role in sex recognition because two potential visual cues (i.e. size dimorphism and dichromatism) appear to complement each other.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 275–288.     

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.     

Size constraints and the evolution of display complexity: why do large lizards have simple displays?

Social, environmental, and perceptual factors have been suggested to account for the evolution of visual signal diversity in lizards. Previous investigations have inferred that signal complexity may also be related to body size. In this study, we use three complementary comparative analyses to investigate whether body size has influenced macro-evolutionary trends in display modifier repertoire size for 110 species of iguanian lizards. We found evidence that signal complexity, as measured by repertoire size, is negatively associated with body size. However, this relationship was not strictly linear. Rather, body size seems to impose a threshold on signal evolution. Specifically, the evolution of large repertoire size appears to be less likely above a particular size threshold, which results in large-bodied lizards having a significantly lower probability of evolving elaborate displays. This relationship may reflect the influence of body size on resource use and the emergent social dynamics it promotes. Large lizards tend to be herbivorous and typically do not defend foraging patches. Consistent with this hypothesis is the previously reported finding of a similar size threshold dividing herbivorous from insectivorous lizards. We suggest to fully understand the evolutionary processes acting on communicative systems, it is important that we identify both the selective forces involved and the nature of their influence. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 145–161.     

Morphological, functional and evolutionary aspects of tail autotomy and regeneration in the 'living fossil'Sphenodon (Reptilia: Rhynchocephalia)

Tail autotomy and regeneration are less known in Sphenodon ('Reptilia': Rhynchocephalia) than in Squamata. We examined museum specimens, Sphenodon guntheri ( N  = 8) and Sphenodon punctatus ( N  = 172), wild Sphenodon punctatus ( N  = 19) and Sphenodon sp. skeletons ( N  = 8). In S. punctatus , unlike in typical Squamata, sexes had similar relative (intact) tail lengths, and regeneration frequencies; tail and body growth was isometric. Tail breakage was usually intravertebral, usually followed by ablation of a variably sized terminal vertebral piece, partly deviating from lizards. Hypothetically, imperfect autotomy results from sphenodon's primitiveness. As in squamates, tail-losers were morphologically more left-side dominant than tail-retainers. Individual directional asymmetries in digit morphology and in digit injury were correlated (in lizards observed only at population level); tail-losers had more fluctuating asymmetry but their exclusion did not facilitate morphological taxonomic distinctions (no 'Seligmann effect'). In S. punctatus , extents and directions of sexual dimorphism paralleled differences between tail-retainers and tail-losers, females resembling tail-losers, also accounting for character interdependence (developmental constraints; employing a method similar to phylogenetic contrasts). The variation in the location of tail injury was correlated with the continuum of variation between injured and intact (pholidotic) morphotypes. These last two phenomena remain to be explored in Squamates.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 721–743.     

Plastic resource polymorphism: effects of resource availability on Arctic char (Salvelinus alpinus) morphology

Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char ( Salvelinus alpinus ) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 341–351.     

Functional basis for sexual differences in bite force in the lizard Anolis carolinensis

In many species of lizards, males attain greater body size and have larger heads than female lizards of the same size. Often, the dimorphism in head size is paralleled by a dimorphism in bite force. However, the underlying functional morphological basis for the dimorphism in bite force remains unclear. Here, we test whether males are larger, and have larger heads and bite forces than females for a given body size in a large sample of Anolis carolinensis . Next, we test if overall head shape differs between the sexes, or if instead specific aspects of skull shape can explain differences in bite force. Our results show that A. carolinensis is indeed dimorphic in body and head size and that males bite harder than females. Geometric morphometric analyses show distinct differences in skull shape between males and females, principally reflecting an enlargement of the jaw adductor muscle chamber. Jaw adductor muscle mass data confirm this result and show that males have larger jaw adductors (but not jaw openers) for a given body and head size. Thus, the observed dimorphism in bite force in A. carolinensis is not merely the result of an increase in head size, but involves distinct morphological changes in skull structure and the associated jaw adductor musculature.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 111–119.     

Ultraviolet reflectance and cryptic sexual dichromatism in the ocellated lizard, Lacerta (Timon) lepida (Squamata: Lacertidae)

Ultraviolet (UV) colorations have garnered extensive theoretical and empirical treatment in recent years, although the majority of studies have concerned themselves with avian taxa. However, many lizards have acute visual systems with retinal photoreceptors that are sensitive to UV wavelengths, and also display UV-reflecting colour patches. In the present study, we used UV photography and full-spectrum reflectance spectrophotometry to describe intra- and intersexual colour variation in adult ocellated lizards Lacerta ( Timon ) lepida and to obtain evidence of UV-based ornamentation. We also investigated whether any colour traits correlate with morphological traits potentially related to individual quality. The results obtained show that the prominent eyespots and blue outer ventral scales (OVS) that ocellated lizards have on their flanks reflect strongly in the UV range and are best described as UV/blue in coloration. The eyespots of males are larger and cover a larger surface area than those of females. However, these differences can be entirely accounted for by sex differences in body size, with males being generally larger than females. We also found differences in the shape of reflectance curves from males and females, with the eyespots and blue OVS of males being more UV-shifted than those of females. Other body regions have extremely low UV reflectance and are not sexually dichromatic. Eyespot size and the total surface area covered by eyespots increases with body size in males but not in females, suggesting that they may be signalling an intrinsic individual characteristic such as body size or male fighting ability. We also discuss the alternative and non-exclusive hypothesis that eyespots may function in lizards of both sexes as protective markings against predators.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 766–780.     

Survival estimation in a long-lived monitor lizard: radio-tracking of Varanus mertensi

The population dynamics of varanids (large monitor lizards) is poorly understood. We report on the most detailed study to date of a population of one of Australia’s largest semi-aquatic varanids, Varanus mertensi. Survival of V. mertensi was derived from known-fate modelling of radio-tracked individuals over two and a half years. We demonstrate empirically what intuition suggests; that apparent survival probability in long-lived lizards is high over short sampling periods, with body size and gender influencing these estimates. Survival estimation in long-lived species such as varanids clearly requires long-term studies.     

Reversible frequency-dependent predation of a puffer, Takifugu niphobles (Pisces: Tetraodontidae), related to spatial distribution of colour-polymorphic prey

The puffer Takifugu niphobles is a top predator of hard-shelled prey such as molluscs; its predatory tactics may affect the evolution of prey coloration. Two hypotheses concerning its foraging were tested: (1) T. niphobles shows frequency dependence in foraging colour-polymorphic prey, and (2) such dependence reverses in response to changes in prey distribution. Captive fish were provided with 70 artificial prey, coloured either dark brown or pale brown, at four frequencies (1 : 4, 2 : 3, 3 : 2, 4 : 1) and in two distribution patterns (uniform and aggregated). When prey were uniformly distributed, frequency and feeding rate significantly influenced colour preference: the common morph was consumed more. When prey were aggregated, frequency significantly affected preference only when the feeding rate was low, in which case the rare morph was consumed more. Thus both hypotheses were supported. The impact of T. niphobles 's frequency-dependent predation and its reversal on the colour evolution of prey species, especially molluscs, is discussed.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 197–202.     

Size matters: the (negative) allometry of copulatory duration in mammals

The amount of time taken to copulate varies enormously among mammals. Because copulation likely exposes animals to an increased risk of predation, and uses time and energy that could be spent on foraging, smaller mammals (which are vulnerable to more predators and have a shorter time-to-starvation than larger mammals) should spend less time copulating than do larger mammals. Furthermore, if extended copulation reflects competition among males, then the duration of mating (after correction for body size) should be greater in mammals in which females mate with more than one male. We tested these predictions using comparative data from 113 mammalian species in 85 genera, 40 families, and 14 orders, while controlling for the effect of phylogeny. We found: (1) the relationship between duration of copulation and body size to be negative, not positive; (2) no relationship with inferred multiple mating by females (based upon relative testes mass). We suggest that small mammals may find the sustained maneuvering and body positioning of copulation easier than do large mammals. This hypothesis is supported by an apparently isometric relationship between duration of copulation and ratio of power to mass.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 185–193.     

Relationships between bone growth rate, body mass and resting metabolic rate in growing amniotes: a phylogenetic approach

We explored the factors that explain the variation in resting metabolic rates (RMR) in growing amniotes by using the phylogenetic comparative method. For this, we measured raw RMR (mL O₂ h⁻¹), body mass, body mass growth rate, and periosteal bone growth rate in a sample of 44 growing individuals belonging to 13 species of amniotes. We performed variation partitioning analyses, which showed that phylogeny explains a significant fraction of the variation of mass-specific RMR (mL O₂ h⁻¹ g⁻¹), and that the cost of growth is much higher than the cost of maintenance. Moreover, we tested the hypothesis of the independence of energy allocation, and found that maintenance metabolism and growth rates are not significantly related. Finally, we calculated the statistical parameters of the relationship between geometry-corrected RMR (mL O₂ h⁻¹ g^−0.67) and bone growth rate. This relationship could potentially be used in palaeobiology to infer RMR from bone tissue samples of fossil species by assuming Amprino's rule (according to which bone tissue types reflect bone growth rates). These estimates would be especially interesting for Mesozoic non-avian theropod dinosaurs and Permian and Triassic therapsids to investigate, respectively, the origin of avian and mammalian endothermy.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 63–76.     

Convergent body flattening in a clade of tropical rock-using lizards (Scincidae: Lygosominae)

The repeated occurrence of similar morphologies in organisms from similar habitats provides good evidence of convergent selection, and convergent patterns of evolutionary change. In lizards, a flattened morphology has often been noted; however, whether this trait is convergent in specific habitats has never been tested using phylogenetic methods. The present study examined patterns of morphological convergence in 18 species of tropical Lygosomine skinks from three broad habitat categories (generalist, leaf litter-dwelling, and rock-using species). In general, although there where relatively few morphological differences of species from different habitats, phylogenetic analyses revealed that rock-using species have consistently and repeatedly evolved a dorsoventrally flattened head and body. The adaptive basis of this flattened morphology is consistent with both biomechanical predictions of performance (e.g. climbing locomotion) and ecology (e.g. use of rock crevices, camouflage) of species that occupy rocky habitats.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 399–411.     

Body size in proboscideans, with notes on elephant metabolism

Mass estimates for a number of fossil proboscideans were computed using regression analyses on appendicular bones to body mass, for seven specimens of modern elephants, for which body masses had been recorded prior to death. The marked differences in physical proportions between extant Loxodonta and Elephas , implying substantial differences in body mass at any given shoulder height, were not present in their long bone parameters. Length and least circumferences proved to be the best parameters for prediction of body mass. Some extinct proboscideans, notably certain Mammuthus and Deinotherium , were much larger than extant elephants. Both the basal and the field metabolic rates of extant elephants are lower than predicted for a hypothetical mammal, in accordance with their body size and subsistence on low-quality foods. The feeding quantities often ascribed to extant wild elephants are exaggerated, and would in fact have sufficed to nourish much larger species.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 523–549.     

Adaptive significance of food income in European snakes: body size is related to prey energetics

Feeding strategies and diet patterns have been extensively investigated in vertebrates and, more specifically, in snakes. Although it has been hypothesized that prey species may differ in terms of energy content, almost no theoretical or practical study has been carried out to determine actual nutritional values of the common prey types of wild snakes. Our model taxa were a selection of widely distributed and well known European snake species, which have all been studied in depth: approximately 76% of their diet is composed of mammals, reptiles, and insects. We therefore selected a single model species for each of these categories and proceeded with the analyses. Nutritional values were determined using a standard procedure: lizards and mice were richer in proteins than insects (crickets); insects and mice were richer in lipids than lizards, and mice and crickets have a higher energy content than lizards; lizards were rich in ashes. We then applied our experimental results to a selected sample of European terrestrial snakes (11 populations, ten species, seven genera, two families) characterized by different body size (50–160 cm total length) and reproductive strategies (oviparous versus viviparous), aiming to correlate these parameters with patterns of energy income. A direct relationship was found between body mass/body length ratio (BCI, body condition index) and meal energetics: the higher the BCI, the higher was the metabolic requirement, whereas BCI was independent of species or of reproductive system effect. Large‐sized snakes thus need a highly diversified and more energy‐rich diet than smaller snakes, supporting previous hypotheses. The simple applicability of this method could be of valuable support in further comparative research work, reducing experimental costs and stimulating further ecological, behavioural, and, possibly, phylogenetic comparisons. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 307–317.     

Physiological variation along a geographical gradient: is growth rate correlated with routine metabolic rate in Rana temporaria tadpoles?

Shorter season length and lower temperature towards higher latitudes and altitudes often select for intraspecific clines in development and growth rates. However, the physiological mechanisms enabling these clines are not well understood. We studied the relationship between routine metabolic rate (RMR) and larval life-history traits along a 1500-km latitudinal gradient across Sweden. In a laboratory common garden experiment, we exposed eight common frog Rana temporaria populations to two experimental temperatures (15 and 18 °C) and measured RMR using flow-through respirometry. We found significant differences among populations in RMR, but there was no evidence for a linear relationship between latitude and RMR in either temperature treatment. However, we found a concave relationship between latitude and RMR at the lower experimental temperature. RMR was not correlated with growth rate at population or at individual levels. The results obtained suggest that, unlike in growth and development rates, there is no latitudinal cline in RMR in R. temporaria tadpoles.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 217–224.