Bieke Vanhooydonck

Performance capacity, fighting tactics and the evolution of life-stage male morphs in the green anole lizard (**Anolis carolinensis**)

The evolution of alternative male phenotypes is probably driven by male–male competition for access to reproductive females, but few studies have examined whether whole–organism performance capacities differ between male morphs, and if so whether any such differences affect fighting ability. We show how ontogenetic changes in performance and morphology have given rise to two distinct life–stage male morphs exhibiting different fighting tactics within the green anole lizard (Anolis carolinensis). Field studies show a bimodal distribution of adult males within a single population: larger ‘heavyweight’ males have relatively large heads and high bite forces for their size, whereas smaller ‘lightweight’ males have smaller heads and lower bite forces. In staged fights between size–matched heavyweight males, males with greater biting ability won more frequently, whereas in lightweight fights, males with greater jumping velocity and acceleration won more often. Because growth in reptiles is indeterminate, and the anole males examined are sexually mature, we propose that the heavyweight morph arose through selection against males with small heads and poor bite forces at the lightweight–heavyweight size transition. Our findings imply that one may not be able to predict male fighting success (and hence potential mating success) by examining aspects of male ‘quality’ at only one life stage.

Rapid large-scale evolutionary divergence in morphology and performance associated with exploitation of a different dietary resource

Although rapid adaptive changes in morphology on ecological time scales are now well documented in natural populations, the effects of such changes on whole-organism performance capacity and the consequences on ecological dynamics at the population level are often unclear. Here we show how lizards have rapidly evolved differences in head morphology, bite strength, and digestive tract structure after experimental introduction into a novel environment. Despite the short time scale (≈36 years) since this introduction, these changes in morphology and performance parallel those typically documented among species and even families of lizards in both the type and extent of their specialization. Moreover, these changes have occurred side-by-side with dramatic changes in population density and social structure, providing a compelling example of how the invasion of a novel habitat can evolutionarily drive multiple aspects of the phenotype.

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 animals 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. Corresponding Editor: T. Garland Jr.

LOCOMOTOR COMPENSATION CREATES A MISMATCH BETWEEN LABORATORY AND FIELD ESTIMATES OF ESCAPE SPEED IN LIZARDS: A CAUTIONARY TALE FOR PERFORMANCE-TO-FITNESS STUDIES

Abstract A key assumption in evolutionary studies of locomotor adaptation is that standard laboratory measures of performance accurately reflect what animals do under natural circumstances. One widely examined measure of performance is maximum sprint speed, which is believed to be important for eluding predators, capturing prey, and defending territories. Previous studies linking maximum sprint speed to fitness have focused on laboratory measurements, and we suggest that such analyses may be appropriate for some species and intraspecific classes, but not others. We provide evidence for a general inverse relationship between maximum laboratory sprint speed and the percentage of maximum capacity that animals use when escaping from a threat in the field (the model of locomotor compensation). Further, absolute values of field escape speed and maximum laboratory speed are not significantly related when comparing across a diverse group of Anolis and lacertid lizards. We show that this pattern of locomotor compensation holds both within (i.e., among intraspecific classes) and among lizard species (with some exceptions). We propose a simple method of plotting field escape speed (y‐axis) versus maximum laboratory speed (x‐axis) among species and/or intraspecific classes that allows researchers to determine whether their study organisms are good candidates for relating laboratory performance to fitness. We suggest that species that reside directly on, or near the “best fitness line”(field escape speed = maximum laboratory speed) are most likely to bear fruit for such studies.

Evolutionary trade-offs in locomotor capacities in lacertid lizards: are splendid sprinters clumsy climbers?

We tested the hypothesis that an evolutionary trade‐off exists between the capacity to run on level terrain and the ability to climb inclined structures in lacertid lizards. Biomechanical and physiological models of lizard locomotor performance suggest that the morphological design requirements of a ground‐dwelling vs. scansorial life style are difficult to reconcile. This conflict is thought to preclude simultaneous evolution of maximal locomotor performance on level and inclined terrain. This notion has been corroborated by comparative studies on lizard species from other groups (Anolis, Chamaeleo, Sceloporus), but is not supported by our data on 13 species from the family Lacertidae. We found no indication of a negative association between maximal sprint speed of lizards over a level racetrack (indicative of ground‐dwelling locomotor performance), on an inclined stony surface (indicative of climbing performance over rock faces) and inclined mesh surface (indicative of clambering performance among vegetation). Moreover, morphological characteristics associated with fast sprinting capacities (e.g. long hind limbs) apparently enhance, rather than hinder climbing and clambering performance. We conclude that in our sample of lacertid lizards, the evolution of fast sprinting capacity on level terrain has not inflicted major restrictions on climbing and clambering performance.

Omnivory in lacertid lizards: adaptive evolution or constraint?

Feeding specializations such as herbivory are an often cited example of convergent and adaptive evolution. However, some groups such as lizards appear constrained in the evolution of morphological specializations associated with specialized diets. Here we examine whether the inclusion of plant matter into the diet of omnivorous lacertid lizards has resulted in morphological specializations and whether these specializations reflect biomechanical compromises as expected if omnivores are constrained by functional trade‐offs. We examined external head shape, skull shape, tooth structure, intestinal tract length and bite performance as previous studies have suggested correlations between the inclusion of plants into the diet and these traits. Our data show that omnivorous lacertid lizards possess modifications of these traits that allow them to successfully exploit plant material as a food source. Conversely, few indications of a compromise phenotype could be detected, suggesting that the evolution towards herbivory is only mildly constrained by functional trade‐offs.

A biomechanical analysis of intra- and interspecific scaling of jumping and morphology in Caribbean Anolis lizards

SUMMARY Scaling models predict how functional variables change as animals grow or increase in size evolutionarily. However, few experimental studies have found support for the predictions of these models. Here, we use a force plate to investigate the scaling of functional variables associated with jumping within (for three species) and across adults of 12 species of Anolis lizards. Both ontogenetically (with the exception of Anolis carolinensis) and across the 12 species examined, limb dimensions increased geometrically, making Anolis lizards an ideal study system to test the predictions of geometric scaling models. However, both the ontogenetic and interspecific scaling of functional variables deviated in several aspects from model predictions. Unexpectedly, the scaling of functional variables such as acceleration differed for different species. Whereas acceleration capacity increases with hindlimb length for A. carolinensis, no relationship was detected for the other two species. Interspecifically, the inclusion of two large species in our analysis appears to drive the absence of a correlation between acceleration capacity and hindlimb length across species. These data suggest that selection for enhanced jumping performance is relaxed in larger anoles and support the notion that no scaling model seems to be able to comprehensively predict changes in function with size across species; rather, natural selection seems to drive changes in the scaling relationships of some key variables such as force output or acceleration capacity.

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

Abstract 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.

Morphology, performance, behavior and ecology of three color morphs in males of the lizard Podarcis melisellensis

Males of a Croatian population of the lacertid lizard Podarcis melisellensis exhibit a striking polymorphism, with coloration of the throat and abdomen ranging from completely white, to yellow or orange. In a first attempt to explore the potential ecological and evolutionary significance of this polymorphism, we compared the three forms of males in aspects of their morphology, whole-animal performance, behavior, and ecology. Orange males are, on average, larger in snout-vent length and have disproportionately larger heads than either white or yellow males. This is reflected in orange males having higher bite force capacity and theoretically an increased access to harder prey. Residual limb length, maximal sprint speed and maximal exertion do not differ among color morphs. Body temperatures in the field are similar in the three morphs, but yellow males are caught at sites with slightly higher air temperatures than are orange and white males. Behavioral observations show no differences in time budgets or in the timing of activities among morphs. Microhabitat use is also similar in the three color morphs, but orange males were more often initially seen on rocky substrates. Our findings suggest that the observed polymorphism likely does not originate from a divergence in niche or use of resources, but possibly reflects an underlying polymorphism in mating tactics.

Correlations between habitat use and body shape in a phrynosomatid lizard (Urosaurus ornatus): a population-level analysis

Recent ecomorphological studies have shown that the predicted correlations between morphology and ecology on broad taxonomic levels are often obscured when comparing more closely related groups. Among species, comparisons of lizards often indicate very little support for adaptive radiations into novel habitats. As few population level studies have been performed, we compared body, head and limb shape between four populations of Urosaurus ornatus living in structurally distinct habitats (cliffs, rocks, trees and boulders). Surprisingly, clear correlations between habitat use and body shape among populations were found, most of which were in good accordance with a priori biomechanical predictions (e.g. flat body and head for extreme climbers; long distal hindlimb segments for jumpers and runners; narrow body and long tail for tree dwelling lizards). This indicates that populations of Urosaurus ornatus are seemingly ‘adapted’ to the habitat they live in. However, quantification of performance and behaviour are needed to determine the adaptive nature of these observations.  2001 The Linnean Society of London