Anthony Herrel

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.

Octopamine in Male Aggression of Drosophila

BACKGROUND In mammals and humans, noradrenaline is a key modulator of aggression. Octopamine, a closely related biogenic amine, has been proposed to have a similar function in arthropods. However, the effect of octopamine on aggressive behavior is little understood. RESULTS An automated video analysis of aggression in male Drosophila has been developed, rendering aggression accessible to high-throughput studies. The software detects the lunge, a conspicuous behavioral act unique to aggression. In lunging, the aggressor rears up on his hind legs and snaps down on his opponent. By using the software to eliminate confounding effects, we now show that aggression is almost abolished in mutant males lacking octopamine. This suppression is independent of whether tyramine, the precursor of octopamine, is increased or also depleted. Restoring octopamine synthesis in the brain either throughout life or in adulthood leads to a partial rescue of aggression. Finally, neuronal silencing of octopaminergic and tyraminergic neurons almost completely abolishes lunges. CONCLUSIONS Octopamine modulates Drosophila aggression. Genetically depleting the animal of octopamine downregulates lunge frequency without a sizable effect on the lunge motor program. This study provides access to the neuronal circuitry mediating this modulation.

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.

Evolution of bite performance in turtles

Abstract Among vertebrates, there is often a tight correlation between variation in cranial morphology and diet. Yet, the relationships between morphological characteristics and feeding performance are usually only inferred from biomechanical models. Here, we empirically test whether differences in body dimensions are correlated with bite performance and trophic ecology for a large number of turtle species. A comparative phylogenetic analysis indicates that turtles with carnivorous and durophagous diets are capable of biting harder than species with other diets. This pattern is consistent with the hypothesis that an evolutionary increase in bite performance has allowed certain turtles to consume harder or larger prey. Changes in carapace length tend to be associated with proportional changes in linear head dimensions (no shape change). However, maximum bite force tends to change in proportion to length cubed, rather than length squared, implying that changes in body size are associated with changes in the design of the jaw apparatus. After the effect of body size is accounted for in the analysis, only changes in head height are significantly correlated with changes in bite force. Additionally, our data suggest that the ability to bite hard might trade off with the ability to feed on fast agile prey. Rather than being the direct result of conflicting biomechanical or physiological demands for force and speed, this trade‐off may be mediated through the constraints imposed by the need to retract the head into the shell for defensive purposes.

Ecomorphological analysis of trophic niche partitioning in a tropical savannah bat community

The exceptional diversity of neotropical bat communities is sustained by an intricate partitioning of available resources among the member species. Trophical specialization is considered an important evolutionary avenue towards niche partitioning in neotropical phyllostomid bats. From an ancestral insectivorous condition, phyllostomids evolved into highly specialized frugivorous, carnivorous, nectarivorous, piscivorous and even sanguivorous species. Previously, correlations between cranial morphology and trophic ecology within this group have been documented. Here, we examine the evolutionary relationships between bite force and head shape in over 20 species of bats from a single tropical savannah bat community. The results show that bite force increases exponentially with body size across all species examined. Despite the significant differences between large dietary groups using traditional analysis (i.e. non–phylogenetic) and the strong evolutionary correlations between body mass and bite force, phylogenetic analyses indicated no differences in bite performance between insectivorous, omnivorous and frugivorous bats. Comparisons of three species with highly specialized feeding habits (nectarivory, piscivory and sanguivory) with the rest of the species in the community indicate that specialization into these niches comes at the expense of bite performance and, hence, may result in a reduction of the trophic niche breadth.

The effects of gape angle and bite point on bite force in bats.

SUMMARY Models of mammalian mastication predict that bite force is affected by both the degree of mouth opening (gape angle) and the point along the tooth row at which force is transferred to a food item (bite point). Despite the widespread use of these models in comparative analyses, experimental data documenting bite force in non-human mammals are extremely limited. The goal of this study is to document variation in non-stimulated bite force associated with change in gape angle and bite point in a broad range of species. We focus on plant-visiting bats because they exhibit a relatively primitive cranial morphology and are good models for generalized mammals. Assessments of the relationship between gape angle and bite force within and among species demonstrate that bite force decreases significantly as gape angle increases. The relationship between bite force and bite point within each of seven species demonstrates that unilateral molar biting universally generates the highest forces while the unilateral canine biting produces the lowest forces. Bilateral canine biting is intermediate. Beyond these general patterns, differences among species suggest that bite force reflects variation in craniofacial architecture. Finally, these data suggest that behavioral variation in gape angle and bite point may be important variables in comparative, functional analyses of feeding.

Ontogeny of Performance in Vertebrates

When competing for food or other resources, or when confronted with predators, young animals may be at a disadvantage relative to adults because of their smaller size. Additionally, the ongoing differentiation and growth of tissues and the development of sensory‐motor integration during early ontogeny may constrain performance. Because ectothermic vertebrates show different growth regimes and energetic requirements when compared to endothermic vertebrates, differences in the ontogenetic trajectories of performance traits in these two groups might be expected. However, both groups of vertebrates show similar patterns of changes in performance with ontogeny. Evidence for compensation, resulting in relatively high levels of performance in juveniles relative to adults, appears common for traits related to locomotor and defensive behaviors. However, there is little evidence for compensation in traits associated with feeding and foraging. We suggest that this difference may be due to different selective regimes operating on locomotor versus feeding traits. As a result, relatively high levels of locomotor performance in juveniles and relatively high levels of feeding performance in adults are observed across a wide range of vertebrate groups.

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.

Sexual Dimorphism of Head Size in Podarcis Hispanica Atrata: Testing the Dietary Divergence Hypothesis By Bite Force Analysis

Sexual dimorphism of relative head size is a widespread phenomenon in lizards, males having larger head/trunk ratios than females. In an attempt to explain this sexual dimorphism several hypotheses have been formulated. The two most frequently cited ones are: 1) sexual selection acting on those structures important in intrasexual competition and 2) natural selection for reduction of food competition between the sexes. In the insular subspecies of Podarcis hispanica (P h. atrata) males tend to have significantly larger heads than similarly sized females. We here test an implicit assumption of the dietary divergence hypothesis, namely that an increase in head size results in an increase in gape width and/or bite force, thereby allowing the larger headed sex to exploit larger prey classes. Using a static bite force model, we calculated the magnitude of bite forces for given directions at given positions on the jaws and for different head sizes. We experimentally determined the hardness of three different prey items and compared the data to the maximal bite force produced by both sexes. Our results suggest an important difference in male and female bite capacity, which may bear significant ecological relevance, and are in agreement with the implicit assumption of the dietary divergence theory.

Ontogenetic Scaling of Bite Force in Lizards and Turtles

Because selection on juvenile life‐history stages is likely strong, disproportionately high levels of performance (e.g., sprint speed, endurance, etc.) might be expected. Whereas this phenomenon has been demonstrated with respect to locomotor performance, data for feeding are scarce. Here, we investigate the relationships among body dimensions, head dimensions, and bite force during growth in lizards and turtles. We also investigate whether ontogenetic changes in bite performance are related to changes in diet. Our analyses show that, for turtles, head dimensions generally increase with negative allometry. For lizards, heads scale as expected for geometrically growing systems. Bite force generally increased isometrically with carapace length in turtles but showed significant positive allometry relative to body dimensions in lizards. However, both lizards and turtles display positive allometric scaling of bite force relative to some measures of head size throughout ontogeny, suggesting (1) strong selection for increased relative bite performance with increasing head size and (2) intrinsic changes in the geometry and/or mass of the jaw adductors during growth. Whereas our data generally do not provide strong evidence of compensation for lower absolute levels of performance, they do show strong links among morphology, bite force, and diet during growth.