Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Morphological correlates of burst speed and field movement patterns: the behavioural adjustment of locomotion in wall lizards (Podarcis muralis)

Locomotion of lizards has clear morphological determinants and is important for developing activities such as feeding, social interaction and predator avoidance. Thus, morphological variation is believed to have fitness consequences through affecting locomotor performance. This paper firstly evaluates the dependence of burst speed on morphology, and secondly examines the movement patterns of free-ranging undisturbed wall lizards ( Podarcis muralis ) engaged in several kinds of activity. Body size was the most important correlate of burst speed as performed at the optimal temperature for running in the laboratory. After removing size effects from performance and morphological traits, the length of some particular limb segments had positive influence on burst speed, but these effects were weak, each trait explaining less than 16% of variance in burst speed. Free-ranging P. muralis exhibited intermittent locomotion, with movement sequences interrupted by frequent short pauses. Field movement patterns greatly differed depending upon the kind of activity and were in most aspects independent of the size and sex of the animal. P. muralis involved in thermoregulation performed short and low-speed displacements; exploratory activities were characterized by frequent, slow and short movements. On the contrary, lizards involved in intraspecific pursuits and predator escape developed comparatively high speeds, although only exceptionally did they attain the size-specific burst speed predicted from the laboratory trials. Speed of escape increased with distance to the refuge and the animals are able to assess predation risks to modulate approach distance, speed and pauses, so maximum exertion is seldom required. The evolution of locomotor capacities exceeding routine needs is discussed in the context of the principle of 'excessive construction'.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 135–146.     

Costs of reproduction in a tropical invariant-clutch producing lizard (Carlia rubrigularis)

Reproduction involves costs and benefits, whereby increased expenditure into current reproduction can reduce future reproductive success. Costs of reproduction, often assessed using the reduction in locomotor speed, have become well established in temperate-zone lizard species. However, substantial differences in biotic conditions and life-history traits between temperate- and tropical-zone lizards suggest that such costs may be different or of less importance for tropical species. This study examined the effects of reproduction on locomotor speed in the tropical invariant-clutch producing lizard Carlia rubrigularis . Counter to predictions and despite a low relative clutch mass (RCM), gravid and post-oviposition females experienced a reduction in locomotor speed with a physiological basis that was unrelated to the level of reproductive investment. In addition, gravid and postoviposition females exhibited locomotor speeds that were inversely related to the timing of oviposition and approached the speed of non-reproductive females after 3 weeks of oviposition. These results suggest that in addition to RCM, selection may act to reduce the period of recovery in species with extended reproductive seasons and which make numerous bouts of reproduction, such as C. rubrigularis .     

Sex, Reproductive Status, and Cost of Tail Autotomy via Decreased Running Speed in Lizards

Autotomy, voluntary shedding of body parts to permit escape, is a theoretically interesting defense because escape benefit is offset by numerous costs, including impaired future escape ability. Reduced sprint speed is a major escape cost in some lizards. We predicted that tail loss causes decreased speed in males and previtellogenic females, but not vitellogenic females already slowed by mass gain. In the striped plateau lizard, Sceloporus virgatus , adults of both sexes are subject to autotomy, and females undergo large increases in body condition (mass/length) during vitellogenesis. Time required for running 1 m was similar in intact autotomized males and previtellogenic females, but increased by nearly half after autotomy. Vitellogenic females were slower than other lizards when intact, but their speed was unaffected by autotomy. Following autotomy, speeds of all groups were similar. Thus, speed costs of autotomy vary with sex and reproductive condition: decreased running speed is not a cost of autotomy in vitellogenic females or presumably gravid females. Costs of autotomy are more complex than previously known. Speed and other costs might interact in unforseen ways, making it difficult to predict whether strategies to compensate for diminished escape ability differ with reproductive condition in females.     

Field use of maximal sprint speed by collared lizards (Crotaphytus collaris): compensation and sexual selection

To understand how selection acts on performance capacity, the ecological role of the performance trait being measured must be determined. Knowing if and when an animal uses maximal performance capacity may give insight into what specific selective pressures may be acting on performance, because individuals are expected to use close to maximal capacity only in contexts important to survival or reproductive success. Furthermore, if an ecological context is important, poor performers are expected to compensate behaviorally. To understand the relative roles of natural and sexual selection on maximal sprint speed capacity we measured maximal sprint speed of collared lizards (Crotaphytus collaris) in the laboratory and field-realized sprint speed for the same individuals in three different contexts (foraging, escaping a predator, and responding to a rival intruder). Females used closer to maximal speed while escaping predators than in the other contexts. Adult males, on the other hand, used closer to maximal speed while responding to an unfamiliar male intruder tethered within their territory. Sprint speeds during foraging attempts were far below maximal capacity for all lizards. Yearlings appeared to compensate for having lower absolute maximal capacity by using a greater percentage of their maximal capacity while foraging and escaping predators than did adults of either sex. We also found evidence for compensation within age and sex classes, where slower individuals used a greater percentage of their maximal capacity than faster individuals. However, this was true only while foraging and escaping predators and not while responding to a rival. Collared lizards appeared to choose microhabitats near refugia such that maximal speed was not necessary to escape predators. Although natural selection for predator avoidance cannot be ruled out as a selective force acting on locomotor performance in collared lizards, intrasexual selection for territory maintenance may be more important for territorial males.     

Effects of substrate structure on speed and acceleration capacity in climbing geckos

We studied the effects of substrate structure on locomotor performance in a climbing gecko, Hemidactylus garnoti . We quantified three performance traits (acceleration capacity, instantaneous speed, and final speed) on three substrates: (i) smooth wood, (ii) a cloth surface, and (iii) a wire mesh. While acceleration capacity and instantaneous speed were highest on the wooden surface, final speed did not differ significantly among substrates. Using scanning electron microscopy (SEM) pictures, we estimated that 98% of the wooden surface is available for adhesion by the setae on the toepads, while this percentage is much lower for the mesh and cloth (41 and 37%, respectively). We suggest that when a gecko climbs up a gap-filled substrate, such as the wire mesh or cloth, adhesion will only happen between part of the toepad and the substrate, resulting in a diminished acceleration capacity. The higher acceleration capacity on the wooden substrate and the fact that the geckos tend to slip less often on this particular surface, may explain the difference in instantaneous speed. As for final speed, geckos might achieve similar final speeds on all three substrates by employing different locomotor strategies. Our results suggest that microhabitat use in nature might have a profound effect on locomotor performance and survival for climbing lizards such as geckos.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 385–393.     

Size dimorphism in Rankinia [Tympanocryptis] diemensis (Family Agamidae): sex-specific patterns and geographic variation

Sexual dimorphism has implications for a range of biological and ecological factors, and intersexual morphological differences within a species provide an ideal opportunity for investigating evolutionary influences on phenotypic variation. We investigated sexual size dimorphism (SSD) in an agamid species, Rankinia [Tympanocryptis] diemensis , to determine whether overall size and/or relative morphological trait size differences exist and whether geographic variation in size dimorphism occurs in this species. Relative morphological trait proportions included a range of head, limb, and inter-limb measurements. We found significant overall intersexual adult size differences; females were the larger sex across all sites but the degree of dimorphism between the sexes did not differ between sites. This female-biased size difference is atypical for agamid lizards, which are usually characterized by large male body size. In this species, large female-biased SSD appears to have evolved as a result of fecundity advantages. The size of relative morphological trait also differed significantly between the sexes, but in the opposite direction: relative head, tail, and limb sizes were significantly larger in males than females. This corresponds to patterns in trait size usually found in this taxonomic group, where male head and limb size is important in contest success such as male–male rivalry. There were site-specific morphological differences in hatchlings, including overall body size, tail, inter-limb, thigh, and hindlimb lengths; however, there were no sex-specific differences indicating the body size differences present in the adult form occur during ontogeny.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 699–709.     

Bite-force performance and head shape in a sexually dimorphic crevice-dwelling lizard, the common chuckwalla [Sauromalus ater (= obesus)]

The Common Chuckwalla [ Sauromalus ater (=  obesus )] is a large, sexually dimorphic lizard with a flattened head that takes refuge from predators in rock crevices. Males use their relatively large heads to bite competing males during territorial fights and to restrain females during copulation. Flattened heads with an antipredator function (i.e. seeking refuge in crevices) and enlarged heads with intrasexual competition and reproductive functions suggest possible antagonism between selective pressures on head morphology in males. To examine this hypothesis, we performed a morphometric analysis and measured the bite-force performance of 49 adult chuckwallas. Males had disproportionately wider heads than females, but did not have deeper heads. Males bit with nearly four times the force of females, consistent with the notion of sexual selection for high bite force in males. Although constrained by crevice-wedging behaviour, head depth was a good predictor of bite force in both sexes. In males, however, osteological head width also was a good predictor of bite force. These results are consistent with the hypothesis that head shape in males is under antagonistic selective pressures, which may partly explain the pattern of head shape dimorphism. The disproportionately wide head of males may reflect anatomical modifications to enhance bite force in response to sexual selection in spite of presumed constraints on head shape for crevice-wedging behaviour  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 215–222.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Sexual dimorphism and interpopulation differences in lizard hind limb length: locomotor performance or chemical signalling?

Intraspecific variation in morphology has often been related to fitness differences through its effects on performance. In lizards, variation in hind limb length can be shaped by natural selection for increased locomotor performance, sexual selection on the number or size of femoral pores involved in chemical signalling, or both. Here, we analyse the selective forces involved in sexual dimorphism and differences in hind limb length between two populations of Psammodromus algirus living at different elevation. Males were more robust and had longer hind limbs and limb segments than females, and low‐elevation lizards had longer limbs than high‐elevation lizards. However, differences in locomotor performance were small and non‐significant, making natural selection for faster runs an unlikely explanation for the observed pattern. On the other hand, males had more femoral pores than females, and lizards had more pores at lower elevation, although the difference was significant only for males (which invest more in chemical signalling). In males, the number of pores, which remains constant along a lizard's life, was not correlated with hind limb length. However, femur length was positively correlated with mean pore size, allowing low‐elevation males to have larger than expected pores, which could increase the effectiveness with which they spread their signals in a dry and warm habitat where chemicals become volatile rapidly. Also, saturation of the sexual coloration of the head was higher for low‐elevation males, suggesting that sexual selection pressures may be more intense. Overall, our results indicate that sexual selection plays a significant role in shaping intraspecific variation in hind limb length. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 318–329.