Revealing a Vacancy Analog of the Crowdion Interstitial in Simple Cubic Crystals.

Vacancies in simple cubic crystals of hard cubes are known to delocalize over one-dimensional chains of several lattice sites. Here, we use computer simulations to examine the structure and dynamics of vacancies in simple cubic crystals formed by hard cubes, right rhombic prisms (slanted cubes), truncated cubes, and particles interacting via a soft isotropic pair potential. We show that these vacancies form a vacancy analog of the crowdion interstitial, generating a strain field which follows a soliton solution of the sine-Gordon equation, and diffusing via a persistent random walk. Surprisingly, we find that the structure of these "voidions" is not significantly affected by changes in density, vacancy concentration, and even particle interaction. We explain this structure quantitatively using a one-dimensional model that includes the free-energy barrier particles have to overcome to slide between lattice sites and the effective pair interaction along this line. We argue that voidions are a robust phenomenon in systems of repulsive particles forming simple cubic crystals.

Phase and vacancy behaviour of hard "slanted" cubes.

We use computer simulations to study the phase behaviour for hard, right rhombic prisms as a function of the angle of their rhombic face (the "slant" angle). More specifically, using a combination of event-driven molecular dynamics simulations, Monte Carlo simulations, and free-energy calculations, we determine and characterize the equilibrium phases formed by these particles for various slant angles and densities. Surprisingly, we find that the equilibrium crystal structure for a large range of slant angles and densities is the simple cubic crystal-despite the fact that the particles do not have cubic symmetry. Moreover, we find that the equilibrium vacancy concentration in this simple cubic phase is extremely high and depends only on the packing fraction and not the particle shape. At higher densities, a rhombic crystal appears as the equilibrium phase. We summarize the phase behaviour of this system by drawing a phase diagram in the slant angle-packing fraction plane.

Climate-related environmental variation in a visual signalling device: the male and female dewlap in Anolis sagrei lizards.

Animals communicate using a variety of signals that differ dramatically among and within species. The astonishing dewlap diversity in anoles has attracted considerable attention in this respect. Yet, the evolutionary processes behind it remain elusive and have mostly been explored for males only. Here, we considered Anolis sagrei males and females to study signal divergence among populations. First, we assessed the degree of variation in dewlap design (size, pattern and colour) and displays by comparing 17 populations distributed across the Caribbean. Second, we assessed whether the observed dewlap diversity is associated with variation in climate-related environmental conditions. Results showed that populations differed in all dewlap characteristics, with the exception of display rate in females. We further found that males and females occurring in 'xeric' environments had a higher proportion of solid dewlaps with higher UV reflectance. In addition, lizards inhabiting 'mesic' environments had primarily marginal dewlaps showing high reflectance in red. For dewlap display, a correlation with environment was only observed in males. Our study provides evidence for a strong relationship between signal design and prevailing environmental conditions, which may result from differential selection on signal efficacy. Moreover, our study highlights the importance of including females when studying dewlaps in an evolutionary context.

How phylogeny and foraging ecology drive the level of chemosensory exploration in lizards and snakes.

The chemical senses are crucial for squamates (lizards and snakes). The extent to which squamates utilize their chemosensory system, however, varies greatly among taxa and species' foraging strategies, and played an influential role in squamate evolution. In lizards, 'Scleroglossa' evolved a state where species use chemical cues to search for food (active foragers), whereas 'Iguania' retained the use of vision to hunt prey (ambush foragers). However, such strict dichotomy is flawed as shifts in foraging modes have occurred in all clades. Here, we attempted to disentangle effects of foraging ecology from phylogenetic trait conservatism as leading cause of the disparity in chemosensory investment among squamates. To do so, we used species' tongue-flick rate (TFR) in the absence of ecological relevant chemical stimuli as a proxy for its fundamental level of chemosensory investigation, that is baseline TFR. Based on literature data of nearly 100 species and using phylogenetic comparative methods, we tested whether and how foraging mode and diet affect baseline TFR. Our results show that baseline TFR is higher in active than ambush foragers. Although baseline TFRs appear phylogenetically stable in some lizard taxa, that is a consequence of concordant stability of foraging mode: when foraging mode shifts within taxa, so does baseline TFR. Also, baseline TFR is a good predictor of prey chemical discriminatory ability, as we established a strong positive relationship between baseline TFR and TFR in response to prey. Baseline TFR is unrelated to diet. Essentially, foraging mode, not phylogenetic relatedness, drives convergent evolution of similar levels of squamate chemosensory investigation.

Evolution of a designless nanoparticle network into reconfigurable Boolean logic.

Natural computers exploit the emergent properties and massive parallelism of interconnected networks of locally active components. Evolution has resulted in systems that compute quickly and that use energy efficiently, utilizing whatever physical properties are exploitable. Man-made computers, on the other hand, are based on circuits of functional units that follow given design rules. Hence, potentially exploitable physical processes, such as capacitive crosstalk, to solve a problem are left out. Until now, designless nanoscale networks of inanimate matter that exhibit robust computational functionality had not been realized. Here we artificially evolve the electrical properties of a disordered nanomaterials system (by optimizing the values of control voltages using a genetic algorithm) to perform computational tasks reconfigurably. We exploit the rich behaviour that emerges from interconnected metal nanoparticles, which act as strongly nonlinear single-electron transistors, and find that this nanoscale architecture can be configured in situ into any Boolean logic gate. This universal, reconfigurable gate would require about ten transistors in a conventional circuit. Our system meets the criteria for the physical realization of (cellular) neural networks: universality (arbitrary Boolean functions), compactness, robustness and evolvability, which implies scalability to perform more advanced tasks. Our evolutionary approach works around device-to-device variations and the accompanying uncertainties in performance. Moreover, it bears a great potential for more energy-efficient computation, and for solving problems that are very hard to tackle in conventional architectures.

Genetic divergence among sympatric colour morphs of the Dalmatian wall lizard (Podarcis melisellensis).

If alternative phenotypes in polymorphic populations do not mate randomly, they can be used as model systems to study adaptive diversification and possibly the early stages of sympatric speciation. In this case, non random mating is expected to support genetic divergence among the different phenotypes. In the present study, we use population genetic analyses to test putatively neutral genetic divergence (of microsatellite loci) among three colour morphs of the lizard Podarcis melisellensis, which is associated with differences in male morphology, performance and behaviour. We found weak evidence of genetic divergence, indicating that gene flow is somewhat restricted among morphs and suggesting possible adaptive diversification.

Characterization of polymorphic microsatellite markers in the Dalmatian wall lizard Podarcis melisellensis (Squamata: Lacertidae).

We describe polymerase chain reaction primers and amplification conditions for 13 highly polymorphic microsatellite DNA loci isolated from the Dalmatian wall lizard, Podarcis melisellensis. The number of alleles per locus ranged from 12 to 41, with levels of observed heterozygosity between 0.62 and 0.94. Most of these loci were successfully cross-amplified in the closely related species P. sicula, but levels of polymorphism were always lower.

Does foraging mode mould morphology in lacertid lizards?

Evolutionary changes in foraging style are often believed to require concurrent changes in a complex suite of morphological, physiological, behavioural and life-history traits. In lizards, species from families with a predominantly sit-and-wait foraging style tend to be more stocky and robust, with larger heads and mouths than species belonging to actively foraging families. Here, we test whether morphology and foraging behaviour show similar patterns of association within the family Lacertidae. We also examine the association of bite force abilities with morphology and foraging behaviour. Lacertid lizards exhibit considerable interspecific variation in foraging indices, and we found some evidence for a covariation between foraging style and body shape. However, the observed relationships are not always in line with the predictions. Also, the significance of the relationships varies with the evolutionary model used. Our results challenge the idea that foraging style is evolutionarily conservative and invariably associated with particular morphologies. It appears that the flexibility of foraging mode and its morphological correlates varies among lizard taxa.

Fight versus flight: physiological basis for temperature-dependent behavioral shifts in lizards.

Previous studies have demonstrated that a behavioral shift from flight to aggressive behavior occurs at low temperatures in some lizards. Our data for the agamid lizard Trapelus pallida demonstrate how the effect of temperature on whole organism performance traits such as sprint speed (much lower performance at lower temperature) and bite force (largely independent of temperature) may explain the shift from flight to fight behavior with decreasing temperature. Moreover, our data hint at the physiological basis for this effect as isolated muscle power output, twitch and tetanus time traits, relevant to sprinting, appear to be strongly temperature-dependent muscle properties. Maximal muscle force production, on the other hand, appears largely independent of temperature. Unexpectedly, differences in the physiological properties of jaw versus limb muscle were observed that enhance the ability of the jaw muscle to generate maximal force at all temperatures tested. Thus our data show how behavioral responses may be determined by the limitations set by temperature on physiological processes.

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.

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.

Proximate causes of intraspecific variation in locomotor performance in the lizard Gallotia galloti.

To understand the evolution of biological traits, information on the degree and origins of intraspecific variation is essential. Because adaptation can take place only if the trait shows heritable variation, it is important to know whether (at least) part of the trait variation is genetically based. We describe intra- and interindividual variation in three performance measures (sprint speed, climbing, and clambering speed) in juvenile Gallotia galloti lizards from three populations and examine how genetic, environmental (incubation temperature), and ontogenetic (age, size) effects interact to cause performance variation. Moreover, we test whether the three performance traits are intercorrelated phenotypically and genetically. Sprint speed is highest in juveniles incubated at the lowest temperature (26 degrees C) irrespective of population. Climbing speed differs among populations, and the differences persist at least until the lizards are 30 wk old. This suggests that the three populations experience different selective pressures. Moreover, mass, snout-vent length, and hindlimb length seem to affect climbing performance differently in the three populations. The variation in sprinting and climbing ability appears to be genetically based. Moreover, the two performance traits are intercorrelated and thus will not evolve independently from each other. Clambering speed (i.e., capacity to climb up an inclined mesh) varies among individuals, but the origin of this variation remains obscure.

Speed and stamina trade-off in lacertid lizards.

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.

Spatio-temporal gait characteristics of level and vertical locomotion in a ground-dwelling and a climbing gecko.

The effects of incline (vertical versus horizontal) on spatio-temporal gait characteristics (stride and step length, frequency, duty factor, degree of sprawling) were measured over a range of speeds in a ground-dwelling (Eublepharis macularius) and a climbing (Gekko gecko) species of gecko. Surprisingly, the climbing species also performs very well when moving on the horizontal substratum. In the present experiments, climbing speeds ranged from 0.6 to 1.2 m s(-1), whereas speeds for level locomotion were between 0.6 and 1.8 m s(-1). In contrast, the vertical climbing capacities of the ground-dweller are limited (speeds below 0.1 m s(-1 )versus level speeds between 0.2 and 1.1 m s(-1)). In general, we demonstrate that very little adjustment in gait characteristics is made by either species when they are forced to move on their non-habitual substratum. Moreover, gait characteristics differ little between the species despite the clear differences in ecological niche. Higher level or climbing speeds are realized mainly (or exclusively in the case of level locomotion in G. gecko) by increasing stride frequency. Stride lengths and duty factors vary with speed in the ground-dweller, but not in the climbing species. Step length and the degree of sprawling are speed-independent (except for hind-limb sprawling in G. gecko on the level). It is argued that this common strategy suits climbing (fixed spatial variables, no floating phases) rather than level locomotion.

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.

Terrestrial locomotion in the black-billed magpie: kinematic analysis of walking, running and out-of-phase hopping.

The inter-limb kinematic patterns of walking, running and out-of-phase hopping in black-billed magpies (Pica pica) were studied using high-speed video recordings. The flexion/extension patterns of the joints were similar between the gait types, suggesting that the within-leg control of the angular excursions is similar. This result is further supported by the fact that running and hopping are alternative gaits at speeds higher than walking; however, magpies show a preference for hopping. Moreover, only small differences occur between the kinematic patterns of the two limbs during out-of-phase hopping, during which the legs are believed to have different functions. The hindlimb kinematic patterns of magpies are like those of other flying and more terrestrial bird species; however, striking differences are found in comparison with humans at the level of the internal angles. This is probably due to the differences in the morphology and configuration of their legs.

Spatio-temporal gait characteristics of the hind-limb cycles during voluntary bipedal and quadrupedal walking in bonobos (Pan paniscus).

Spatio-temporal gait characteristics (step and stride length, stride frequency, duty factor) were determined for the hind-limb cycles of nine bonobos (Pan paniscus) walking quadrupedally and bipedally at a range of speeds. The data were recalculated to dimensionless quantities according to the principle of dynamic similarity. Lower leg length was used as the reference length. Interindividual variability in speed modulation strategy of bonobos appears to be low. Compared to quadrupedal walking, bipedal bonobos use smaller steps to attain a given speed (differences increase with speed), resulting in shorter strides at a higher frequency. In the context of the ("hybrid") dynamic pattern approach to locomotion (Latach, 1998) we argue that, despite these absolute differences, intended walking speed is the basic control variable which elicits both quadrupedal and bipedal walking kinematics in a similar way. Differences in the initial status of the dynamic system may be responsible for the differences in step length between both gaits. Comparison with data deduced from the literature shows that the effects of walking speed on stride length and frequency are similar in bonobos, common chimpanzees, and humans. This suggests that (at least) within extant homininae, spatio-temporal gait characteristics are highly comparable, and this in spite of obvious differences in mass distribution and bipedal posture.

Absolute versus per unit body length speed of prey as an estimator of vulnerability to predation.

To study whether absolute (m/s) or relative (body lengths/s) speed should be used to compare the vulnerability of differently sized animals, we developed a simple computer simulation. Human 'predators' were asked to 'catch' (mouse-click) prey of different sizes, moving at different speeds across a computer screen. Using the simulation, a prey's chances of escaping predation depended on its speed (faster prey were more difficult to catch than slower prey of the same body size), but also on its size (larger prey were easier to catch than smaller prey at the same speed). Catching time, the time needed to catch a prey, also depended on both prey speed and prey size. Relative prey speed (body lengths/s or body surface/s) was a better predictor of catching time than was absolute prey speed (m/s). Our experiment demonstrates that, in contrast to earlier assertions, per unit body length speed of prey may be more 'ecologically relevant' than absolute speed. Copyright 1998 The Association for the Study of Animal Behaviour.