Is Habitat Preference Associated with Locomotor Performance in Multiocellated Racerunners(Eremias multiocellata) from a Desert Steppe?

Locomotor performance in lizards is strongly affected by structural habitat. Understanding this relationship allows us to predict species distributions across habitat types. However, little information is available about the ecological role of the locomotion of multiocellated racerunner （Eremias multiocellata） in the desert steppe ecosystem of Inner Mongolia, China. Herein, we studied the effects of habitat structure on the locomotor performance of this lizard species in the field. We found that the sprint speed of this lizard declined significantly with increasing vegetation coverage. Manipulative experiments were further conducted to examine the effects of branch barriers and surface substrates on the sprint speed of the lizard. We found that the sprint speed was significantly influenced by the surface substrates and branch barriers, and there were no interactions between them. Branch barriers impeded sprint speed, and E. multiocellata showed better locomotor performance on sandy rather than loamy substrates. Our results indicate that E. multiocellata tends to occupy open areas with sandy substrates, but its locomotor performance is not closely associated with habitat preference.

Diet, Food Intake of Phrynocephalus frontalis (Agamidae) and Its Potential Role in Desert Habitat

We examined the dietary diversity and food intake of Phrynocephalus frontalis, compared the difference of insect diversity in the natural habitats with different lizard densities, and discussed the potential role of this lizard in the desert ecosystem. The results show that： （1） arthropodans of the orders Coleoptera, Hymenoptera and Hemiptera were major dietary components of P. frontalis; （2） coleoptera larvae always formed the predominant component of lizard diets; （3） dietary diversities of P. frontalis were not significantly different between summer and autumn or between the two sexes; （4） the similarity in trophic niches between seasons was 0.756, whereas the similarity in trophic niches between sexes was 0.994; （5） stomach content weight of lizards varied significantly among different seasons, but there was no significant difference in stomach content weight between sexes; （6） insect diversity differed significantly among the groups of the habitat with different degrees of lizard density, and the habitat with moderate lizards density had the highest insect diversity. We infer that P. frontalis prey mainly on insects and change their diet and food intake with season; males and females consumed similar preys in types and weights. As an important predator, P. frontalis could affect the insect community in the arid ecosystem of Hunshandak Desert on the Mongolian Plateau.

Effects of Sand Grain Size on Habitat Selection in Steppe Toad-headed Lizard(Phrynocephalus frontalis)

To understand the effects of sand grain size on habitat selection, we conducted a field study on the steppe toad-headed lizard in Hunshandake Desert northern China. Methods of mark-recapture and transect survey were used to investigate the density of steppe toad-headed lizards and the environmental variables. The comparison on lizard densities among the habitats with different environmental factors revealed that： 1） population density of the steppe toad-headed lizard differed significantly among the habitats with different sand grain size indexes（SGSIs, representing roughness of sand substrate）： the highest lizard density was found in the group with an SGSI of 〉 0.30, whereas the lowest density was found in the group with an SGSI of 0–0.15; and 2） vegetation cover, soil moisture, invertebrate diversity index, and abundance had no significant effects on the lizard density. These results implied that the sand grain size was the most important determinant of habitat selection for steppe toad-headed lizards in Hunshandake Desert. Steppe toad-headed lizards could avoid structural habitats that have negative effects on their maximal sprinting capabilities. Considering the changing sand grain size in the development phase of sand dunes, the sand lizard could be used as an indicator of the process of desertification.

Thermal-physiological Strategies Underlying the Sympatric Occurrence of Three Desert Lizard Species

Sympatric reptiles are the ideal system for investigating temperature-driven coexistence. Understanding thermally physiological responses of sympatric lizards is necessary to reveal the physiological mechanisms that underpin the sympatric occurrence of reptiles. In this study, we used three lizard species, Eremias argus, E. multiocellata, and Phrynocephalus przewalskii, which are sympatric in the Inner Mongolia desert steppe, as a study system. By comparing their resting metabolic rates(RMR) and locomotion at different body temperatures, we aimed to better understand their physiological responses to thermal environments, which may explain the sympatric occurrence of these lizards. Our results showed that E. argus had significantly higher RMR and sprint speed than E. multiocellata, and higher RMR than P. przewalskii. In addition, the optimal temperature that maximized metabolic rates and locomotion for E. argus and E. multiocellata was 36°C, whereas for P. przewalskii it was 39°C. Our study revealed the physiological responses to temperatures that justify the sympatric occurrence of these lizards with different thermal and microhabitat preferences and active body temperatures. Eremias argus and E. multiocellata, which have lower body temperatures than P. przewalskii, depend on higher RMR and locomotion to compensate for their lower body temperatures in field conditions. Our study also highlights the importance of using an integrative approach, combining behavior and physiology, to explore the basis of sympatric occurrence in ectothermic species.

Spatial Distribution Pattern of the Steppe Toad-headed Lizard(Phrynocephalus frontalis) and Its Influencing Factors

Spatial distribution patterns are associated with life history and behavioral adaptations of animals. For studying the spatial distribution pattern of the steppe toad-headed lizard(Phrynocephalus frontalis) and its influencing factors,we conducted experiments in Hunshandake Sandy Land in Inner Mongolia,China in July 2009. By calculating the clustered indices,we found that the lizard was aggregately distributed when the sampling quadrat was smaller than 10 m × 10 m,and uniformly distributed when it was greater than 10 m × 10 m. The Nearest Neighbor Rule showed a clustering distribution pattern for P. frontalis and the distribution pattern was quadrat-sampling dependent. Furthermore,the cluster was determined by environmental factors when the sampling quadrat was smaller than 20 m × 20 m,but it was determined by both environmental factors and characteristics of the lizard when it was larger than 20 m × 20 m. Our results suggested that the steppe toad-headed lizards tended to aggregate into suitable habitat patches in desert areas. Additionally,we discussed that the lizard aggregation could be potentially used as an indictor of movement of sand dunes.

Behavior of Phrynocephalus frontalis to Avoid Traps

Previous studies have shown that reptiles are capable of spatial learning and learn to locate important environmental resources so that they can return to those locations at a future time, when needed. Lizards improve their adaptability and survival by learning the position of their shelter in a complex environment. This behavior raises the question, whether lizards can sense danger, such as a trap, in their surroundings, by determining the location of the trap and avoiding it. In the present study, we used a pitfall trap to test if steppe toad-headed agama, Phrynocephalus frontalis, can learn to recognize the position of the trap and avoid it. Our results revealed that the percentage of activity time in the trap area was significantly reduced （P 〈 0.001） and the time of drop trap was also significantly reduced （P = 0.00631）. The number of burrows dug by lizards distributed in the trap area was the least. Reduced activity time in the trap area was observed to have no obvious relationship with the drop in the number of burrows. The present study, therefore, demonstrates that P. frontalis are capable of learning the avoidance of a trap by locating its position. The findings offer significant insight in the understanding of reptilian behavior, which is important in the study of the role of reptiles in global ecology, especially because they are often very sensitive to environmental changes.

Phenotypic consequences of maternally selected nests:a cross-fostering experiment in a desert lizard

Despite the importance of maternally selected nests in shaping offspring phenotypes,our understanding of how the nest environment affects embryonic development and offspring traits of most non-avian reptiles is rather limited largely due to the logistical difficulty in locating their nests.To identify the relative contributions of environmental(temporal[seasonal]and spatial[nest-site])and intrinsic(clutch)factors on embryonic development and offspring traits,we conducted a cross-fostering experiment by swapping eggs between maternally-selected nests of the toad-headed agama(Phrynocephalus przewalskii)in the field.We found that nest environment explained a large proportion of variation in incubation duration,hatching success,and offspring size and growth.In contrast,clutch only explained a small proportion of variation in these embryonic and offspring traits.More significantly,compared with spatial effects,seasonal effects explained more phenotypic variation in both embryonic development and offspring traits.Eggs laid early in the nesting season had longer incubation durations and produced smaller hatchlings with higher post-hatching growth rates than did later-laid eggs.Consequently,hatchlings from early-laid eggs reached larger body sizes prior to winter.In addition,we found that female toad-headed agama did not select nests specific to reaction norms of their own offspring because hatchlings from original or translocated nests had similar phenotypic traits.Overall,our study demonstrates the importance of seasonal variation in nest environments in determining embryonic development and offspring phenotypes,which has not been widely appreciated at least in non-avian reptiles.