Short-Term Dispersal Response of an Endangered Australian Lizard Varies with Time of Year

Dispersal is an important component in the demography of animal populations. Many animals show seasonal changes in their tendency to disperse, reflecting changes in resource availability, mating opportunities, or in population age structure at the time when new offspring enter the population. Understanding when and why dispersal occurs can be important for the management of endangered species. The pygmy bluetongue lizard is an endangered Australian species that occupies and defends single burrow refuges for extended periods of time, rarely moving far from the burrow entrance. However, previous pitfall trapping data have suggested movement of adult males in spring and of juveniles in autumn of each year. In the current study we compared behaviours of adult lizards each month, over the spring-summer activity period over two consecutive field seasons, to provide deeper understanding of the seasonal dispersal pattern. We released adult pygmy bluetongue lizards into a central area, provided with artificial burrows, within large enclosures, and monitored the behaviour and movements of the released lizards over a four day period. There was a consistent decline in time spent basking, amount of movement around burrow entrances, and rates of dispersal from the central release area from early spring to late summer. Results could be relevant to understanding and managing natural populations and for any translocation attempts of this endangered lizard species.     

Spiders provide important resources for an endangered lizard

Conservation management requires knowledge of how a target species interacts with other species. Some relatively common species can modify the environment to the advantage of rarer, endangered species. Thus, local enhancement of those common species can positively influence remaining populations of the rarer species. The endangered pygmy bluetongue lizard Tiliqua adelaidensis inhabits burrows that are constructed by lycosid and mygalomorph spiders. We recorded 490 burrows in a 1 ha plot at the end of one season, and then observed at regular intervals the formation and loss of burrows, and the changes in occupancy status of each burrow over the next season. We found spiders in 94% of all newly constructed burrows and deduced that they had built the burrows. We found no evidence that lizards dug new burrows or deepened existing burrows. The numbers of both lizards and spiders in the burrows declined over the spring and summer, with lizards moving from their burrows more often early in the season than later. However, there was no strong trend for lizards to replace spiders in burrows. In fact, lizards tended to occupy deeper burrows than spiders, suggesting little negative impact of lizards on spiders. However, spiders had a positive impact on lizards by providing the refuge burrows central for lizard survival. Although lizards readily accept artificial burrows, long-term conservation for the lizards must include viable spider populations to maintain a supply of suitable burrow refuges.     

Responses of the endangered pygmy bluetongue lizard to conspecific scats

Many animals use chemical signals for communication between conspecifics and for territory marking. The pygmy bluetongue lizard is normally solitary, focussing activity around the entrance of its burrow, from where it ambushes prey, and rarely contacts other individuals. In this paper we examined whether lizards in laboratory experiments alter their behaviour in the presence of scats from conspecifics. In the first experiment, when lizards were offered a choice of two vacant burrows with or without a scat close to the entrance, they tongue flicked more often at the burrow entrance when the scat was present, and more often chose to occupy the burrow with the scat. An interpretation is that lizards use scat signals to recognise burrows that may be suitable because they have previously been occupied by a conspecific, but that they approach those burrows cautiously in case a resident is still present and likely to resist a takeover. Scats from male lizards were inspected (by both sexes) for longer than scats of female lizards. In the second experiment, when resident lizards were presented with scats outside of their burrows, they inspected and tongue flicked at those scats more often if the scat came from a male than a female lizard, but there was no definitive evidence from our experiments that lizards differentiated in their response to scats from lizards that were found close to or far from the test lizard. The results were consistent with a communication system in which lizards use scats to advertise their presence, independent of any direct contact.     

Co‐occupancy of spider‐engineered burrows within a grassland community changes temporally

Burrow‐digging organisms act as ecosystem engineers, providing potential habitat to other organisms. In the Mid North region of South Australia, wolf and trapdoor spiders in fragmented grassland communities provide this service. Pygmy bluetongue lizards are an endangered skink, endemic to these grasslands. The lizards obligatorily use burrows dug by these spider groups as refuges, basking sites and ambush points. We investigated the occupancy of these spider burrows by lizards and other organisms within the grassland community, identifying the occasions that burrows were shared by multiple taxa. We found that the lizards and trapdoor spiders are predominantly solitary, while wolf spiders co‐shared burrows more frequently with either weevils or snails. There were numerous taxa that were found to regularly co‐share with other taxa, particularly snails, centipedes and weevils. There was a strong temporal influence on burrow sharing, with most co‐sharing occurring late in summer. This study provides an insight into the use of burrows by the lizards and co‐existing taxa within these grassland communities. The dynamics of burrow‐use by other taxa have the potential to influence long‐term conservation of these lizards as burrow availability is crucial to their survival in these grasslands.     

Hold your breath: Observations of the endangered pygmy bluetongue (Tiliqua adelaidensis) submerged in flooded burrows

Understanding adaptations to extreme weather events by endangered species is critical to inform conservation decisions, particularly when their adaptations relate to artificial habitat supplementation at translocation sites. Apnoea, temporary suspension of breathing, has been observed as an anti-predator adaptation by semi-aquatic reptiles that dive underwater for periods of time to avoid detection. This study reports on the observations of an endangered grassland skink, the pygmy bluetongue (Tiliqua adelaidensis), remaining submerged in rain-induced flooded artificial burrows at an experimental translocation site.     

The Ontogeny of Display Behaviour in Sceloporus undulatus: (Sauria: Iguanidae)

A longitudinal study (from neonate to adult) was made of the display behaviour of 36 fence lizards from 8 clutches. A frame-by-frame analysis of 941 video-taped displays revealed that the hatchlings possessed almost all of their adult display patterns by the first two days of life. Of the total variance in the display behaviour, 7% or less was attributed to experience. Of the variance among lizards, approximately half came from between clutches and half from between clutchmates. Our data indicate that the visual communication signals (head bob displays) of this vertebrate species are almost totally innate.     

Survivorship and Spatial Patterns of an Urban Population of Texas Horned Lizards

Habitat fragmentation has negative consequences on threatened and endangered species by creating isolated populations. The Texas horned lizard (Phrynosoma cornutum) is experiencing population declines and localized extirpations throughout its range and has been classified as a species of greatest conservation need in Oklahoma, USA. Younger age classes have been poorly studied but may be vital to the stability of remaining populations. To address gaps in knowledge concerning subadult (hatchling and juvenile) morphometrics, survivorship, and home range sizes, we studied 2 cohorts of subadults, for 2 years each, covering their hatching and juvenile years (2016–2019). We used a combination of radio-telemetry and novel harmonic radar methodology to study a closed population of Texas horned lizards in 15 ha of native grassland at Tinker Air Force Base, Oklahoma. Population abundance for adults and juveniles was estimated as 56.5 ± 5.5 lizards and density as 7.96 lizards/ha. Our lowest estimates of survival indicated an average survival probability for the hatchling life stage of 0.285 (95% CI = 0.15–0.44), which is lower than for adults on the site. Average home range size increased from hatchling to adult life stages. Our results will have an immediate effect on the planning and assessment of ongoing headstart and management programs for Texas horned lizards. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Maternal exposure to predator scents: offspring phenotypic adjustment and dispersal

Predation is a strong selective pressure generating morphological, physiological and behavioural responses in organisms. As predation risk is often higher during juvenile stages, antipredator defences expressed early in life are paramount to survival. Maternal effects are an efficient pathway to produce such defences. We investigated whether maternal exposure to predator cues during gestation affected juvenile morphology, behaviour and dispersal in common lizards (Zootoca vivipara). We exposed 21 gravid females to saurophagous snake cues for one month while 21 females remained unexposed (i.e. control). We measured body size, preferred temperature and activity level for each neonate, and released them into semi-natural enclosures connected to corridors in order to measure dispersal. Offspring from exposed mothers grew longer tails, selected lower temperatures and dispersed thrice more than offspring from unexposed mothers. Because both tail autotomy and altered thermoregulatory behaviour are common antipredator tactics in lizards, these results suggest that mothers adjusted offspring phenotype to risky natal environments (tail length) or increased risk avoidance (dispersal). Although maternal effects can be passive consequences of maternal stress, our results strongly militate for them to be an adaptive antipredator response that may increase offspring survival prospects.     

Invader danger: lizards faced with novel predators exhibit an altered behavioral response to stress

Animals respond to stressors by producing glucocorticoid stress hormones, such as corticosterone (CORT). CORT acts too slowly to trigger immediate behavioral responses to a threat, but can change longer-term behavior, facilitating an individual's survival to subsequent threats. To be adaptive, the nature of an animal's behavior following elevated CORT levels should be matched to the predominant threats that they face. Seeking refuge following a stressful encounter could be beneficial if the predominant predator is a visual hunter, but may prove detrimental when the predominant predator is able to enter these refuge sites. As a result, an individual's behavior when their CORT levels are high may differ among populations of a single species. Invasive species impose novel pressures on native populations, which may select for a shift in their behavior when CORT levels are high. We tested whether the presence of predatory invasive fire ants (Solenopsis invicta) at a site affects the behavioral response of native eastern fence lizards (Sceloporus undulatus) to elevated CORT levels. Lizards from an uninvaded site were more likely to hide when their CORT levels were experimentally elevated; a response that likely provides a survival advantage for lizards faced with native predatory threats (e.g. birds and snakes). Lizards from a fire ant invaded site showed the opposite response; spending more time moving and up on the basking log when their CORT levels were elevated. Use of the basking log likely reflects a refuge-seeking behavior, rather than thermoregulatory activity, as selected body temperatures were not affected by CORT. Fleeing off the ground may prove more effective than hiding for lizards that regularly encounter small, terrestrially-foraging fire ant predators. This study suggests that invasive species may alter the relationship between the physiological and behavioral stress response of native species.     

The consequences of lifetime and evolutionary exposure to toxic prey: changes in avoidance behaviour through ontogeny

Responses to novel threats (e.g. invasive species) can involve genetic changes or plastic shifts in phenotype. There is controversy over the relative importance of these processes for species survival of such perturbations, but we are realizing they are not mutually exclusive. Native eastern fence lizards (Sceloporus undulatus) have adapted to top-down predation pressure imposed by the invasive red imported fire ant (Solenopsis invicta) via changes in adult (but not juvenile) lizard antipredator behaviour. Here, we examine the largely ignored, but potentially equally important, bottom-up effect of fire ants as toxic prey for lizards. We test how fire ant consumption (or avoidance) is affected by lifetime (via plasticity) and evolutionary (via natural selection) exposure to fire ants by comparing field-caught and laboratory-reared lizards, respectively, from fire ant-invaded and uninvaded populations. More naive juveniles from invaded populations ate fire ants than did adults, reflecting a natural ontogenetic dietary shift away from ants. Laboratory-reared lizards from the invaded site were less likely to eat fire ants than were those from the uninvaded site, suggesting a potential evolutionary shift in feeding behaviour. Lifetime and evolutionary exposure interacted across ontogeny, however, and field-caught lizards from the invaded site exhibited opposite ontogenetic trends; adults were more likely to eat fire ants than were juveniles. Our results suggest that plastic and evolutionary processes may both play important roles in permitting species survival of novel threats. We further reveal how complex interactions can shape adaptive responses to multimodal impacts imposed by invaders: in our system, fire ants impose stronger bottom-up selection than top-down selection, with each selection regime changing differently across lizard ontogeny.     

Behavior and temperature modulate a thermoregulation–predation risk trade‐off in juvenile gopher tortoises

Ectotherms frequently thermoregulate behaviorally to improve physiological processes such as digestion and growth, but basking and other thermoregulatory activities can also increase predation risk. Organismal and environmental characteristics can, in some species, influence predation risk associated with thermoregulation and thereby relax or tighten constraints on thermoregulatory behavior, physiological performance, and, ultimately, life history traits. Providing one of the first such investigations in turtles, we examine whether behavior and thermal environment modulate a thermoregulation–predation risk trade‐off in juvenile gopher tortoises (Gopherus polyphemus). Young gopher tortoises experience very high predation pressure, and their declining species faces many challenges, including human‐induced increased shading of its environment. We hypothesized that in response to simulated predator approach, basking hatchling and juvenile tortoises would: (i) hide inside burrows; (ii) hide for shorter durations in cooler burrows presumably due to greater constraints on physiological performance; and (iii) spend greater time at the surface following disturbance in cool environments because individuals would need to bask more to maintain preferred body temperatures. Basking tortoises always hid inside burrows when approached and exhibited very long flight initiation distances ( = 45 m) that increased with age/size. Individuals fled into burrows even when it was not possible for them to see the approaching researcher, suggesting the use of vibrations (aerial/ground) to detect potential predators and a possible antipredator function for exceptionally large otoliths characteristic of the species. Tortoises hid for short durations ( = 18.3 min), especially in cool burrows, suggesting that they optimize hiding responses to balance physiological costs and antipredator benefits. Additionally, surface activity following disturbance consisted primarily of basking and correlated negatively with burrow temperature. These findings suggest that thermal environment influences predation risk in ectotherms whose surface activity is driven primarily by thermoregulatory requirements and highlight potential benefits of warm, well‐insolated habitats, such as endangered longleaf pine (Pinus palustris) ecosystems, for juvenile gopher tortoises.     

Effects of grazing regime on the relative body mass of the endangered pygmy bluetongue lizard (Tiliqua adelaidensis)

Many animals rely on fat reserves, to keep them alive through extended periods of food shortage, such as the winter, and to provide additional energy for reproduction. Fat reserves, measured relative to an animal's size, are often referred to as the animal's body condition. The present study investigated how different levels of grazing by domestic stock in native grassland habitat affect the body condition of the pygmy bluetongue lizard (Tiliqua adelaidensis), and if these effects are related to changes in the abundance and size of grasshoppers which are the lizards primary source of food. The initial hypothesis was that lizards would have the highest body condition in moderately grazed paddocks, because those paddocks would have more grasshoppers than heavily grazed paddocks, and better visual conditions for lizards to catch those grasshoppers than in ungrazed paddocks. The results, however, showed that both lizard body condition and the abundance of grasshoppers increased with decreasing grazing intensity. The connection between lizard body condition and abundance of grasshoppers was complex. Within an activity season, lizard body condition generally declined from spring to summer, while the number of grasshoppers grew. The mean size of grasshoppers seemed to be more important, as lizard body condition was higher in spring, the time of year with the largest grasshoppers. These results show that the intensity of grazing by domestic stock influences the body condition of pygmy bluetongue lizards, but that this effect is not entirely due to the reduction in the number of grasshoppers resulting from grazing.     

Parasite in peril? A new species of mite in the genus Ophiomegistus Banks (Parasitiformes: Paramegistidae) on an endangered host,the pygmy bluetongue lizard Tiliqua adelaidensis (Peters) (Squamata: Scincidae)

Host‐parasite relationships are generally understudied in wild populations but have a potential to influence host population dynamics and the broader ecosystem, which becomes particularly important when the host is endangered. Herein we describe a new species of parasitic mite from the genus Ophiomegistus (Parasitiformes: Mesostigmata: Paramegistidae) of an endangered South Australian skink; the pygmy bluetongue lizard (Tiliqua adelaidensis). Adult mites were observed on lizard hosts in three different host populations, among which prevalence varied. No temporal trend in prevalence was evident over two spring‐summer seasons of monitoring. We hypothesise that the reliance on burrows as refuges by T. adelaidensis may be essential for the completion of the mite life cycle and also for horizontal transmission. The conservation implications of not only its effect on the host, but also its potential status as an endangered species itself, are considered.     

Promiscuous mating in the endangered Australian lizard Tiliqua adelaidensis: a potential windfall for its conservation

Studies have revealed an unsuspected complexity in social systems within a few lizard species, including group living, long-term monogamy and individual recognition of partners or offspring. Comparisons among these species and their relatives could provide valuable insights, allowing us to investigate traits that are shared across social systems and identify general principles relating to the evolution of sociality. The endangered pygmy bluetongue lizard, Tiliqua adelaidensis, is a member species in the Egernia group, but is thought to show a more solitary social structure than other members in this group. Within this study we used microsatellite markers to determine the mating system of T. adelaidensis. Unlike many other species in the Egernia group, we found a predominately promiscuous mating system in T. adelaidensis. We detected multiple paternity in 75 % of litters. Of the 70 males identified as having fathered juveniles, only 5 were identified as mating with the same female in more than 1 year and only 3 were identified as the father of juveniles with the same female in consecutive years. The genetic evidence suggested that partners were chosen randomly with respect to the level of relatedness among neighbouring lizards. However, mated lizards were geographically closer to each other than expected by random chance. Multiple paternities rely on the opportunity for males to encounter multiple females during the period when they are receptive to mating, and this may depend on population densities. Drivers for the polygamous mating system may be the single occupancy burrow and the central place territorial defence of those burrows in T. adelaidensis. We propose a fourth mating system for the Egernia group: polygyny within stable non-social colonies.     

Cranial morphology and bite force in Chamaeleolis lizards--adaptations to molluscivory?

Anolis lizards have become a model system for the study of adaptive radiations as species with similar morphologies occupying similar habitats have arisen independently on all the larger islands in the Caribbean. However, on both, Cuba and Hispaniola unique forms have evolved that seemingly have no counterparts on any of the other Caribbean islands. Anoles of the genus Chamaeleolis comprise such a unique form and have been termed 'twig giants' because of their cryptic life style, slow locomotor mode, and short limbs. However, some of the most unusual features of these lizards are their large heads and molluscivorous diet. Here, we compare head shape, bite force, and muscle structure among sexes and age classes of Chamaeleolis lizards with Anolis crown giants. Our data show that Chamaeleolis lizards have a dramatically different head shape characterized by tall heads with a pronounced temporal ridge and long snouts. Analyses of bite force, surprisingly, show no differences between adult Chamaeleolis and Anolis crown giants. Juveniles of Chamaeleolis, however, have very tall heads for their size and bite harder than Anolis juveniles do. This can be related to the propensity of juveniles of this genus to eat snails, food items for which high bite forces are crucial. This observation is corroborated by the presence of well-developed jaw adductors in juveniles. Thus, our data suggest that the unusual phenotype of adults with large and tall heads may be due to selection on the juvenile life history stages.     

Intraspecific alkaloid variation in ladybird eggs and its effects on con- and hetero-specific intraguild predators

Sexual size dimorphism (SSD) is a common phenomenon in animals. In many species females are substantially larger than males. Because body size plays a central role in modulating the body temperature (T _b) of ectotherms, intersexual differences in body size may lead to important intersexual differences in thermoregulation. In addition, because SSD is realized by differences in growth rate and because growth rate is strongly temperature dependent in ectotherms, a conflict between male reproductive behaviour and thermoregulation may affect the expression of SSD. In this study, we investigated the thermal implications of SSD in a reptile exhibiting spectacular female-biased SSD: the northern map turtle (Graptemys geographica). Over three seasons, we collected >150,000 measurements of T _b in free-ranging adult and juvenile northern map turtles using surgically implanted miniature temperature loggers. Northern map turtles exhibited seasonal patterns of thermoregulation typical of reptiles in northern latitudes, but we found that large adult females experienced a lower daily maximum T _b and a narrower daily range of T _b than adult males and small juvenile females. In addition, despite more time spent basking, large adult females were not able to thermoregulate as accurately as small turtles. Our findings strongly suggest that body size limits the ability to thermoregulate accurately in large females. By comparing thermoregulatory patterns between adult males and juvenile females of similar body size, we found no evidence that male reproductive behaviours are an impediment to thermoregulation. We also quantified the thermal significance of basking behaviour. We found, contrary to previous findings, that aerial basking allows northern map turtles to raise their T _b substantially above water temperature, indicating that basking behaviour likely plays an important role in thermoregulation.     

Using social networks to deduce whether residents or dispersers spread parasites in a lizard population

1. Heterogeneity of host behaviour can play an important role in the spread of parasites and pathogens around wildlife populations. Social networks have previously been suggested to represent transmission pathways within a population, but where the dynamics of host-parasite interactions are difficult to observe, networks may also be used to provide insights into transmission processes. 2. Pygmy bluetongue lizards, Tiliqua adelaidensis, occupy individual territories, live exclusively in burrows constructed by spiders in Australian native grasslands and are hosts to a tick, Bothriocroton hydrosauri, and a nematode, Pharyngodon wandillahensis. 3. On five monthly occasions, the locations of all individual lizards in three study plots were used to construct weighted, undirected networks based on proximity of adjacent burrows. 4. The networks were used to explore alternative hypotheses about the spread of each parasite through the population: that stable population members that remained in the same burrow over the study period played a major role in influencing the pattern of infection or that dispersing individuals played a more significant role. 5. For ticks, host individuals that were infected were more connected in the network than uninfected hosts and this relationship remained significant for connections to residents in the population, but not for connections to dispersers. 6. For nematodes, infected and uninfected hosts did not differ in their overall strength of connection in the network, but infected hosts were more connected to dispersers than were uninfected hosts, suggesting that lizards moving across the population are the major agents for the transmission of nematodes. 7. This study shows how network analyses can provide new insights into alternative pathways of parasite spread in wildlife populations, where it is difficult to make direct observations of transmission-related behaviours.     

Maternal basking behaviour determines offspring sex in a viviparous reptile

Two primary dichotomies within vertebrate life histories involve reproductive mode (oviparity versus viviparity) and sex determination (genotypic sex determination versus environmental sex determination). Although reptiles show multiple evolutionary transitions in both parameters, the co-occurrence of viviparity and environmental-dependent sex determination have heretofore been regarded as incompatible. Our studies on the viviparous lizard Niveoscincus ocellatus show that the extent of basking by a female influences the sex of her offspring. Critically, our data reveal this effect both in the field (via correlations between date of birth and litter sex ratio) and in a laboratory experiment (females with reduced basking opportunities produced more male offspring). Changes in thermoregulatory behaviour thus allow pregnant female lizards to modify the sex of their offspring.     

Clustering of related individuals in a population of the Australian lizard, Egernia frerei

Stable social aggregations are rarely recorded in lizards, but have now been reported from several species in the Australian scincid genus Egernia. Most of those examples come from species using rock crevice refuges that are relatively easy to observe. But for many other Egernia species that occupy different habitats and are more secretive, it is hard to gather the observational data needed to deduce their social structure. Therefore, we used genotypes at six polymorphic microsatellite DNA loci of 229 individuals of Egernia frerei, trapped in 22 sampling sites over 3500 ha of eucalypt forest on Fraser Island, Australia. Each sampling site contained 15 trap locations in a 100 x 50 m grid. We estimated relatedness among pairs of individuals and found that relatedness was higher within than between sites. Relatedness of females within sites was higher than relatedness of males, and was higher than relatedness between males and females. Within sites we found that juvenile lizards were highly related to other juveniles and to adults trapped at the same location, or at adjacent locations, but relatedness decreased with increasing trap separation. We interpreted the results as suggesting high natal philopatry among juvenile lizards and adult females. This result is consistent with stable family group structure previously reported in rock dwelling Egernia species, and suggests that social behaviour in this genus is not habitat driven.