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Parasite load can vary with seasonality, but this is rarely quantified. The garden skink (Lampropholis guichenoti) is host to multiple species of endoparasite. To measure seasonal effects of parasite transmission we established three captive groups of wild-caught individuals in which 2 of 16 individuals (12.5%) were initially infected with nematodes. We collected three faecal samples from each lizard, a sample at the beginning and at the end of the non-activity season and at the end of the following activity season. We measured parasite load (ascarid group) by counting parasite eggs per gram of faeces using a microscope. We found that parasite load was significantly higher in the activity season than in the non-activity season. The prevalence of parasites increased from 15.9% in the non-activity season to 72.5% in the activity season. The activity season is characterised by greater host activity and warmer ambient temperatures, which promote parasite egg survival in the environment as well as egg development. Taken together, this facilitates parasite transmission and could ultimately explain the higher parasite load during the activity season.

Conspicuously coloured signals may evolve via sexual selection to be ornaments or armaments, thereby conferring a fitness advantage to their bearer. Conversely, conspicuous colours may also evolve under natural selection as either aposematic signals or deimatic displays that deter attacks from predators. While conspicuous colour patches may evolve for one purpose (e.g. quality indicators), they may later be co-opted for another (e.g. anti-predator defence). Phrynocephalus mystaceus is a cryptic agamid lizard with flaps in both sexes that when folded against the head are inconspicuous, but when deployed are predicted to be highly conspicuous and to increase the appearance of body size. We tested whether head flaps play a role in social signalling via courtship or as status signals during contests in both sexes. We also tested whether the head flaps have an anti-predator function by simulating predatory encounters. Head flaps were never deployed in courtship or during contests and, therefore, are unlikely to be under sexual selection. However, head flaps and their deployment during simulated predatory encounters were consistent with the predictions associated with deimatic display theory. First, head flaps were similar in form and function between sexes. Second, they were highly conspicuous to both avian and snake predators. Third, there was a rapid transition from crypsis to conspicuousness when they deployed their head flaps during a late stage of predation, the subjugation phase, consistent with an ambush. Confirmation of the deimatic display hypothesis will require future testing of receiver responses.

Sex is a pervasive factor that underpins functional phenotypic variation across a range of traits. Although sex can usually be distinguished morphologically, in some species this is not possible. The development of genetic markers for sex identification is, thus, key if we are to incorporate sex into an understanding of ecological or evolutionary process. Here we develop genetic markers for the identification of sex within an iconic Australian lizard group, the Egernia group, which is notable for its complex social behaviour. We used restriction-site associated DNA sequencing to characterise sex-specific genetic sequences for a key member of the group, Liopholis whitii, and designed primers for four of these putative sex-specific sequences. These primers amplified across some, but not all, species of the group. Our results provided several important insights. They suggest conservatism of a XX/XY sex determination system within the group as well as sex-specific genomic regions that appear independent of the conserved genomic regions identified in other skink species. More broadly, the development of sex markers for the Egernia group opens up a range of potential research questions related to the role that sex plays in the mediation of social behaviour and, through this, the emergence and stability of social life.

Australian lizards are a diverse group distributed across the continent and inhabiting a wide range of environments. Together, they exhibit a remarkable diversity of reproductive morphologies, physiologies, and behaviours that is broadly representative of vertebrates in general. Many reproductive traits exhibited by Australian lizards have evolved independently in multiple lizard lineages, including sociality, complex signalling and mating systems, viviparity, and temperature-dependent sex determination. Australian lizards are thus outstanding model organisms for testing hypotheses about how reproductive traits function and evolve, and they provide an important basis of comparison with other animals that exhibit similar traits. We review how research on Australian lizard reproduction has contributed to answering broader evolutionary and ecological questions that apply to animals in general. We focus on reproductive traits, processes, and strategies that are important areas of current research, including behaviours and signalling involved in courtship; mechanisms involved in mating, egg production, and sperm competition; nesting and gestation; sex determination; and finally, birth in viviparous species. We use our review to identify important questions that emerge from an understanding of this body of research when considered holistically. Finally, we identify additional research questions within each topic that Australian lizards are well suited for reproductive biologists to address.

Thermal conditions during embryonic development affect offspring phenotype in ectotherms. Therefore, rising environmental temperatures can have important consequences for an individual's fitness. Nonetheless, females have some capacity to compensate for potential negative consequences that adverse developmental environments may have on their offspring. Recent studies show that oviparous reptiles exhibit behavioral plasticity in nest site selection, which can buffer their embryos against high incubation temperatures; however, much less is known about these responses in viviparous reptiles. We subjected pregnant viviparous skinks, Saiphos equalis, to current or projected midcentury (2050) temperatures to test (i) how elevated temperatures affect female thermoregulatory and foraging behavior; (ii) whether temperatures experienced by females during pregnancy negatively affect the morphology, performance, and behavior of hatchlings; and (iii) whether behavioral thermoregulation during pregnancy is costly to females. Females from the elevated temperature treatment compensated by going deeper belowground to seek cooler temperatures and eating less, and they consequently had a lower body mass relative to snout-to-vent length (condition estimator) compared with females from the current thermal treatment. The temperatures experienced by females in the elevated temperature treatment were high enough to affect foraging and locomotor performance but not the morphology and growth rate of hatchlings. By seeking cooler temperatures, mothers can mitigate some of the effects of high temperatures on their offspring (e.g., reduced body size and growth). However, this protective behavior of females may come at an energetic cost to them. This study adds to growing evidence of lizards' vulnerability to global warming, particularly during reproduction when females are already paying a substantial cost.

This work was supported by the Australian Research Council (ARC DP130102998 grant to MJW and RWB), Natural Sciences and Engineering Research Council of Canada (scholarship to JLR), the Australasian Society for the Study of Animal Behavior, the Australian Museum, and Macquarie University (scholarship to JLR). The ability to produce viable offspring without recently mating, either through sperm storage or parthenogenesis, can provide fitness advantages under a suite of challenging ecological scenarios. Using genetic analysis, we demonstrate that three wild-caught female Tree Skinks (Egernia striolata) reproduced in captivity with no access to males for over a year, and that this is best explained by sperm storage. To the best of our knowledge, this is the first time female sperm storage has been documented in any monogamous family-living reptile, including social Australian egerniine skinks (from the subfamily Egerniinae). Furthermore, by using paternal reconstruction of genotypes we show that captive-born offspring produced by the same females in the preceding year, presumably without sperm storage, were sired by different males. We qualitatively compared aspects of these females' mates and offspring between years. The parents of each litter were unrelated, but paternal and offspring genotypes from litters resulting from stored sperm were more heterozygous than those inferred to be from recent matings. Family-living egerniine skinks generally have low rates of multiple paternity, yet our study suggests that female sperm storage, potentially from outside social partners, offers the real possibility of benefits. Possible benefits include increasing genetic compatibility of mates and avoiding inbreeding depression via cryptic female choice. Sperm storage in Tree Skinks, a family-living lizard with a monogamous mating system, suggests that females may bet-hedge through extra-pair copulation with more heterozygous males, reinforcing the idea that females could have more control on reproductive outcomes than previously thought. Postprint

Elevated temperatures during development affect a wide range of traits in ectotherms. Less well understood is the impact of global warming on brain development, which has only rarely been studied experimentally. Here, we evaluate current progress in the field and search for common response patterns among ectotherm groups. Evidence suggests that temperature may have a positive effect on neuronal activity and growth in developing brains, but only up to a threshold, above which temperature is detrimental to neuron development. These responses appear to be taxon dependent but this assumption may be due to a paucity of data for some taxonomic groups. We provide a framework with which to advance this highly promising field in the future.

Gene expression levels are key molecular phenotypes at the interplay between genotype and environment. Mounting evidence suggests that short-term changes in environmental conditions, such as those encountered in captivity, can substantially affect gene expression levels. Yet, the exact magnitude of this effect, how general it is, and whether it results in parallel changes across populations are not well understood. Here, we take advantage of the well-studied cane toad, Rhinella marina, to examine the effect of short-term captivity on brain gene expression levels, and determine whether effects of captivity differ between long-colonized and vanguard populations of the cane toad's Australian invasion range. We compared the transcriptomes of wild-caught toads immediately assayed with those from toads captured from the same populations but maintained in captivity for seven months. We found large differences in gene expression levels between captive and wild-caught toads from the same population, with an over-representation of processes related to behaviour and the response to stress. Captivity had a much larger effect on both gene expression levels and gene expression variability in toads from vanguard populations compared to toads from long-colonized areas, potentially indicating an increased plasticity in toads at the leading edge of the invasion. Overall, our findings indicate that short-term captivity can induce large and population-specific transcriptomic changes, which has significant implications for studies comparing phenotypic traits of wild-caught organisms from different populations that have been held in captivity.

While foraging or during social interactions, animals may benefit from judging relative quantity. Individuals may select larger prey or a patch with more food and, likewise, it may pay to track the number and type of individuals and social interactions. We tested for spontaneous quantity discrimination in the gidgee skink (Egernia stokesii), a family-living lizard. Lizards were presented with food quantities differing in number or size and were able to select the larger quantity of food items when they differed in number but not when items differed in size. We show, for the first time, superior spontaneous discrimination of items differing in number over size in a lizard species, which contrasts with previous findings. Our simple method, however, did not include controls for the use of continuous quantities, and further tests are required to determine the role of such information during quantity discrimination. Our results provide support for the use of the parallel individuation system for the discrimination of small quantities (four or fewer items). Lizards might, however, still use the approximate number system if items in larger quantities (more than four) are presented. Overall, we uncovered evidence that species might possess specific cognitive abilities potentially adapted to their niche with respect to quantity information (discrete and/or continuous) and the processing system used when judging quantities. Importantly, our results highlight the need for testing multiple species using similar testing procedures to gain a better understanding of the underlying causes leading to differences across species.

Conspicuous social and sexual signals are predicted to experience pronounced character release when natural selection via predation is relaxed. However, we have few good examples of this phenomenon in the wild and none in species with dynamic color change. Here, we show that Jackson’s chameleons inadvertently introduced from Kenya to Hawaii (Oahu), where there are no coevolved, native lizard predators, experienced pronounced character release of color signals. Hawaiian chameleons displayed more conspicuous social color signals than Kenyan chameleons during male contests and courtship, were less cryptic in response to bird and snake predators, and showed greater change between display and antipredator color states. Hawaiian chameleon display colors were also more conspicuous in their local than ancestral habitats, consistent with local adaptation of social signals. These results demonstrate that relaxed predation pressure can result in character release of dynamic social signals in introduced species experiencing strong sexual selection.

Sexual selection shapes the adaptive landscape in complex ways that lead to trait integration. Much of our understanding of selection comes from studies of morphological traits. However, few studies attempt to quantify the form and direction of selection on performance even though it is predicted to be a more direct target of selection in nature. We measured sexual selection on performance traits (bite force, sprint speed and endurance) in an Australian lizard, the Eastern water skink (Eulamprus quoyii). We first staged 123 contests between size-matched males to investigate whether performance traits were important in determining contest outcome. In a second experiment, we established six breeding populations in large replicate semi-natural enclosures to estimate whether performance traits predicted reproductive success. Our results show that none of the performance measures were important in predicting contest outcome and were not generally strong predictors of reproductive success. However, our analyses suggest a complex fitness landscape driven by males adopting different alternative reproductive tactics (ARTs). We provide a rare test of the role performance plays in sexual selection and highlight the need to test common assumptions regarding the link between maximal performance and fitness. Our results suggest that performance traits may not necessarily be direct targets of sexual selection, but rather indirect targets through their integration with morphological and/or behavioural traits, highlighting a need for more explicit tests of the predicted links between performance and fitness.

Recently, there has been a surge in cognition research using non-avian reptile systems. As a diverse group of animals, non-avian reptiles [turtles, the tuatara, crocodylians, and squamates (lizards, snakes and amphisbaenids)] are good model systems for answering questions related to cognitive ecology, from the role of the environment on the brain, behaviour and learning, to how social and life-history factors correlate with learning ability. Furthermore, given their variable social structure and degree of sociality, studies on reptiles have shown that group living is not a pre-condition for social learning. Past research has demonstrated that non-avian reptiles are capable of more than just instinctive reactions and basic cognition. Despite their ability to provide answers to fundamental questions in cognitive ecology, and a growing literature, there have been no recent systematic syntheses of research in this group. Here, we systematically, and comprehensively review studies on reptile learning. We identify 92 new studies investigating learning in reptiles not included in previous reviews on this topic - affording a unique opportunity to provide a more in-depth synthesis of existing work, its taxonomic distribution, the types of cognitive domains tested and methodologies that have been used. Our review therefore provides a major update on our current state of knowledge and ties the collective evidence together under nine umbrella research areas: (i) habituation of behaviour, (ii) animal training through conditioning, (iii) avoiding aversive stimuli, (iv) spatial learning and memory, (v) learning during foraging, (vi) quality and quantity discrimination, (vii) responding to change, (viii) solving novel problems, and (ix) social learning. Importantly, we identify knowledge gaps and propose themes which offer important future research opportunities including how cognitive ability might influence fitness and survival, testing cognition in ecologically relevant situations, comparing cognition in invasive and non-invasive populations of species, and social learning. To move the field forward, it will be immensely important to build upon the descriptive approach of testing whether a species can learn a task with experimental studies elucidating causal reasons for cognitive variation within and among species. With the appropriate methodology, this young but rapidly growing field of research should advance greatly in the coming years providing significant opportunities for addressing general questions in cognitive ecology and beyond.

Urban wildlife faces a novel set of challenges resulting in selective pressure that can lead to population-level changes. We studied Australian water dragons (Intellagama lesueurii) from urban and natural populations to test if urban populations differed in body size, shape, and performance capacity. If urban-derived morphology has arisen through selection, we predicted distinct morphological differences between wild dragons from urban and natural areas in both adult and hatchling life-stages. Urban hatchlings were morphologically distinct (shorter body lengths and longer limbs) from natural populations, while urban adult males continued this trend but only for body size (shorter body lengths). We then experimentally reared hatchlings originating from urban and natural populations within urban- and natural-style enclosures (2 × 2 factorial design) for a year to determine if differences in morphology and performance capacity (sprint speed, endurance, and clinging ability) were related to either the individual’s origin population or developmental environment. Yearlings reared in urban-style enclosures, irrespective of population origin, had smaller body sizes compared to those from natural-style enclosures, suggesting developmental environment was affecting their morphology. Despite this difference in body size, yearling dragon performance capacity was not significantly different between treatments. Overall, this study provides evidence of a complex relationship driving urban-divergent morphology – whereby urban dragons emerge as smaller hatchlings with longer limbs (innate traits) and are then further influenced by the urban environments that they develop in (phenotypic plasticity); however, and potentially owing to behavioral, ecological, and demographical differences, these changes appear to be sex-specific.

Early developmental environments influence life-history traits and survival of reptiles. In fact, rising global temperatures have already caused widespread extinctions among lizards. Viviparous species might be more susceptible to increasing temperatures because of their inability to meet their energetic demands following rapid environmental changes. Nevertheless, we do not yet fully understand how lizards with different reproductive modes can respond to climate change. We investigated the effect of both maternal thermal environment during pregnancy and incubation temperature on hatchling morphology and physiological performance of two populations of the lizard Saiphos equalis differing in their mode of reproduction, to test whether reproductive mode affects the ability to buffer against rising temperatures. Gravid females from both populations were subjected to current or projected end-of-century (future) thermal environments, to evaluate differences in the body size, growth rate, thermal preference, and locomotor performance of their offspring. Our results show that independently of the mode of reproduction, high temperatures accelerated gestation periods. Thermal environments did not affect hatchling thermal preference, but viviparous hatchlings consistently preferred lower temperatures. Unlike viviparous lizards, oviparous hatchlings incubated under future temperatures were smaller and had a lower growth rate compared to current-incubated hatchlings. Finally, thermal environments did not affect hatchling endurance and speed when controlling for body size. Our results show that global warming is likely to have a negative impact on S. equalis, but suggest that some of its effects may be ameliorated by maternal responses during pregnancy, particularly in viviparous populations.

One of the most widely studied abilities in lizards is discrimination learning. The protocols used to test lizards are often novel or adapted from other taxa without proper validation. We need to ensure that tests of discrimination learning are appropriate and properly applied in lizards so that robust inferences can be made about cognitive ability. Here, we present a new protocol for testing lizard discrimination learning that incorporates a target training procedure, uses many daily trials for efficiency and reinforcement, and has a robust, validated, learning criterion. We trained lizards to touch a cue card using operant conditioning and tested lizards separately on a colour, and pattern discrimination test. Lizards successfully learnt to touch a cue card and to discriminate between light and dark blue but had issues discriminating the patterns. After modifying the test procedure, some lizards reached criterion, revealing possible issues with stimulus processing and interference of generalisation. Here, we describe a protocol for operant conditioning and two-choice discrimination learning in lizards with a robust learning criterion that can help researcher better design future studies on discrimination learning in lizards.

Anticipatory behaviour is the expectation of a near-future event based on information processed in the past and influences an animal's tactical decisions, particularly when there are significant fitness consequences. The grass lizard (Takydromus viridipunctatus) perches on blades of grass at night which likely reduces the probability of predation by terrestrial predators such as snakes, rodents and shrews. During twilight (starting 30 min before sunrise), they move from above the grass to within grass clumps and this is thought to afford the lizard protection while reducing detection by avian predators. Here, we examined how lizards shift their behaviour as a function of visual detectability to their primary predator, the cattle egret (Bubulcus ibis). We show that the lizards shift from their perch site during twilight at the earliest time at which egrets depart communal roosts. At the same time, visual modelling shows a dramatic increase in the detectability of the lizards to the visual system of egrets. Therefore, anticipatory behaviour in response to environmental cues acts to reduce predation risk as lizards become more conspicuous and predators become more active. Grass lizard anticipatory behaviour appears to be finely tuned by natural selection to adjust to temporal changes in predation risk.

To make decisions, animals gather information from the environment in order to avoid costs (e.g., reduced survival) and increase benefits (e.g., foraging success). When time is limited or information is insufficient, most animals face a speed-accuracy trade-off (SAT) - they have to balance the benefits of making quick decisions against the costs of inaccurate decisions. Here, we investigated the relationship between decision accuracy and decision speed in gidgee skinks (Egernia stokesii) performing a food-based spontaneous quantity discrimination task. Rather than a SAT we found a speed-accuracy alignment; lizards made decisions that were fast and accurate, rather than inaccurate. Furthermore, we found only within-, but no between-individual, differences in decision making indicating behavioural plasticity in the absence of individual decision styles. Finally, latency to choice was highly repeatable, more so than choice accuracy. Previous work has shown that learning, the costs of a bad decision and task difficulty frequently result in SATs. The lack of a SAT in our lizards might be a direct consequence of our simple testing methodology which prevented learning by not allowing lizards to consume the chosen quantity. To fully understand how SATs develop, different methodologies that control the costs and benefits of decisions should be compared.

Background Quantity discrimination, the ability to discriminate a magnitude of difference or discrete numerical information, plays a key role in animal behavior. While quantitative ability has been well documented in fishes, birds, mammals, and even in previously unstudied invertebrates and amphibians, it is still poorly understood in reptiles and has never been tested in an aquatic turtle despite the fact that evidence is accumulating that reptiles possess cognitive skills and learning ability. To help address this deficiency in reptiles, we investigated the quantitative ability of an Asian freshwater turtle, Mauremys sinensis, using red cubes on a white background in a trained quantity discrimination task. While spontaneous quantity discrimination methods are thought to be more ecologically relevant, training animals on a quantity discrimination task allows more comparability across taxa. Results We assessed the turtles’ quantitative performance in a series of tests with increasing quantity ratios and numerosities. Surprisingly, the turtles were able to discriminate quantities of up to 9 versus 10 (ratio = 0.9), which shows a good quantitative ability that is comparable to some endotherms. Our results showed that the turtles’ quantitative performance followed Weber’s law, in which success rate decreased with increasing quantity ratio across a wide range of numerosities. Furthermore, the gradual improvement of their success rate across different experiments and phases suggested that the turtles possess learning ability. Conclusions Reptile quantitative ability has long been ignored and therefore is likely under-estimated. More comparative research on numerical cognition across a diversity of species will greatly contribute to a clearer understanding of quantitative ability in animals and whether it has evolved convergently in diverse taxa.

When faced with a predator, some animals engage in a deimatic display to startle the predator momentarily, resulting in a pause or retreat, thereby increasing their chance of escape. Frillneck lizards (Chlamydosaurus kingii) are characterised by a large, pronounced frill that extends from the base of the head to beyond the neck and, when displayed, can be up to six times the width of the head. We used behavioural assays with a model avian predator to demonstrate that their display conforms to deimatic display theory. First, juveniles and adults deployed the frill in encounters with a model predator. Second, the display revealed three colour patches (white and red–orange patches on the frill; yellow mouth palate) that facilitate a transition from a cryptic to a conspicuous state as perceived by a raptor visual system. Third, the display was performed with movements that amplified its effect. The frill area was larger in males than in females, which suggests that the frill might also be co-opted for male–male contests. If future research confirms a role of the frill in male agonistic interactions, frillneck lizards will be a rare case in which a structure has a dual function in a deimatic display and a sexually selected signal.

Given that graduate employment is a clear outcome of education success, there is an urgent need to conceptualize course design strategically to maximize students’ chances of employment. In this pap...

Our knowledge of the conservation status of reptiles, the most diverse class of terrestrial vertebrates, has improved dramatically over the past decade, but still lags behind that of the other tetrapod groups. Here, we conduct the first comprehensive evaluation (~92% of the world’s ~1,714 described species) of the conservation status of skinks (Scincidae), a speciose reptile family with a worldwide distribution. Using International Union for Conservation of Nature (IUCN) criteria, we report that ~20% of species are threatened with extinction, and nine species are Extinct or Extinct in the Wild. The highest levels of threat are evident in Madagascar and the Neotropics, and in the subfamilies Mabuyinae, Eugongylinae and Scincinae. The vast majority of threatened skink species were listed based primarily on their small geographic ranges (Criterion B, 83%; Criterion D2, 13%). Although the population trend of 42% of species was stable, 14% have declining populations. The key threats to skinks are habitat loss due to agriculture, invasive species, and biological resource use (e.g., hunting, timber harvesting). The distributions of 61% of species do not overlap with protected areas. Despite our improved knowledge of the conservation status of the world’s skinks, 8% of species remain to be assessed, and 14% are listed as Data Deficient. The conservation status of almost a quarter of the world’s skink species thus remains unknown. We use our updated knowledge of the conservation status of the group to develop and outline the priorities for the conservation assessment and management of the world’s skink species.

A key goal of conservation is to protect biodiversity by supporting the long-term persistence of viable, natural populations of wild species. Conservation practice has long been guided by genetic, ecological and demographic indicators of risk. Emerging evidence of animal culture across diverse taxa and its role as a driver of evolutionary diversification, population structure and demographic processes may be essential for augmenting these conventional conservation approaches and decision-making. Animal culture was the focus of a ground-breaking resolution under the Convention on the Conservation of Migratory Species of Wild Animals (CMS), an international treaty operating under the UN Environment Programme. Here, we synthesize existing evidence to demonstrate how social learning and animal culture interact with processes important to conservation management. Specifically, we explore how social learning might influence population viability and be an important resource in response to anthropogenic change, and provide examples of how it can result in phenotypically distinct units with different, socially learnt behavioural strategies. While identifying culture and social learning can be challenging, indirect identification and parsimonious inferences may be informative. Finally, we identify relevant methodologies and provide a framework for viewing behavioural data through a cultural lens which might provide new insights for conservation management. Fundação Caloust Gulbenkian - FCT info:eu-repo/semantics/publishedVersion

The habenula is a small structure in the brain that acts as a relay station for neural information, helping to modulate behaviour in response to variable and unpredictable stimuli. Broadly, it is evolutionarily conserved in structure and connectivity across vertebrates and is the most prominent bilaterally asymmetric structure in the brain. Nonetheless, comparative evolutionary studies of the habenula are virtually non-existent. Here, we examine the volumes of the medial and lateral habenular subregions, in both hemispheres, across a group of Australian agamid lizards in the genus Ctenophorus. In males, we found bilaterally asymmetrical selection on the lateral habenula to become smaller with increasing intensity of sexual selection, possibly as a mechanism to increase aggressive responses. In females, we found bilaterally symmetrical selection on both the medial and lateral subregions to become smaller with increasing sexual selection. This is consistent with sexual selection increasing motivation to reproduce and the habenula’s well-characterized role in controlling and modifying responses to rewarding stimuli. However, as there are currently no studies addressing habenular function in reptiles, it is difficult to draw more precise conclusions. As has happened recently in biomedical neuroscience, it is time for the habenula to receive greater attention in evolutionary neuroscience.

Assisted reproductive technologies for population and genetic management for threatened herpetofauna have grown substantially in the past decade. Here we describe experiments to optimise sperm cryopreservation in a model squamate, the eastern water skink Eulamprus quoyii. Small, concentrated volumes of highly motile spermatozoa were reliably collected from adult male E. quoyii by non-lethal ventral massage. Samples were used to: (1) test whether protein-rich diluents, namely Beltsville poultry semen extender (BPSE) and TES and Tris (TEST) yolk buffer (TYB), improve post-thaw quality metrics compared with Dulbecco’s phosphate-buffered saline (DPBS); and (2) compare the efficacy of these diluents in combination with either 1.35 M glycerol or 1.35 M dimethyl sulfoxide (DMSO) at two freezing rates, fast (approximately –20°C min−1) versus slow (–6°C min−1). Glycerol and DMSO performed equally well in preserving spermatozoa under slow freezing rates. Under these conditions, the use of the complex diluents BPSE and TYB significantly improved post-thaw total motility compared with DPBS. Complex interactions occurred between cryodiluent type, cryoprotectant and freezing rate when testing fast versus slow freezing rates among treatment groups. Under slow freezing rates, DMSO was better at preserving membrane integrity and motility, regardless of diluent type, but successful fast freezing required complex diluents to support motility and membrane integrity, which has implications for implementation in a field setting.

We tested learning and behavioral flexibility in family-living gidgee skinks (Egernia stokesii) using a multistage visual discrimination task that included acquisition and reversal stages using simple and compound stimuli composed of black shapes superimposed on a colored background. We evaluated how lizards learn compound cues through a probe test. Lizards showed behavioral flexibility through reversal learning using simple stimuli (only color or shape). Our lizards used compound stimuli to learn a discrimination but had problems reversing and generalizing across novel compound stimuli. In the probe test, lizards chose the correct stimulus in a novel pairing with a distractor feature even without previous experience with compound stimuli. Our results suggest that some lizards are likely able to attend selectively to the relevant features of our compound stimuli while ignoring irrelevant features instead of using the configuration of a cue card as a whole to learn to discriminate between compound stimuli. We hope that our work will spark interest in further studies looking at how lizards (and reptiles in general) learn to solve visual discrimination problems. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Animals that habitually cross the boundary between water and land face specific challenges with respect to locomotion, respiration, insulation, fouling and waterproofing. Many semi-aquatic invertebrates and plants have developed complex surface microstructures with water-repellent properties to overcome these problems, but equivalent adaptations of the skin have not been reported for vertebrates that encounter similar environmental challenges. Here, we document the first evidence of evolutionary convergence of hydrophobic structured skin in a group of semi-aquatic tetrapods. We show that the skin surface of semi-aquatic species of Anolis lizards is characterized by a more elaborate microstructural architecture (i.e. longer spines and spinules) and a lower wettability relative to closely related terrestrial species. In addition, phylogenetic comparative models reveal repeated independent evolution of enhanced skin hydrophobicity associated with the transition to a semi-aquatic lifestyle, providing evidence of adaptation. Our findings invite a new and exciting line of inquiry into the ecological significance, evolutionary origin and developmental basis of hydrophobic skin surfaces in semi-aquatic lizards, which is essential for understanding why and how the observed skin adaptations evolved in some and not other semi-aquatic tetrapod lineages., Summary: Multiple Anolis lineages independently evolved a similar skin surface microarchitecture with water-repellent properties as an adaptation to a semi-aquatic lifestyle.

Biological invasions are a contemporary global threat because invasive species can have substantial negative economic and ecological impacts. Invasive species can outcompete native species through two main mechanisms: interference competition (direct, negative interactions like aggression) and/or exploitative competition (indirect, negative interactions resulting from species using the same, limited resources like food). The invasive Italian wall lizard (Podarcis siculus) was introduced into Lisbon, Portugal, 20 years ago, and is believed to be locally displacing the native green Iberian wall lizard (Podarcis virescens). We experimentally tested for competition between these two lizard species by establishing heterospecific (one pair of each species) and conspecific (two pairs of the same species; control) treatments in enclosures containing a high- and a low-quality refuge. Lizards were fed from food dishes every other day. We tested if species showed interference (aggressive behaviour, stealing food and shelter exclusion) or exploitative competition (tolerance between species but differences in food consumption efficiency). We found evidence for exploitative competition: the invasive species arrived first at food stations, consumed more food and gained more weight than the native species. We suggest that exploitative competition may, in part, explain the observed displacement of P. virescens from contact areas with the invasive P. siculus. Deciphering the competitive mechanisms between invasive and native species is vital for understanding the invasion process. To become successful invaders, alien species must often outcompete native species they encounter in a new location. But how can an alien species outcompete a resident with a long evolutionary history in an environment in which the resident is expected to be better adapted? We studied an invasive and native congeneric pair of sympatric lizards to understand how they interact and potentially compete in a controlled environment. The invasive Italian wall lizard and a native congener were very tolerant of each other; however, the invasive species was first to arrive at food, ate more and grew faster, suggesting exploitative competition. This contrasts with previous studies in other introduced locations where the Italian wall lizard was more aggressive towards native lizards, suggestive of interference competition. Our results help explain why the Italian wall lizard is so successful and suggests it may compete in different ways, possibly in response to local environmental conditions and which species it may be competing with, but with the same outcome: the displacement of native species.

Inhibitory control, the inhibition of prepotent actions, is essential for higher-order cognitive processes such as planning, reasoning, and self-regulation. Individuals and species differ in inhibitory control. Identifying what influences inhibitory control ability within and between species is key to understanding how it evolved. We compared performance in the cylinder task across five lizard species: tree skinks (Egernia striolata), gidgee skinks (Egernia stokesii), eastern blue-tongue skinks (Tiliqua s. scincoides), sleepy lizards (Tiliqua r. asper), and eastern water skinks (Eulamprus quoyii). In our task, animals had to inhibit the prepotent motor response of directly approaching a reward placed within a semi-transparent mesh cylinder and instead reach in through the side openings. Additionally, in three lizard species, we compared performance in the cylinder task to reversal learning to determine the task specificity of inhibitory ability. Within species, neither sex, origin, body condition, neophobia, nor pre-experience with other cognitive tests affected individual performance. Species differed in motor response inhibition: Blue-tongue skinks made fewer contacts with the semi-transparent cylinder wall than all other species. Blue-tongue skinks also had lower body condition than the other species which suggest motivation as the underlying cause for species differences in task performance. Moreover, we found no correlation between inhibitory ability across different experiments. This is the first study comparing cylinder task performance among lizard species. Given that inhibitory control is probably widespread in lizards, motor response inhibition as exercised in the cylinder task appears to have a long evolutionary history and is likely fundamental to survival and fitness.SignificanceThe study of lizard cognition is receiving increasing attention. Lizards are a diverse group with a wide range of ecological attributes and represent a model system through which we can test a wide range of hypotheses relating to cognitive evolution. Furthermore, considering their evolutionary history, studying non-avian reptile cognition can help understand the evolution of different cognitive abilities including inhibitory control. Here, we provide a comparison of inhibitory control ability in five lizard species. Consequently, we are able to, firstly, validate a method (the cylinder task) initially developed for the use in mammals and birds, for use in lizards, and secondly, collect valuable data on inhibitory control in a poorly studied group with respect to cognitive ability. Our study suggests non-cognitive factors as a major influence on cylinder task performance, which is in agreement with previous studies of other vertebrates.

Magnetic resonance imaging (MRI) is an established technique for neuroanatomical analysis, being particularly useful in the medical sciences. However, the application of MRI to evolutionary neuroscience is still in its infancy. Few magnetic resonance brain atlases exist outside the standard model organisms in neuroscience and no magnetic resonance atlas has been produced for any reptile brain. A detailed understanding of reptilian brain anatomy is necessary to elucidate the evolutionary origin of enigmatic brain structures such as the cerebral cortex. Here, we present a magnetic resonance atlas for the brain of a representative squamate reptile, the Australian tawny dragon (Agamidae: Ctenophorus decresii), which has been the subject of numerous ecological and behavioral studies. We used a high-field 11.74T magnet, a paramagnetic contrasting-enhancing agent and minimum-deformation modeling of the brains of thirteen adult male individuals. From this, we created a high-resolution three-dimensional model of a lizard brain. The 3D-MRI model can be freely downloaded and allows a better comprehension of brain areas, nuclei, and fiber tracts, facilitating comparison with other species and setting the basis for future comparative evolution imaging studies. The MRI model and atlas of a tawny dragon brain (Ctenophorus decresii) can be viewed online and downloaded using the Wiley Biolucida Server at wiley.biolucida.net.

This project was funded by an ARC Discovery grant (DP130102998) to Martin Whiting and Richard Byrne and by Macquarie University. Behavioural flexibility, the ability to adjust behaviour to environmental change by adapting existing skills to novel situations, is key to coping with, for example, complex social interactions, seasonal changes in food availability or detecting predators. We tested the tree skink, Egernia striolata, a family-living skink from eastern Australia, in a set-shifting paradigm of eight colour/shape discriminations including reversals, an intradimensional acquisition of a new colour/shape and extradimensional shift from colour to shape (and vice versa). Skinks could learn to discriminate between colour/shape pairs and reverse this initial stimulus–reward association; however, they showed no significant decrease in the probability of making a correct choice in the extradimensional shift suggesting that they did not form an attentional set. Subjects appear to have learnt each stage as a new problem instead of generalizing stimuli into specific dimensions (set formation). In conclusion, tree skinks solved a discrimination reversal by focusing their attention towards visual stimuli and flexibly adjusting their choice behaviour accordingly. These lizards learned to use multidimensional visual stimuli to find a food reward but did not generalize stimuli into dimensions. Furthermore, this study is the first to test for set shifting in a lizard species and thereby allows us to extend set-shifting theory to a new taxon for comparison with primates, rodents, a bird and a turtle. Postprint