sphenodon punctatus
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<p>Anthropogenic climate change is progressing at a rate unprecedented in the past 65 million years and is a significant conservation concern. The associated biotic and abiotic impacts are expected to have substantial effects on global biodiversity, with some species potentially more vulnerable than others. The tuatara (Sphenodon punctatus) is a New Zealand endemic reptile and of particular interest as it is a slowly reproducing, range-restricted, cold-adapted ectotherm with temperature-dependent sex determination. Consequently, tuatara could be particularly vulnerable to rising air temperatures and conservation translocations have been key components of tuatara conservation efforts. Knowledge of how the tuatara might be affected by warmer climates will help inform where future conservation efforts are best directed, practices to avoid and which sites might be most suitable for the establishment of populations. The translocation of 176 adult tuatara in October 2012 from Stephens Island in New Zealand’s Cook Strait to four latitudinally distant North Island sites offered the opportunity to study the responses of tuatara in a range of environments. The comparatively warmer, drier climates of several sites provided surrogates for temporal climate change, enabling an assessment of how a warming climate might impact tuatara, and how they might respond. Using field observations, laboratory analysis and controlled experiments I investigated the short-term success of the translocations, the influence of translocation and climate on tuatara enteric bacterial communities and parasites, as well as how warmer climates might influence nocturnal activity, thermoregulatory opportunities and learning ability. I found several translocated populations to be progressing favourably, and found evidence that tuatara may exhibit enhanced growth at warmer, less densely-populated sites, suggesting that further translocations to lower latitude sites might be a viable conservation strategy. However, high population density at one translocation site was a concern and management recommendations were made to enable the dispersal of individuals. I detected Salmonella Saintpaul for the first time in a live tuatara, Campylobacter spp. was identified as a likely common commensal organism, and no measurable impact of translocation or climate on bacterial prevalence was observed, suggesting no substantial risk of climate warming to the susceptibility of tuatara to these bacteria. Tick populations were negatively impacted by translocation-associated factors following release but subsequently recovered at most sites and mites were not found on any translocated tuatara. Diurnal and nocturnal activities were positively influenced by air temperature, up to an upper threshold, and assessment of the site-specific thermal climates suggested that tuatara at warmer sites may benefit from increased opportunities for emergence and the attainment of preferred body temperatures throughout the year, though a higher frequency of restrictive air temperatures over summer may also reduce emergence opportunities. Experimental work showed that warmer air temperatures may enhance learning in tuatara, which could improve their ability to cope with challenging environments under climate change. However, body size was also an influential component of learning ability and further research is needed to build on these initial findings. I conclude that tuatara may experience overall benefits from further translocations to warmer sites and warming climates at currently cooler sites, which suggests that other cold-adapted reptiles with similar thermal tolerances may also see initial benefits under climate warming, though further monitoring is required to determine longer-term translocation success. Equally, while warmer air temperatures were not found to be detrimental to tuatara, they still pose a risk to population viability and further work is required on the impacts of associated abiotic factors like drought, and how populations of this long-lived species may be affected if and when climate warming exceeds the upper temperature rise of ~5°C predicted by the 2100s.</p>

<p>Anthropogenic climate change is progressing at a rate unprecedented in the past 65 million years and is a significant conservation concern. The associated biotic and abiotic impacts are expected to have substantial effects on global biodiversity, with some species potentially more vulnerable than others. The tuatara (Sphenodon punctatus) is a New Zealand endemic reptile and of particular interest as it is a slowly reproducing, range-restricted, cold-adapted ectotherm with temperature-dependent sex determination. Consequently, tuatara could be particularly vulnerable to rising air temperatures and conservation translocations have been key components of tuatara conservation efforts. Knowledge of how the tuatara might be affected by warmer climates will help inform where future conservation efforts are best directed, practices to avoid and which sites might be most suitable for the establishment of populations. The translocation of 176 adult tuatara in October 2012 from Stephens Island in New Zealand’s Cook Strait to four latitudinally distant North Island sites offered the opportunity to study the responses of tuatara in a range of environments. The comparatively warmer, drier climates of several sites provided surrogates for temporal climate change, enabling an assessment of how a warming climate might impact tuatara, and how they might respond. Using field observations, laboratory analysis and controlled experiments I investigated the short-term success of the translocations, the influence of translocation and climate on tuatara enteric bacterial communities and parasites, as well as how warmer climates might influence nocturnal activity, thermoregulatory opportunities and learning ability. I found several translocated populations to be progressing favourably, and found evidence that tuatara may exhibit enhanced growth at warmer, less densely-populated sites, suggesting that further translocations to lower latitude sites might be a viable conservation strategy. However, high population density at one translocation site was a concern and management recommendations were made to enable the dispersal of individuals. I detected Salmonella Saintpaul for the first time in a live tuatara, Campylobacter spp. was identified as a likely common commensal organism, and no measurable impact of translocation or climate on bacterial prevalence was observed, suggesting no substantial risk of climate warming to the susceptibility of tuatara to these bacteria. Tick populations were negatively impacted by translocation-associated factors following release but subsequently recovered at most sites and mites were not found on any translocated tuatara. Diurnal and nocturnal activities were positively influenced by air temperature, up to an upper threshold, and assessment of the site-specific thermal climates suggested that tuatara at warmer sites may benefit from increased opportunities for emergence and the attainment of preferred body temperatures throughout the year, though a higher frequency of restrictive air temperatures over summer may also reduce emergence opportunities. Experimental work showed that warmer air temperatures may enhance learning in tuatara, which could improve their ability to cope with challenging environments under climate change. However, body size was also an influential component of learning ability and further research is needed to build on these initial findings. I conclude that tuatara may experience overall benefits from further translocations to warmer sites and warming climates at currently cooler sites, which suggests that other cold-adapted reptiles with similar thermal tolerances may also see initial benefits under climate warming, though further monitoring is required to determine longer-term translocation success. Equally, while warmer air temperatures were not found to be detrimental to tuatara, they still pose a risk to population viability and further work is required on the impacts of associated abiotic factors like drought, and how populations of this long-lived species may be affected if and when climate warming exceeds the upper temperature rise of ~5°C predicted by the 2100s.</p>

<p>Animals are regularly exposed to environmental, social and physiological challenges. In reaction to these challenges, individuals adjust their physiology and behaviour to maintain essential processes and optimise fitness. The most widely used indicators of physiological stress in vertebrates are glucocorticoid hormones (corticosterone (CORT) or cortisol), which are commonly referred to as ‘stress hormones’. The use of CORT as a tool to understand how individuals respond to natural or human-caused challenges is central to stress physiology research. Here, I investigated intrinsic and extrinsic factors associated with CORT secretion, CORT secretion as an indicator of physiological response to challenges/stressors, and the value of CORT secretion as conservation tool in an iconic protected reptile (the tuatara, Sphenodon punctatus).  A capture-restraint time series revealed a significant CORT response over a 24 h period in male and female (non-gravid and gravid) tuatara. Baseline CORT and the CORT response to capture and restraint (i.e. a standardised capture-stress protocol) were similar between sexes; however, female reproductive condition was correlated with CORT secretion in that higher baseline CORT and a lower CORT response were observed in gravid females. An observational study incorporating data across a wide range of ambient temperatures (from four island sites) confirmed that body temperature (Tb) is positively correlated with baseline CORT in gravid females only, and revealed a positive correlation between the CORT response and higher Tb in all adults. A supporting experimental study showed that acute ambient temperature increase (in which mean Tb reached 21.4±0.4°C) elicits a significant CORT response to capture-restraint in gravid females. These results confirmed that gravid females are not secreting CORT maximally during nesting, but actively modulate secretion. An inter-island comparison of CORT secretion (for four populations) revealed that baseline CORT secretion was similar among populations during the non-breeding and breeding seasons; however, the CORT response to capture-restraint varied significantly among populations. Inter-population variation in testosterone (T) was observed in males (but not females) and was positively linked with increased baseline CORT from the non-breeding season to the breeding season, suggesting male reproductive activity may drive seasonal change of baseline CORT. Significant correlations were observed between the CORT response to capture-restraint (but not baseline CORT) and habitat elements of latitude, tuatara density and seabird abundance and 2) demogenetic factors of sex ratio and genetic diversity. The measurement of CORT as a physiological monitoring tool indicated that short- and long- term dynamics of CORT secretion in tuatara are not altered through translocation to a new island, as the acute CORT response remained stable throughout exposure to cumulative stressors and long-term dynamics of CORT secretion in translocated populations simultaneously mirrored those in source populations.  These findings deliver the most detailed study of CORT secretion patterns in tuatara to date. Moreover, as the first study to apply CORT secretion data as an conservation physiology monitoring tool in tuatara, these results serve as a baseline reference for future research and monitoring of conservation efforts.</p>

<p>Animals are regularly exposed to environmental, social and physiological challenges. In reaction to these challenges, individuals adjust their physiology and behaviour to maintain essential processes and optimise fitness. The most widely used indicators of physiological stress in vertebrates are glucocorticoid hormones (corticosterone (CORT) or cortisol), which are commonly referred to as ‘stress hormones’. The use of CORT as a tool to understand how individuals respond to natural or human-caused challenges is central to stress physiology research. Here, I investigated intrinsic and extrinsic factors associated with CORT secretion, CORT secretion as an indicator of physiological response to challenges/stressors, and the value of CORT secretion as conservation tool in an iconic protected reptile (the tuatara, Sphenodon punctatus).  A capture-restraint time series revealed a significant CORT response over a 24 h period in male and female (non-gravid and gravid) tuatara. Baseline CORT and the CORT response to capture and restraint (i.e. a standardised capture-stress protocol) were similar between sexes; however, female reproductive condition was correlated with CORT secretion in that higher baseline CORT and a lower CORT response were observed in gravid females. An observational study incorporating data across a wide range of ambient temperatures (from four island sites) confirmed that body temperature (Tb) is positively correlated with baseline CORT in gravid females only, and revealed a positive correlation between the CORT response and higher Tb in all adults. A supporting experimental study showed that acute ambient temperature increase (in which mean Tb reached 21.4±0.4°C) elicits a significant CORT response to capture-restraint in gravid females. These results confirmed that gravid females are not secreting CORT maximally during nesting, but actively modulate secretion. An inter-island comparison of CORT secretion (for four populations) revealed that baseline CORT secretion was similar among populations during the non-breeding and breeding seasons; however, the CORT response to capture-restraint varied significantly among populations. Inter-population variation in testosterone (T) was observed in males (but not females) and was positively linked with increased baseline CORT from the non-breeding season to the breeding season, suggesting male reproductive activity may drive seasonal change of baseline CORT. Significant correlations were observed between the CORT response to capture-restraint (but not baseline CORT) and habitat elements of latitude, tuatara density and seabird abundance and 2) demogenetic factors of sex ratio and genetic diversity. The measurement of CORT as a physiological monitoring tool indicated that short- and long- term dynamics of CORT secretion in tuatara are not altered through translocation to a new island, as the acute CORT response remained stable throughout exposure to cumulative stressors and long-term dynamics of CORT secretion in translocated populations simultaneously mirrored those in source populations.  These findings deliver the most detailed study of CORT secretion patterns in tuatara to date. Moreover, as the first study to apply CORT secretion data as an conservation physiology monitoring tool in tuatara, these results serve as a baseline reference for future research and monitoring of conservation efforts.</p>

<p>Successful conservation of terrestrial biodiversity requires understanding and predicting the impacts of rapid climate warming on the suitability of both current and potential future habitats. Most predictions of range shifts and other population-scale effects of climate change rely to some extent on statistical links between a species' known geographical distribution and the suite of environmental conditions experienced within that space. However, species' responses to climate change are likely to be more complex than can be represented by the projection of current species-environment relationships into unknown environments. An important goal in biodiversity conservation is the development of quantitative tools with which to assess habitat suitability independently of distributions.  In populations of oviparous species, climate change and habitat modification may have distinct effects on different life stages. Temperatures that are well within the thermal tolerance range of adults, for example, may affect embryonic development rates, hatching phenology, or offspring survival and phenotype. I examined how environmental variation may affect the thermal suitability of habitat for facilitating embryonic development and maintaining balanced sex ratios in tuatara (Sphenodon punctatus), an endemic New Zealand reptile with temperature-dependent sex determination (TSD). Once widespread throughout New Zealand, populations are now restricted to offshore islands and fenced mainland sanctuaries, though establishment of additional populations via translocation is ongoing. Due to intensive conservation efforts, tuatara are not classified as an endangered species, but, like other species in which hatchling sex is determined by the incubation environment, populations are potentially at risk from the detrimental effects of sex-ratio bias.  I conducted two seasons of field work on the island of Takapourewa to quantify the relationship between rapid vegetation succession and selection of nesting areas. I then used a variety of predictive models to link data on nesting behaviour collected in the field with the microclimate conditions experienced by nesting female tuatara and developing embryos. Using mechanistically modelled soil temperature data, I generated predictions of incubation temperatures, offspring sex ratios, and hatching dates for two populations of tuatara on environmentally distinct islands, Takapourewa and Hauturu, under current and projected future climate scenarios. Finally, I classified the thermal suitability of sites on Hauturu for facilitating successful embryonic development and created geospatial surfaces defining suitable nesting locations adjacent to tuatara habitats.  Offspring sex ratios on both islands are unlikely to become male-biased if the magnitude of climate warming observed over the next century more closely matches the minimum, rather than the maximum, projected warming scenario. On Takapourewa, the timing of nesting will be critical in determining whether sex ratios become male-biased under a scenario of maximum climate warming. Earlier nesting may also lead to shifts in hatching phenology under either scenario of climate warming. Warmer annual temperatures on Hauturu are more likely to lead to heavily male-biased offspring sex ratios under the maximum warming scenario. Female tuatara on Hauturu do not need to travel away from their current habitats to locate suitable nesting sites. Monitoring the population to quantify nesting behaviour on the island will be important for determining whether females' choices of incubation microclimates can compensate for the sex ratio-biasing effects of climate change.</p>

<p>Successful conservation of terrestrial biodiversity requires understanding and predicting the impacts of rapid climate warming on the suitability of both current and potential future habitats. Most predictions of range shifts and other population-scale effects of climate change rely to some extent on statistical links between a species' known geographical distribution and the suite of environmental conditions experienced within that space. However, species' responses to climate change are likely to be more complex than can be represented by the projection of current species-environment relationships into unknown environments. An important goal in biodiversity conservation is the development of quantitative tools with which to assess habitat suitability independently of distributions.  In populations of oviparous species, climate change and habitat modification may have distinct effects on different life stages. Temperatures that are well within the thermal tolerance range of adults, for example, may affect embryonic development rates, hatching phenology, or offspring survival and phenotype. I examined how environmental variation may affect the thermal suitability of habitat for facilitating embryonic development and maintaining balanced sex ratios in tuatara (Sphenodon punctatus), an endemic New Zealand reptile with temperature-dependent sex determination (TSD). Once widespread throughout New Zealand, populations are now restricted to offshore islands and fenced mainland sanctuaries, though establishment of additional populations via translocation is ongoing. Due to intensive conservation efforts, tuatara are not classified as an endangered species, but, like other species in which hatchling sex is determined by the incubation environment, populations are potentially at risk from the detrimental effects of sex-ratio bias.  I conducted two seasons of field work on the island of Takapourewa to quantify the relationship between rapid vegetation succession and selection of nesting areas. I then used a variety of predictive models to link data on nesting behaviour collected in the field with the microclimate conditions experienced by nesting female tuatara and developing embryos. Using mechanistically modelled soil temperature data, I generated predictions of incubation temperatures, offspring sex ratios, and hatching dates for two populations of tuatara on environmentally distinct islands, Takapourewa and Hauturu, under current and projected future climate scenarios. Finally, I classified the thermal suitability of sites on Hauturu for facilitating successful embryonic development and created geospatial surfaces defining suitable nesting locations adjacent to tuatara habitats.  Offspring sex ratios on both islands are unlikely to become male-biased if the magnitude of climate warming observed over the next century more closely matches the minimum, rather than the maximum, projected warming scenario. On Takapourewa, the timing of nesting will be critical in determining whether sex ratios become male-biased under a scenario of maximum climate warming. Earlier nesting may also lead to shifts in hatching phenology under either scenario of climate warming. Warmer annual temperatures on Hauturu are more likely to lead to heavily male-biased offspring sex ratios under the maximum warming scenario. Female tuatara on Hauturu do not need to travel away from their current habitats to locate suitable nesting sites. Monitoring the population to quantify nesting behaviour on the island will be important for determining whether females' choices of incubation microclimates can compensate for the sex ratio-biasing effects of climate change.</p>

<p>Intraspecific competition is of importance in the wild and captivity, as the interaction among individuals for resources can affect growth, survival, and ultimately fitness. Tuatara, Sphenodon punctatus, are endemic to New Zealand and the sole representatives of the reptile Order Rhynchocephalia, and their recovery plan outlines the importance of head-started individuals to supplement existing populations and provide stock to start new populations. Head-starting is a widespread conservation tool for raising juveniles in captivity prior to release in the wild, with the aim of reducing juvenile mortality and providing populations with more individuals. However, mortality differs between sexes and juvenile tuatara show enormous variation in size in captivity. I investigated aggression and competition for space and food in a tuatara head-starting facility to determine whether intraspecific competition may affect mortality and growth. Pairs of one-year-old tuatara, S. punctatus, were chosen according to sex and relative size, e.g. a big male and a small female or two similar sized females. Seven scenarios were replicated four times with different pairs. Behaviour (including two feeding trials) was recorded over a six day period via security cameras and direct observations. The number of aggressive conflicts differed among scenarios, and male-male dyads were significantly more aggressive than female-female dyads. Dominance hierarchies were established in 18 of 28 experimental pairs, with bigger animals being dominant. Conflicts include chasing, biting or colliding at full speed. One year old juveniles did not compete for space. They did not use space exclusively, but stopped clustering and had developed aggressive behaviour, suggesting that they are not territorial yet but in an early stage of transition towards territoriality as seen in older juveniles and adults. Space use and avoidance in space and time did not differ among social scenarios and the latter were negligible, but they marked a novel enclosure with urine and faeces. Juveniles competed directly and indirectly for food. Dominant individuals were likely to secure more food than submissive individuals. Females acquired less food when paired with males of bigger or similar size, and acquired about equal shares when paired with a smaller male. While bigger males acquired slightly more food when paired with smaller males, this was not the case in differently sized females. Interference behaviours such as chasing and food stealing were mostly directed from bigger towards smaller individuals. Captive group housing has consequences for competition and aggression, and may directly influence survival. As juvenile tuatara mortality is female-biased, and aggression against females in bigger male-biased groups common, I recommend keeping sexes separate, and assorting groups by size with more spacious enclosures for male groups. These modifications should improve health and numbers of juveniles for release, improve recruitment into the reproductive adult population, and ultimately create more successful head-starting facilities.</p>

<p>Population viability for small, isolated populations is determined by many factors, particularly demographic stochasticity. Coexistence of communities is promoted through resource partitioning, particularly if species share similar niche requirements. Demographic characteristics, long-term trends and patterns of partitioning were investigated for two reptile species: tuatara (Sphenodon punctatus) and Duvaucel's gecko (Hoplodactylus duvaucelii), using mark recapture techniques on North Brother Island, New Zealand. Capture time and location were recorded as well as snout-vent length, mass and sex of individuals. Adult population size, sex ratio, survival and recapture probability for both species were estimated. Intervention will be needed to prevent population collapse for tuatara, as the population is male-biased (3.24 males: 1 female), with sub-adults exhibiting a stronger bias (4.1 males: 1 female). The total population size is estimated at 390-437 adults, with high adult survival (95%). The Duvaucel's gecko population is stable enough to be harvested for translocation, as the population was estimated at 583-677 adults, with an even sex ratio. Adult survival was high (92%) and longevity is at least 43-50 years. Patterns in partitioning suggest tuatara are excluding Duvaucel's gecko as tuatara occupy vegetated areas and few animals were caught at the same time in the same place as a member of the other species (~10%). Long-term site fidelity appears to occur in both species as the majority of animals were captured previously within 10m (tuatara) or 15m (Duvaucel's gecko) of their 2008 location, and travelled less than 2m per year on average. Tuatara show an overall decline in body condition since 1957, which is more rapid in females, and may be related to intraspecific interactions and density-dependent effects. Gecko body condition is not declining, suggesting no negative effects at the population level are occurring as a result of competitive exclusion. This study indicates that characteristics that have implications for population viability have the capacity to differ, even for species with similar niche requirements occupying the same habitat, and supports the considerable value of long-term monitoring.</p>

<p>Population viability for small, isolated populations is determined by many factors, particularly demographic stochasticity. Coexistence of communities is promoted through resource partitioning, particularly if species share similar niche requirements. Demographic characteristics, long-term trends and patterns of partitioning were investigated for two reptile species: tuatara (Sphenodon punctatus) and Duvaucel's gecko (Hoplodactylus duvaucelii), using mark recapture techniques on North Brother Island, New Zealand. Capture time and location were recorded as well as snout-vent length, mass and sex of individuals. Adult population size, sex ratio, survival and recapture probability for both species were estimated. Intervention will be needed to prevent population collapse for tuatara, as the population is male-biased (3.24 males: 1 female), with sub-adults exhibiting a stronger bias (4.1 males: 1 female). The total population size is estimated at 390-437 adults, with high adult survival (95%). The Duvaucel's gecko population is stable enough to be harvested for translocation, as the population was estimated at 583-677 adults, with an even sex ratio. Adult survival was high (92%) and longevity is at least 43-50 years. Patterns in partitioning suggest tuatara are excluding Duvaucel's gecko as tuatara occupy vegetated areas and few animals were caught at the same time in the same place as a member of the other species (~10%). Long-term site fidelity appears to occur in both species as the majority of animals were captured previously within 10m (tuatara) or 15m (Duvaucel's gecko) of their 2008 location, and travelled less than 2m per year on average. Tuatara show an overall decline in body condition since 1957, which is more rapid in females, and may be related to intraspecific interactions and density-dependent effects. Gecko body condition is not declining, suggesting no negative effects at the population level are occurring as a result of competitive exclusion. This study indicates that characteristics that have implications for population viability have the capacity to differ, even for species with similar niche requirements occupying the same habitat, and supports the considerable value of long-term monitoring.</p>

<p>Intraspecific competition is of importance in the wild and captivity, as the interaction among individuals for resources can affect growth, survival, and ultimately fitness. Tuatara, Sphenodon punctatus, are endemic to New Zealand and the sole representatives of the reptile Order Rhynchocephalia, and their recovery plan outlines the importance of head-started individuals to supplement existing populations and provide stock to start new populations. Head-starting is a widespread conservation tool for raising juveniles in captivity prior to release in the wild, with the aim of reducing juvenile mortality and providing populations with more individuals. However, mortality differs between sexes and juvenile tuatara show enormous variation in size in captivity. I investigated aggression and competition for space and food in a tuatara head-starting facility to determine whether intraspecific competition may affect mortality and growth. Pairs of one-year-old tuatara, S. punctatus, were chosen according to sex and relative size, e.g. a big male and a small female or two similar sized females. Seven scenarios were replicated four times with different pairs. Behaviour (including two feeding trials) was recorded over a six day period via security cameras and direct observations. The number of aggressive conflicts differed among scenarios, and male-male dyads were significantly more aggressive than female-female dyads. Dominance hierarchies were established in 18 of 28 experimental pairs, with bigger animals being dominant. Conflicts include chasing, biting or colliding at full speed. One year old juveniles did not compete for space. They did not use space exclusively, but stopped clustering and had developed aggressive behaviour, suggesting that they are not territorial yet but in an early stage of transition towards territoriality as seen in older juveniles and adults. Space use and avoidance in space and time did not differ among social scenarios and the latter were negligible, but they marked a novel enclosure with urine and faeces. Juveniles competed directly and indirectly for food. Dominant individuals were likely to secure more food than submissive individuals. Females acquired less food when paired with males of bigger or similar size, and acquired about equal shares when paired with a smaller male. While bigger males acquired slightly more food when paired with smaller males, this was not the case in differently sized females. Interference behaviours such as chasing and food stealing were mostly directed from bigger towards smaller individuals. Captive group housing has consequences for competition and aggression, and may directly influence survival. As juvenile tuatara mortality is female-biased, and aggression against females in bigger male-biased groups common, I recommend keeping sexes separate, and assorting groups by size with more spacious enclosures for male groups. These modifications should improve health and numbers of juveniles for release, improve recruitment into the reproductive adult population, and ultimately create more successful head-starting facilities.</p>