Anthropogenic and climatic factors interact to influence reproductive timing and effort.

Reproduction, although absolutely essential to a species' persistence, is in itself challenging. As anthropogenic change increasingly affects every landscape on Earth, it is critical to understand how specific pressures impact the reproductive efforts of individuals, which directly contribute to the success or failure of populations. However, organisms rarely encounter a single burden at a time, and the interactions of environmental challenges can have compounding effects. Understanding environmental and physiological pressures is difficult because they are often context-dependent and not generalizable, but long-term monitoring across variable landscapes and weather patterns can improve our understanding of these complex interactions. We tested the effects of urbanization, climate, and individual condition on the reproductive investment of wild side-blotched lizards (Uta stansburiana) by measuring physiological/reproductive metrics from six populations in urban and rural areas over six consecutive years of variable precipitation. We observed that reproductive stage affected body condition, corticosterone concentration, and oxidative stress. We also observed that reproductive patterns differed between urban and rural populations depending on rainfall, with rural animals increasing reproductive investment during rainier years compared to urban conspecifics, and that reproductive decisions appeared to occur early in the reproductive process. These results demonstrate the plastic nature of a generalist species optimizing lifetime fitness under varying conditions.

Blood chemistry and biliverdin differ according to reproduction and tourism in a free-living lizard.

Changes in the physiological health of species are an essential indicator of changing conditions and environmental challenges. Reponses to environmental challenges can often induce stress, influence physiology, and change metabolism in organisms. Here we tested blood chemistry parameters indicative of stress and metabolic activity using an i-STAT point-of-care blood analyzer in seven populations of free-ranging rock iguanas exposed to varying levels of tourism and supplemental feeding. We found significant differences in blood chemistry (glucose, oxygen, carbon dioxide, hematocrit, hemoglobin, calcium, potassium, and biliverdin levels) among populations exposed to varying levels of tourism, and some variation between sexes and reproductive states. However, different variables are not directly related to one another, suggesting that the causal physiological pathways driving tourism-induced differences are influenced by mechanisms that are not detected by common analyses of blood chemistry. Future work should investigate upstream regulators of these factors affected by tourism. Regardless, these blood metrics are known to be both stress sensitive and related to metabolic activity, suggesting that exposure to tourism and associated supplemental feeding by tourists are generally driven by stress-related changes in blood chemistry, biliverdin, and metabolism.

Complex tourism and season interactions contribute to disparate physiologies in an endangered rock iguana.

To promote survival and fitness, organisms use a suite of physiological systems to respond to both predictable and unpredictable changes in the environment. These physiological responses are also influenced by changes in life history state. The continued activation of physiological systems stemming from persistent environmental perturbations enable animals to cope with these challenges but may over time lead to significant effects on the health of wildlife. In the present study, we tested how varying environmental perturbations driven by tourism and associated supplemental feeding affects the energetics, corticosterone and immunity of six discrete populations of the northern Bahamian rock iguana (Cyclura cychlura inornata and Cyclura cychlura figginsi). We studied populations within and outside the reproductive season and quantified tourist numbers during sample collection. Specifically, we measured clutch size, body condition, plasma energy metabolites, reactive oxygen species, baseline corticosterone concentrations and immune function of male and female iguanas from each population to address whether (i) disparate physiologies are emerging across a gradient of tourism and feeding, (ii) both subspecies respond similarly and (iii) responses vary with season/reproductive condition. We found significant effects of tourism level, season and their interaction on the physiology of both C. c. inornata and C. c. figginsi, supporting the idea that tourism is leading to the divergence of phenotypes. Specifically, we found elevated plasma energy metabolites, oxidative stress and a measure of innate immunity (bactericidal ability), but reduced corticosterone concentrations with increasing tourism in both subspecies of rock iguanas. These physiological metrics differ according to the level of tourism in both subspecies and persist across seasons despite variation with natural seasonal and reproductive changes. These findings suggest that anthropogenic disturbance results in disparate physiologies in northern Bahamian rock iguanas.

Glucose tolerance of iguanas is affected by high-sugar diets in the lab and supplemental feeding by ecotourists in the wild.

There is great interspecific variation in the nutritional composition of natural diets, and the varied nutritional content is physiologically tolerated because of evolutionarily based balances between diet composition and processing ability. However, as a result of landscape change and human exposure, unnatural diets are becoming widespread among wildlife without the necessary time for evolutionary matching between the diet and its processing. We tested how a controlled, unnatural high glucose diet affects glucose tolerance using captive green iguanas, and we performed similar glucose tolerance tests on wild Northern Bahamian rock iguanas that are either frequently fed grapes by tourists or experience no such supplementation. We evaluated both short and longer-term blood glucose responses and corticosterone (CORT) concentrations as changes have been associated with altered diets. Experimental glucose supplementation in the laboratory and tourist feeding in the wild both significantly affected glucose metabolism. When iguanas received a glucose-rich diet, we found greater acute increases in blood glucose following a glucose challenge. Relative to unfed iguanas, tourist-fed iguanas had significantly lower baseline CORT, higher baseline blood glucose, and slower returns to baseline glucose levels following a glucose challenge. Therefore, unnatural consumption of high amounts of glucose alters glucose metabolism in laboratory iguanas with short-term glucose treatment and free-living iguanas exposed to long-term feeding by tourists. Based on these results and the increasing prevalence of anthropogenically altered wildlife diets, the consequences of dietary changes on glucose metabolism should be further investigated across species, as such changes in glucose metabolism have health consequences in humans (e.g. diabetes).

Investigating the relationship between corticosterone and glucose in a reptile.

The glucocorticoid hormone corticosterone (CORT) has classically been used in ecophysiological studies as a proxy for stress and energy mobilization, but rarely are CORT and the energy metabolites themselves concurrently measured. To examine CORT's role in mobilizing glucose in a wild reptile, we conducted two studies. The first study measured natural baseline and stress-induced blood-borne CORT and glucose levels in snakes during spring emergence and again when snakes return to the denning sites in autumn. The second study manipulated the hypothalamic-pituitary-adrenal (HPA) axis in male snakes in the autumn by taking a baseline blood sample, then subjecting individuals to one of five treatments (no injection, saline, CORT, adrenocorticotropin hormone and metyrapone). Subsequent samples were taken at 30 and 60 min. In both studies, we found that glucose levels do increase with acute stress, but that the relationship was not directly related to CORT elevation. In the second study, we found that none of the HPA axis manipulations directly affected blood glucose levels, further indicating that CORT may play a complex but not direct role in glucose mobilization in snakes. This study highlights the need for testing mechanisms in wild organisms by combining in situ observations with manipulative studies.

Energetic investment associated with vitellogenesis induces an oxidative cost of reproduction.

Oxidative stress is a potential cost of reproduction, but conclusive evidence for this relationship is lacking. The goal of this study was to serially assess across a seasonal gradient the relationship between reproduction, circulating plasma energy metabolites and oxidative state. Here, we examine a study animal ideally suited to test for the oxidative costs of reproduction: the Allen Cays Rock Iguana. Female rock iguanas reproduce at varying frequencies, often skipping years, allowing for a comparison between reproductive and non-reproductive females during the same narrow, annual breeding season. This feature of iguana life history enabled us to address not just sex and seasonal differences in physiology, but also potential oxidative costs of reproduction in females. Male and female iguanas were sampled during the early (vitellogenic), late (gravid) and post-reproductive seasons. Ultrasound examinations were performed on females to quantify reproductive investment, and blood samples were collected for physiology assays, which included reactive oxygen metabolites (d-ROMs), antioxidants, triglycerides, free glycerol and glucose. The early reproductive season was characterized by significant increases in reproductive female's triglycerides, free glycerol and oxidative stress compared to late and post-reproductive periods and non-reproductive females and males during all sampling periods. Antioxidants were significantly elevated during the early reproductive season for reproductive females, non-reproductive females and males when compared to late and post-season. Follicle number in early reproductive females was positively related to d-ROMs, triglycerides and free glycerol, negatively related to antioxidants and showed no relationship with glucose. Measures of oxidative stress, d-ROMs and oxidative index were positively correlated with circulating levels of the lipid metabolite free glycerol during the early reproductive period, but this relationship weakened in the late season and disappeared in the post-season. Broadly, this study supports the hypothesis that the relationship between reproduction and oxidative stress is driven by energy investment, being greatest during early reproduction when vitellogenesis is occurring.

Town and Country Reptiles: A Review of Reptilian Responses to Urbanization.

The majority of the world population is now inhabiting urban areas, and with staggering population growth, urbanization is also increasing. While the work studying the effects of changing landscapes and specific urban pressures on wildlife is beginning to amass, the majority of this work focuses on avian or mammalian species. However, the effects of urbanization likely vary substantially across taxonomic groups due to differences in habitat requirements and life history. The current article aims first to broaden the review of urban effects across reptilian species; second, to summarize the responses of reptilian fauna to specific urban features; and third, to assess the directionality of individual and population level responses to urbanization in reptile species. Based on our findings, urban research in reptilian taxa is lacking in the following areas: (1) investigating interactive or additive urban factors, (2) measuring multiple morphological, behavioral, and physiological endpoints within an animal, (3) linking individual to population-level responses, and (4) testing genetic/genomic differences across an urban environment as evidence for selective pressures.

Metabolic responses to different immune challenges and varying resource availability in the side-blotched lizard (Uta stansburiana).

The energetic cost of immunity depends on many factors, including the type of challenge, the timing of the response, and the state of the animal. We measured changes in the standard metabolic rates of side-blotched lizards (Uta stansburiana Baird and Girard, 1852) in response to different immune challenges and nutritional states. In the first experiment, lizards were randomly assigned to one of four treatments: lipopolysaccharide (LPS) injection (to stimulate the response to a pathogen), cutaneous biopsy (as a proxy to a superficial wound), both injection and biopsy, or neither (control). Four and five days later, we measured the standard metabolic rates of the lizards. In response to healing a cutaneous wound, lizards reduced metabolic rate and lost body mass. Healing rate was also inversely related to weight loss, but LPS had no effect on body mass or metabolic rate. In the second experiment, a new set of lizards were randomly assigned to a high-food or low-food diet and administered a cutaneous biopsy. As in the first experiment, we observed a reduction in metabolic rate after wounding; moreover, this decrease was positively correlated with the rate of healing. We observed higher rates of metabolism in lizards that ate more food, but food consumption was unrelated to the decrease in metabolic rate following the biopsy. These experiments demonstrate the dynamic nature of the immune response in response to immune challenge and the state of the organism.