Geographic variation in larval cold tolerance and exposure across the invasion front of a widely established forest insect.

Under global climate change, high and low temperature extremes can drive shifts in species distributions. Across the range of a species, thermal tolerance is based on acclimatization, plasticity, and may undergo selection, shaping resilience to temperature stress. In this study, we measured variation in cold temperature tolerance of early instar larvae of an invasive forest insect, Lymantria dispar dispar L. (Lepidoptera: Erebidae), using populations sourced from a range of climates within the current introduced range in the Eastern United States. We tested for population differences in chill coma recovery (CCR) by measuring recovery time following a period of exposure to a nonlethal cold temperature in 2 cold exposure experiments. A 3rd experiment quantified growth responses after CCR to evaluate sublethal effects. Our results indicate that cold tolerance is linked to regional climate, with individuals from populations sourced from colder climates recovering faster from chill coma. While this geographic gradient is seen in many species, detecting this pattern is notable for an introduced species founded from a single point-source introduction. We demonstrate that the cold temperatures used in our experiments occur in nature during cold spells after spring egg hatch, but impacts to growth and survival appear low. We expect that population differences in cold temperature performance manifest more from differences in temperature-dependent growth than acute exposure. Evaluating intraspecific variation in cold tolerance increases our understanding of the role of climatic gradients on the physiology of an invasive species, and contributes to tools for predicting further expansion.

Teaching biology students data exploration and visualization in a data-driven world.

As biologists accumulate or encounter increasingly large and complex data sets, our field creates the need for students to develop skills in data exploration and visualization. Many biology courses lack the time for students to develop the skills needed to parse complex datasets and visualize them appropriately. We developed a new upper-level undergraduate biology course to focused on data exploration and communication without requiring previous coding experience. We emphasized data visualization principles and best practices and taught students how to manage and visualize data via Tableau and R. We also explored scientific ethics, how to refute misinformation, and inequities that can occur in data collection and usage.

Growth and development of an invasive forest insect under current and future projected temperature regimes.

Temperature and its impact on fitness are fundamental for understanding range shifts and population dynamics under climate change. Geographic climate heterogeneity, behavioral and physiological plasticity, and thermal adaptation to local climates make predicting the responses of species to climate change complex. Using larvae from seven geographically distinct wild populations in the eastern United States of the non-native forest pest Lymantria dispar dispar (L.), we conducted a simulated reciprocal transplant experiment in environmental chambers using six custom temperature regimes representing contemporary conditions near the southern and northern extremes of the US invasion front and projections under two climate change scenarios for the year 2050. Larval growth and development rates increased with climate warming compared with current thermal regimes and tended to be greater for individuals originally sourced from southern rather than northern populations. Although increases in growth and development rates with warming varied somewhat by region of the source population, there was not strong evidence of local adaptation, southern populations tended to outperform those from northern populations in all thermal regimes. Our study demonstrates the utility of simulating thermal regimes under climate change in environmental chambers and emphasizes how the impacts from future increases in temperature can vary based on geographic differences in climate-related performance among populations.

R Markdown as a dynamic interface for teaching: Modules from math and biology classrooms.

Advancing technologies, including interactive tools, are changing classroom pedagogy across academia. Here, we discuss the R Markdown interface, which allows for the creation of partial or complete interactive classroom modules for courses using the R programming language. R Markdown files mix sections of R code with formatted text, including LaTeX, which are rendered together to form complete reports and documents. These features allow instructors to create classroom modules that guide students through concepts, while providing areas for coding and text response by students. Students can also learn to create their own reports for more independent assignments. After presenting the features and uses of R Markdown to enhance teaching and learning, we present examples of materials from two courses. In a Computational Modeling course for math students, we used R Markdown to guide students through exploring mathematical models to understand the principle of herd immunity. In a Data Visualization and Communication course for biology students, we used R Markdown for teaching the fundamentals of R programming and graphing, and for students to learn to create reproducible data investigations. Through these examples, we demonstrate the benefits of R Markdown as a dynamic teaching and learning tool.

Assessing drivers of localized invasive spread to inform large-scale management of a highly damaging insect pest.

Studies of biological invasions at the macroscale or across multiple scales can provide important insights for management, particularly when localized information about invasion dynamics or environmental contexts is unavailable. In this study, we performed a macroscale analysis of the roles of invasion drivers on the local scale dynamics of a high-profile pest, Lymantria dispar dispar L., with the purpose of improving the prioritization of vulnerable areas for treatment. Specifically, we assessed the relative effects of various anthropogenic and environmental variables on the establishment rate of 8010 quadrats at a localized scale (5 × 5 km) across the entire L. dispar transition zone (the area encompassing the leading population edge, currently from Minnesota to North Carolina). We calculated the number of years from first detection of L. dispar in a quadrat to the year when probability of establishment of L. dispar was greater than 99% (i.e., waiting time to establishment after first detection). To assess the effects of environmental and anthropogenic variables on each quadrat's waiting time to establishment, we performed linear mixed-effects regression models for the full transition zone and three subregions within the zone. Seasonal temperatures were found to be the primary drivers of local establishment rates. Winter temperatures had the strongest effects, especially in the northern parts of the transition zone. Furthermore, the effects of some factors on waiting times to establishment varied across subregions. Our findings contribute to identifying especially vulnerable areas to further L. dispar spread and informing region-specific criteria by invasion managers for the prioritization of areas for treatment.

Demographic consequences of changing body size in a terrestrial salamander.

Changes in climate can alter individual body size, and the resulting shifts in reproduction and survival are expected to impact population dynamics and viability. However, appropriate methods to account for size-dependent demographic changes are needed, especially in understudied yet threatened groups such as amphibians. We investigated individual- and population-level demographic effects of changes in body size for a terrestrial salamander using capture-mark-recapture data. For our analysis, we implemented an integral projection model parameterized with capture-recapture likelihood estimates from a Bayesian framework. Our study combines survival and growth data from a single dataset to quantify the influence of size on survival while including different sources of uncertainty around these parameters, demonstrating how selective forces can be studied in populations with limited data and incomplete recaptures. We found a strong dependency of the population growth rate on changes in individual size, mediated by potential changes in selection on mean body size and on maximum body size. Our approach of simultaneous parameter estimation can be extended across taxa to identify eco-evolutionary mechanisms acting on size-specific vital rates, and thus shaping population dynamics and viability.

Hurricane-induced demographic changes in a non-human primate population.

Major disturbance events can have large impacts on the demography and dynamics of animal populations. Hurricanes are one example of an extreme climatic event, predicted to increase in frequency due to climate change, and thus expected to be a considerable threat to population viability. However, little is understood about the underlying demographic mechanisms shaping population response following these extreme disturbances. Here, we analyse 45 years of the most comprehensive free-ranging non-human primate demographic dataset to determine the effects of major hurricanes on the variability and maintenance of long-term population fitness. For this, we use individual-level data to build matrix population models and perform perturbation analyses. Despite reductions in population growth rate mediated through reduced fertility, our study reveals a demographic buffering during hurricane years. As long as survival does not decrease, our study shows that hurricanes do not result in detrimental effects at the population level, demonstrating the unbalanced contribution of survival and fertility to population fitness in long-lived animal populations.

Unexpected spatial population ecology of a widespread terrestrial salamander near its southern range edge.

Under the current amphibian biodiversity crisis, common species provide an opportunity to measure population dynamics across a wide range of environmental conditions while examining the processes that determine abundance and structure geographical ranges. Studying species at their range limits also provides a window for understanding the dynamics expected in future environments under increasing climate change and human modification. We quantified patterns of seasonal activity, density and space use in the eastern red-backed salamander (Plethodon cinereus) near its southern range edge and compare the spatial ecology of this population to previous findings from the core of their range. This southern population shows the expected phenology of surface activity based on temperature limitations in warmer climates, yet maintains unexpectedly high densities and large home ranges during the active season. Our study suggests that ecological factors known to strongly affect amphibian populations (e.g. warm temperature and forest fragmentation) do not necessarily constrain this southern population. Our study highlights the utility of studying a common amphibian as a model system for investigating population processes in environments under strong selective pressure.

Evolutionary genomics of gypsy moth populations sampled along a latitudinal gradient.

The European gypsy moth (Lymantria dispar L.) was first introduced to Massachusetts in 1869 and within 150 years has spread throughout eastern North America. This large-scale invasion across a heterogeneous landscape allows examination of the genetic signatures of adaptation potentially associated with rapid geographical spread. We tested the hypothesis that spatially divergent natural selection has driven observed changes in three developmental traits that were measured in a common garden for 165 adult moths sampled from six populations across a latitudinal gradient covering the entirety of the range. We generated genotype data for 91,468 single nucleotide polymorphisms based on double digest restriction-site associated DNA sequencing and used these data to discover genome-wide associations for each trait, as well as to test for signatures of selection on the discovered architectures. Genetic structure across the introduced range of gypsy moth was low in magnitude (FST  = 0.069), with signatures of bottlenecks and spatial expansion apparent in the rare portion of the allele frequency spectrum. Results from applications of Bayesian sparse linear mixed models were consistent with the presumed polygenic architectures of each trait. Further analyses indicated spatially divergent natural selection acting on larval development time and pupal mass, with the linkage disequilibrium component of this test acting as the main driver of observed patterns. The populations most important for these signals were two range-edge populations established less than 30 generations ago. We discuss the importance of rapid polygenic adaptation to the ability of non-native species to invade novel environments.

Thermal Sensitivity of Gypsy Moth (Lepidoptera: Erebidae) During Larval and Pupal Development.

As global temperatures rise, thermal limits play an increasingly important role in determining the persistence and spread of invasive species. Gypsy moth (Lymantria dispar L. Lepidoptera: Erebidae) in North America provides an ideal system for studying the effect of high temperatures on invasive species performance. Here, we used fluctuating temperature regimes and exposed gypsy moth at specific points in development (first-fourth instar, pupa) to cycles of favorable (22-28°C) or high-temperature treatments (30-36°C, 32-38°C, 34-40°C) for either 2 or 7 d. We measured survival, growth, and prolonged effects of exposure on development time and pupal mass. Survival generally decreased as the experimental temperature treatment and duration of exposure increased for all instars and pupae, with a narrow threshold for lethal effects. In response to increasing temperature and magnified by longer exposure times, growth abruptly declined for third instars and development time increased for pupae. For those surviving the 2-d exposure treatment, development time to pupation increased for all instars, but we did not find consistent effects on final pupal mass. These negative effects of high temperature provide important data on the susceptibility of gypsy moth to heat at different points in development. This work improves our understanding of thermal limits to growth and development in gypsy moth and can aid in determining invasion potential under current and future climates.

Physiological responses to elevated temperature across the geographic range of a terrestrial salamander.

Widespread species often possess physiological mechanisms for coping with thermal heterogeneity, and uncovering these mechanisms provides insight into species' responses to climate change. The emergence of non-invasive corticosterone (CORT) assays allows us to rapidly assess physiological responses to environmental change on a large scale. We lack, however, a basic understanding of how temperature affects CORT, and whether temperature and CORT interactively affect performance. Here, we examined the effects of elevated temperature on CORT and whole-organism performance in a terrestrial salamander, Plethodon cinereus, across a latitudinal gradient. Using water-borne hormone assays, we found that raising ambient temperature from 15 to 25°C increased CORT release at a similar rate for salamanders from all sites. However, CORT release rates were higher overall in the warmest, southernmost site. Elevated temperatures also affected physiological performance, but the effects differed among sites. Ingestion rate increased in salamanders from the warmer sites but remained the same for those from cooler sites. Mass gain was reduced for most individuals, although this reduction was more dramatic in salamanders from the cooler sites. We also found a temperature-dependent relationship between CORT and food conversion efficiency (i.e. the amount of mass gained per unit food ingested). CORT was negatively related to food conversion efficiency at 25°C but was unrelated at 15°C. Thus, the energetic gains of elevated ingestion rates may be counteracted by elevated CORT release rates experienced by salamanders in warmer environments. By integrating multiple physiological metrics, we highlight the complex relationships between temperature and individual responses to warming climates.

Thermal Physiology and Developmental Plasticity of Pigmentation in the Harlequin Bug (Hemiptera: Pentatomidae).

Traits that promote the maintenance of body temperatures within an optimal range provide advantages to ectothermic species. Pigmentation plasticity is found in many insects and enhances thermoregulatory potential as increased melanization can result in greater heat retention. The thermal melanism hypothesis predicts that species with developmental plasticity will have darker pigmentation in colder environments, which can be an important adaptation for temperate species experiencing seasonal variation in climate. The harlequin bug (Murgantia histrionica, Hemiptera: Pentatomidae, Hahn 1834) is a widespread invasive crop pest with variable patterning where developmental plasticity in melanization could affect performance. To investigate the impact of temperature and photoperiod on melanization and size, nymphs were reared under two temperatures and two photoperiods simulating summer and fall seasons. The size and degree of melanization of adults were quantified using digital imagery. To assess the effect of coloration on the amount of heat absorption, we monitored the temperature of adults in a heating experiment. Overall, our results supported the thermal melanism hypothesis and temperature had a comparatively larger effect on coloration and size than photoperiod. When heated, the body temperature of individuals with darker pigmentation increased more relative to the ambient air temperature than individuals with lighter pigmentation. These results suggest that colder temperatures experienced late in the season can induce developmental plasticity for a phenotype that improves thermoregulation in this species. Our work highlights environmental signals and consequences for individual performance due to thermal melanism in a common invasive species, where capacity to respond to changing environments is likely contributing to its spread.

Geographic Variation in Larval Metabolic Rate Between Northern and Southern Populations of the Invasive Gypsy Moth.

Thermal regimes can diverge considerably across the geographic range of a species, and accordingly, populations can vary in their response to changing environmental conditions. Both local adaptation and acclimatization are important mechanisms for ectotherms to maintain homeostasis as environments become thermally stressful, which organisms often experience at their geographic range limits. The spatial spread of the gypsy moth (Lymantria dispar L.) (Lepidoptera: Erebidae) after introduction to North America provides an exemplary system for studying population variation in physiological traits given the gradient of climates encompassed by its current invasive range. This study quantifies differences in resting metabolic rate (RMR) across temperature for four populations of gypsy moth, two from the northern and two from southern regions of their introduced range in North America. Gypsy moth larvae were reared at high and low thermal regimes, and then metabolic activity was monitored at four temperatures using stop-flow respirometry to test for an acclimation response. For all populations, there was a significant increase in RMR as respirometry test temperature increased. Contrary to our expectations, we did not find evidence for metabolic adaptation to colder environments based on our comparisons between northern and southern populations. We also found no evidence for an acclimation response of RMR to rearing temperature for three of the four pairwise comparisons examined. Understanding the thermal sensitivity of metabolic rate in gypsy moth, and understanding the potential for changes in physiology at range extremes, is critical for estimating continued spatial spread of this invasive species both under current and potential future climatic constraints.

Novel insights on population and range edge dynamics using an unparalleled spatiotemporal record of species invasion.

Quantifying the complex spatial dynamics taking place at range edges is critical for understanding future distributions of species, yet very few systems have sufficient data or the spatial resolution to empirically test these dynamics. This paper reviews how data from a large-scale pest management programme have provided important contributions to the fields of population dynamics and invasion biology. The invasion of gypsy moth (Lymantria dispar) is well-documented from its introduction near Boston, Massachusetts USA in 1869 to its current extent of over 900,000 km2 in Eastern North America. Over the past two decades, the USDA Forest Service Slow the Spread (STS) programme for managing the future spread of gypsy moth has produced unrivalled spatiotemporal data across the invasion front. The STS programme annually deploys a grid of 60,000-100,000 pheromone-baited traps, currently extending from Minnesota to North Carolina. The data from this programme have provided the foundation for investigations of complex population dynamics and the ability to examine ecological hypotheses previously untestable outside of theoretical venues, particularly regarding invasive spread and Allee effects. This system provides empirical data on the importance of long-distance dispersal and time-lags on population establishment and spatial spread. Studies showing high rates of spatiotemporal variation of the range edge, from rapid spread to border stasis and even retraction, highlight future opportunities to test mechanisms that influence both invasive and native species ranges. The STS trap data have also created a unique opportunity to study low-density population dynamics and quantify Allee effects with empirical data. Notable contributions include evidence for spatiotemporal variation in Allee effects, demonstrating empirical links between Allee effects and spatial spread, and testing mechanisms of population persistence and growth rates at range edges. There remain several outstanding questions in spatial ecology and population biology that can be tested within this system, such as the scaling of local ecological processes to large-scale dynamics across landscapes. The gypsy moth is an ideal model of how important ecological questions can be answered by thinking more broadly about monitoring data.

The association between male-biased sex ratio and indicators of stress in red-spotted newts.

In populations with a male-biased operational sex ratio, coercive mating by males can have fitness consequences for females. One component of reduced fitness for females in populations with a male-biased OSR may be greater activation of the stress response, resulting in higher corticosterone release rates (CORT; a glucocorticoid stress hormone in amphibians). We test the hypothesis that a male-biased sex ratio affects female activity and release rates of CORT and testosterone (T) in male and female red-spotted newts (Notophthalmus viridescens). First, we evaluated if chemical cues from a male-biased sex ratio affect activity and CORT release rates in females. We predicted that females exposed to chemical cues of three males would be less active and have higher CORT release rates than those exposed to chemical cues of one male. Second, we measured CORT release rates of red-spotted newts in field enclosures with either a male-biased or a female-biased sex ratio. We predicted that females in the male-biased treatment would have higher CORT and T release rates than those in a female-biased treatment, owing to higher levels of male harassment. We also predicted that males would have higher CORT and T release rates in male-biased treatments due to higher levels of male-male competition. Females were not less active in response to chemical cues from more males over fewer males, but there was a positive relationship between female activity and CORT when they were exposed to the cues of three males. We also found that females, but not males, in the male-biased sex ratio treatment had higher CORT and T release rates than those in the female-biased treatment. Our results support the hypothesis that a male-biased sex ratio leads to a higher stress response, which may underlie the observed decrease in immune function and body condition in previous work exposing female red-spotted newts to a male-biased sex ratio. This study furthers our understanding of the mechanistic basis for costs associated with a male-biased sex ratio in a pond-breeding amphibian.

Performance of Wild and Laboratory-Reared Gypsy Moth (Lepidoptera: Erebidae): A Comparison between Foliage and Artificial Diet.

The effects of long-term mass rearing of laboratory insects on ecologically relevant traits is an important consideration when applying research conclusions to wild populations or developing management strategies. Laboratory strains of the gypsy moth, Lymantria dispar (L.), an invasive forest pest in North America, have been continuously reared since 1967. Selection on these strains has enhanced a variety of traits, resulting in faster development, shorter diapause, and greater fecundity. As in many mass-reared insects, laboratory strains of the gypsy moth are also reared exclusively on artificial diets that lack much of the phytochemical and nutritional complexity associated with natural foliage. We tested for differences in growth and development of wild gypsy moth populations from across the invasive range in comparison to laboratory strains when reared on artificial diet and a preferred foliage host species, northern red oak (Quercus rubra L.). Overall, caterpillars reared on foliage had higher survival and faster development rates, with smaller differences among populations. When reared on artificial diet, laboratory strains had the highest performance as expected. The response from the wild populations was mixed, with two populations performing poorly on artificial diet and another performing nearly as well as the laboratory strains. Performance on diet was enhanced when larvae received cubed portions changed regularly, as opposed to filled cups. Understanding these relationships between food source and population performance is important for informing studies that examine population comparisons using wild and laboratory-reared strains.

Detection of pathogenic Batrachochytrium dendrobatidis using water filtration, animal and bait testing.

The pathogen Batrachochytrium dendrobatidis (Bd) can be challenging to detect at endangered amphibian reintroduction sites. Pre-release Bd detection can be confounded by imperfect animal sampling and the absence of animals. In Study 1, we used historical Bd-positive sites, to concurrently evaluate water filtrates and mouth bar (tadpoles) or skin swab (caudates) samples for Bd using molecular beacon realtime PCR. In Study 2, during a natural outbreak, we used PCR to detect Bd from zoospore-attracting keratin baits (three avian, three snake species). In Study 1, no captured animals (n=116) exhibited clinical signs, although 10.6% were positive, representing three of seven species sampled. In contrast, 5.4% of water filters (n=56) were Bd-positive. In Study 2, after short incubation times, a single duck down feather tested Bd-positive. In conclusion, Bd was detected in asymptomatic amphibians and water filtrate at two sites, and from water only, at two other sites. With continued refinement, semi-quantitative Bd water filtrate screening could better define zoospore-specific disease risk, allowing better characterization of the free-living phase of the organism's life cycle. Finally, these results suggest wild aquatic birds (e.g., waterfowl) should be systematically explored as a means of Bd spread. Since large numbers of aquatic birds migrate, even low Bd transfer rates could be a significant means for disease dissemination.

Sex ratio bias and extinction risk in an isolated population of Tuatara (Sphenodon punctatus).

Understanding the mechanisms underlying population declines is critical for preventing the extinction of endangered populations. Positive feedbacks can hasten the process of collapse and create an 'extinction vortex,' particularly in small, isolated populations. We provide a case study of a male-biased sex ratio creating the conditions for extinction in a natural population of tuatara (Sphenodon punctatus) on North Brother Island in the Cook Strait of New Zealand. We combine data from long term mark-recapture surveys, updated model estimates of hatchling sex ratio, and population viability modeling to measure the impacts of sex ratio skew. Results from the mark-recapture surveys show an increasing decline in the percentage of females in the adult tuatara population. Our monitoring reveals compounding impacts on female fitness through reductions in female body condition, fecundity, and survival as the male-bias in the population has increased. Additionally, we find that current nest temperatures are likely to result in more male than female hatchlings, owing to the pattern of temperature-dependent sex determination in tuatara where males hatch at warmer temperatures. Anthropogenic climate change worsens the situation for this isolated population, as projected temperature increases for New Zealand are expected to further skew the hatchling sex ratio towards males. Population viability models predict that without management intervention or an evolutionary response, the population will ultimately become entirely comprised of males and functionally extinct. Our study demonstrates that sex ratio bias can be an underappreciated threat to population viability, particularly in populations of long-lived organisms that appear numerically stable.

Behavioral and physiological female responses to male sex ratio bias in a pond-breeding amphibian.

INTRODUCTION: The phenomenon of sexual conflict has been well documented, and in populations with biased operational sex ratios the consequences for the rarer sex can be severe. Females are typically a limited resource and males often evolve aggressive mating behaviors, which can improve individual fitness for the male while negatively impacting female condition and fitness. In response, females can adjust their behavior to minimize exposure to aggressive mating tactics or minimize the costs of mating harassment. While male-male competition is common in amphibian mating systems, little is known about the consequences or responses of females. The red-spotted newt (Notophthalmus viridescens) is a common pond-breeding amphibian with a complex, well-studied mating system where males aggressively court females. Breeding populations across much of its range have male-biased sex ratios and we predicted that female newts would have behavioral mechanisms to mitigate mating pressure from males. We conducted four experiments examining the costs and behavioral responses of female N. viridescens exposed to a male-biased environment. RESULTS: In field enclosures, we found that female newts exposed to a male-biased environment during the five-month breeding season ended with lower body condition compared to those in a female-biased environment. Shorter-term exposure to a male-biased environment for five weeks caused a decrease in circulating total leukocyte and lymphocyte abundance in blood, which suggests females experienced physiological stress. In behavioral experiments, we found that females were more agitated in the presence of male chemical cues and females in a male-biased environment spent more time in refuge than those in a female-biased environment. CONCLUSIONS: Our results indicate that male-biased conditions can incur costs to females of decreased condition and potentially increased risk of infection. However, we found that females can also alter their behavior and microhabitat use under a male-biased sex ratio. Consistent with surveys showing reduced detection probabilities for females, our research suggests that females avoid male encounters using edge and substrate habitat. Our work illustrates the integrated suite of impacts that sexual conflict can have on the structure and ecology of a population.

Life history benefits of residency in a partially migrating pond-breeding amphibian.

Species with partial migration, where a portion of a population migrates and the other remains residential, provide the opportunity to evaluate conditions for migration and test mechanisms influencing migratory decisions. We conducted a five-year study of two populations of red-spotted newts (Notophthalmus viridescens), composed of individuals that either remain as residents in the breeding pond over the winter or migrate to the terrestrial habitat. We used multistate mark-recapture methods to (1) test for differences in survival probability between migrants and residents, (2) determine if migrants breed every year or skip opportunities for reproduction, and (3) estimate the frequency of individuals switching migratory tactic. We used estimates of life history parameters from the natural populations in combination with previous experimental work to evaluate processes maintaining partial migration at the population level and to assess mechanisms influencing the decision to migrate. Based on capture-recapture information on over 3000 individuals, we found that newts can switch migratory tactics over their lifetime. We conclude that migrants and residents coexist through conditional asymmetries, with residents having higher fitness and inferior individuals adopting the migrant tactic. We found that newts are more likely to switch from residency to migrating than the reverse and males were more likely to remain as residents. Migration differences between the sexes are likely driven by reproduction benefits of residency for males and high energetic costs of breeding resulting in lower breeding frequencies for females. Environmental conditions also influence partial migration within a population; we found support for density-dependent processes in the pond strongly influencing the probability of migrating. Our work illustrates how migration can be influenced by a complex range of individual and environmental factors and enhances our understanding of the conditions necessary for the evolution and maintenance of partial migration within populations.