High sugar diet alters immune function and the gut microbiome in juvenile green iguanas (Iguana iguana).

The present work aimed to study whether a high sugar diet can alter immune responses and the gut microbiome in green iguanas. Thirty-six iguanas were split into four treatment groups using a 2x2 design. Iguanas either received a sugar supplemented diet or a control diet, and either received a lipopolysaccharide (LPS) injection or a phosphate buffer solution (PBS) injection. Iguanas were given their respective diet treatment through the entire study (∼3 months) and received a primary immune challenge one month and two months into the experiment. Blood samples and cloacal swabs were taken at various points in the experiment and used to measure changes in the immune system (bacterial killing ability, lysis and agglutination scores, LPS specific IgY concentrations), and alterations in the gut microbiome. We found that sugar diet reduces bacterial killing ability following an LPS challenge, and sugar and the immune challenge temporarily alters gut microbiome composition while reducing alpha diversity. While sugar did not directly reduce lysis and agglutination following the immune challenge, the change in these scores over a 24-hour period following an immune challenge was more drastic (it decreased) relative to the control diet group. Moreover, sugar increased constitutive agglutination outside of the immune challenges (i.e., pre-challenge levels). In this study, we provide evidence that a high sugar diet affects the immune system of green iguanas (in a disruptive manner) and alters the gut microbiome.

Vertebrate Phenological Plasticity: From Molecular Mechanisms to Ecological and Evolutionary Implications.

Seasonal variation in the availability of essential resources is one of the most important drivers of natural selection on the phasing and duration of annually recurring life-cycle events. Shifts in seasonal timing are among the most commonly reported responses to climate change and the capacity of organisms to adjust their timing, either through phenotypic plasticity or evolution, is a critical component of resilience. Despite growing interest in documenting and forecasting the impacts of climate change on phenology, our ability to predict how individuals, populations, and species might alter their seasonal timing in response to their changing environments is constrained by limited knowledge regarding the cues animals use to adjust timing, the endogenous genetic and molecular mechanisms that transduce cues into neural and endocrine signals, and the inherent capacity of animals to alter their timing and phasing within annual cycles. Further, the fitness consequences of phenological responses are often due to biotic interactions within and across trophic levels, rather than being simple outcomes of responses to changes in the abiotic environment. Here, we review the current state of knowledge regarding the mechanisms that control seasonal timing in vertebrates, as well as the ecological and evolutionary consequences of individual, population, and species-level variation in phenological responsiveness. Understanding the causes and consequences of climate-driven phenological shifts requires combining ecological, evolutionary, and mechanistic approaches at individual, populational, and community scales. Thus, to make progress in forecasting phenological responses and demographic consequences, we need to further develop interdisciplinary networks focused on climate change science.

Maternal survival costs in an asocial mammal.

Maternal characteristics, social dynamics, and environmental factors can all influence reproduction and survival and shape trade-offs that might arise between these components of fitness. Short-lived mammals like the golden-mantled ground squirrel (GMGS; Callospermophilus lateralis) tend to maximize effort toward current reproduction at the expense of survival but may be complicated by other aspects of the species' life history and environment. Here, we use 25 years of data (1995-2020) collected from a population of GMGS at the Rocky Mountain Biological Research Laboratory in Gothic, Colorado, to test the effect of several maternal characteristics (e.g., age, experience, and timing of litter emergence), social context (e.g., litter sex ratio and kin density), and environmental context (e.g., date of bare ground and length of vegetative growing season) on survival of reproductive female GMGS using Cox proportional hazard models. Our results indicated that social dynamics (i.e., density) and environmental conditions (i.e., standardized first day of permanent snow cover and length of growing season) explained significant variation in annual maternal survival, while maternal characteristics did not. A higher density of related breeding females and the total number of females (both related and unrelated to the focal mother) were associated with an increase in the mortality hazard. A later standardized date of the first day of permanent snow cover and a shorter growing season both reduced the maternal mortality hazard. Together, our results suggest that factors extrinsic to the squirrels affect maternal survival and thus may also influence local population growth and dynamics in GMGS and other short-lived, territorial mammal species.

Impacts of COVID-19 on ecology and evolutionary biology faculty in the United States.

We surveyed ecologists and evolutionary biologists in American universities to understand how they are coping with the COVID-19 pandemic. Female respondents, assistant professors, and those who care for at least one child or teenager, were significantly more dissatisfied with their work-life balance during this pandemic than others, and further expected these negative impacts to be long lived. Online teaching support, relaxed expectations on publications, the possibility of pausing the tenure clock, and an acknowledgment of "no business as usual" by administrators were thought to be effective policies in mitigating these negative impacts. This survey serves as a manifesto to what our professional community is currently experiencing, and should be used to inform academic policies directed at improving faculty productivity and welfare.

Fishing in hot waters threatens phenotypic diversity.

In Focus: Morrongiello, J. R., Sweetman, P. C., & Thresher, R. E. (2019). Fishing constrains phenotypic responses of marine fish to climate variability. Journal of Animal Ecology, 88, 1645-1656. Forces of unnatural selection, such as climate change and harvest, are rarely studied in concert, yet hold the great potential to act synergistically on individual performance, susceptibility to harvest, tolerance to warming temperatures, and ultimately population persistence and resilience. In this paper, Morrongiello et al. (2019) used long-term monitoring of a site-attached temperate reef fish, the purple wrasse (Notolabrus fucicola), to test novel predictions about how fisheries management and climate variability could alter individual growth rates and thermal reaction norms within and across stocks. Otolith growth increments were collected from three south-east Australian populations between 1980 and 1999, pre- and post-harvest, throughout an intensive warming spell. Using hierarchical models to partition variation in growth within and between individuals and populations, Morrongiello et al. detected increased average growth rate with warming, a release from density dependence post-harvest, and a fishing-by-warming interaction that decreased diversity in thermal growth reaction norms because large individuals that tend to better tolerate warm temperatures were effectively culled from the population. This study outlines the importance of determining which phenotypes are more resilient to increasing temperatures, how fisheries should manage for them, and how such collective knowledge could help preserve and even promote resilience of managed populations to increasing temperatures in ecosystems threatened by climate change.

Seasonal climate effects on the survival of a hibernating mammal.

Global climate change and associated regional climate variability is impacting the phenology of many species, ultimately altering individual fitness and population dynamics. Yet, few studies have considered the effects of pertinent seasonal climate variability on phenology and fitness. Hibernators may be particularly susceptible to changes in seasonal climate since they have a relatively short active season in which to reproduce and gain enough mass to survive the following winter. To understand whether and how seasonal climate variability may be affecting hibernator fitness, we estimated survival from historical (1964-1968) and contemporary (2014-2017) mark-recapture data collected from the same population of Uinta ground squirrels (UGS, Urocitellus armatus), a hibernator endemic to the western United States. Despite a locally warming climate, the phenology of UGS did not change over time, yet season-specific climate variables were important in regulating survival rates. Specifically, older age classes experienced lower survival when winters or the following springs were warm, while juveniles benefited from warmer winter temperatures. Although metabolic costs decrease with decreasing temperature in the hibernacula, arousal costs increase with decreasing temperature. Our results suggest that this trade-off is experienced differently by immature and mature individuals. We also observed an increase in population density during that time period, suggesting resources are less limited today than they used to be. Cheatgrass is now dominating the study site and may provide a better food source to UGS than native plants did historically.

Interacting effects of unobserved heterogeneity and individual stochasticity in the life history of the southern fulmar.

Individuals are heterogeneous in many ways. Some of these differences are incorporated as individual states (e.g. age, size, breeding status) in population models. However, substantial amounts of heterogeneity may remain unaccounted for, due to unmeasurable genetic, maternal or environmental factors. Such unobserved heterogeneity (UH) affects the behaviour of heterogeneous cohorts via intra-cohort selection and contributes to inter-individual variance in demographic outcomes such as longevity and lifetime reproduction. Variance is also produced by individual stochasticity, due to random events in the life cycle of wild organisms, yet no study thus far has attempted to decompose the variance in demographic outcomes into contributions from UH and individual stochasticity for an animal population in the wild. We developed a stage-classified matrix population model for the southern fulmar breeding on Ile des Pétrels, Antarctica. We applied multievent, multistate mark-recapture methods to estimate a finite mixture model accounting for UH in all vital rates and Markov chain methods to calculate demographic outcomes. Finally, we partitioned the variance in demographic outcomes into contributions from UH and individual stochasticity. We identify three UH groups, differing substantially in longevity, lifetime reproductive output, age at first reproduction and in the proportion of the life spent in each reproductive state. -14% of individuals at fledging have a delayed but high probability of recruitment and extended reproductive life span. -67% of individuals are less likely to reach adulthood, recruit late and skip breeding often but have the highest adult survival rate. -19% of individuals recruit early and attempt to breed often. They are likely to raise their offspring successfully, but experience a relatively short life span. Unobserved heterogeneity only explains a small fraction of the variances in longevity (5.9%), age at first reproduction (3.7%) and lifetime reproduction (22%). UH can affect the entire life cycle, including survival, development and reproductive rates, with consequences over the lifetime of individuals and impacts on cohort dynamics. The respective role of UH vs. individual stochasticity varies greatly among demographic outcomes. We discuss the implication of our finding for the gradient of life-history strategies observed among species and argue that individual differences should be accounted for in demographic studies of wild populations.

Wolf in sheep's clothing: Model misspecification undermines tests of the neutral theory for life histories.

Understanding the processes behind change in reproductive state along life-history trajectories is a salient research program in evolutionary ecology. Two processes, state dependence and heterogeneity, can drive the dynamics of change among states. Both processes can operate simultaneously, begging the difficult question of how to tease them apart in practice. The Neutral Theory for Life Histories (NTLH) holds that the bulk of variations in life-history trajectories is due to state dependence and is hence neutral: Once previous (breeding) state is taken into account, variations are mostly random. Lifetime reproductive success (LRS), the number of descendants produced over an individual's reproductive life span, has been used to infer support for NTLH in natura. Support stemmed from accurate prediction of the population-level distribution of LRS with parameters estimated from a state dependence model. We show with Monte Carlo simulations that the current reliance of NTLH on LRS prediction in a null hypothesis framework easily leads to selecting a misspecified model, biased estimates and flawed inferences. Support for the NTLH can be spurious because of a systematic positive bias in estimated state dependence when heterogeneity is present in the data but ignored in the analysis. This bias can lead to spurious positive covariance between fitness components when there is in fact an underlying trade-off. Furthermore, neutrality implied by NTLH needs a clarification because of a probable disjunction between its common understanding by evolutionary ecologists and its translation into statistical models of life-history trajectories. Irrespective of what neutrality entails, testing hypotheses about the dynamics of change among states in life histories requires a multimodel framework because state dependence and heterogeneity can easily be mistaken for each other.

The Conundrum of Heterogeneities in Life History Studies.

What causes interindividual variation in fitness? Evidence of heritability of latent individual fitness traits has resparked a debate about the causes of variation in life histories in populations: neutralism versus empirical adaptationism. This debate about the processes underlying observed variation pits neutral stochastic demographic processes against evolutionarily relevant differences among individual fitness traits. Advancing this debate requires careful consideration of differences among inference approaches used by proponents of each hypothesis. Here we draw parallels between several disciplines focusing on processes generating variation in individuals' life-course, and we contrast methodologies to disentangle these processes. We draw on other disciplines to clarify terminology, risks of flawed inference, and expand the panel of hypotheses and formalizations of processes generating variation in life histories.

Effects of exploitation on an overabundant species: the lesser snow goose predicament.

Invasive and overabundant species are an increasing threat to biodiversity and ecosystem functioning world-wide. As such, large amounts of money are spent each year on attempts to control them. These efforts can, however, be thwarted if exploitation is compensated demographically or if populations simply become too numerous for management to elicit an effective and rapid functional response. We examined the influence of these mechanisms on cause-specific mortality in lesser snow geese using multistate capture-reencounter methods. The abundance and destructive foraging behaviours of snow geese have created a trophic cascade that reduces (sub-) Arctic plant, insect and avian biodiversity, bestowing them the status of 'overabundant'. Historically, juvenile snow geese suffered from density-related degradation of their saltmarsh brood-rearing habitat. This allowed harvest mortality to be partially compensated by non-harvest mortality (process correlation between mortality sources: ρ = -0.47; 90% BCI: -0.72 to -0.04). Snow goose family groups eventually responded to their own degradation of habitat by dispersing to non-degraded areas. This relaxed the pressure of density dependence on juvenile birds, but without this mechanism for compensation, harvest began to have an additive effect on overall mortality (ρ = 0.60; 90% BCI: -0.06 to 0.81). In adults, harvest had an additive effect on overall mortality throughout the 42-year study (ρ = 0.24; 90% BCI: -0.59 to 0.67). With the aim of controlling overabundant snow geese, the Conservation Order amendment to the International Migratory Bird Treaty was implemented in February of 1999 to allow for harvest regulations that had not been allowed since the early 1900s (e.g. a spring harvest season, high or unlimited bag limits and use of electronic calls and unplugged shotguns). Although harvest mortality momentarily increased following these actions, the increasing abundance of snow geese has since induced a state of satiation in harvest that has driven harvest rates below the long-term average. More aggressive actions will thus be needed to halt the growth and spread of the devastating trophic cascade that snow geese have triggered. Our approach to investigating the impacts of population control efforts on cause-specific mortality will help guide more effective management of invasive and overabundant species world-wide.

The influence of snowmobile trails on coyote movements during winter in high-elevation landscapes.

Competition between sympatric carnivores has long been of interest to ecologists. Increased understanding of these interactions can be useful for conservation planning. Increased snowmobile traffic on public lands and in habitats used by Canada lynx (Lynx canadensis) remains controversial due to the concern of coyote (Canis latrans) use of snowmobile trails and potential competition with lynx. Determining the variables influencing coyote use of snowmobile trails has been a priority for managers attempting to conserve lynx and their critical habitat. During 2 winters in northwest Wyoming, we backtracked coyotes for 265 km to determine how varying snow characteristics influenced coyote movements; 278 km of random backtracking was conducted simultaneously for comparison. Despite deep snow (>1 m deep), radio-collared coyotes persisted at high elevations (>2,500 m) year-round. All coyotes used snowmobile trails for some portion of their travel. Coyotes used snowmobile trails for 35% of their travel distance (random: 13%) for a mean distance of 149 m (random: 59 m). Coyote use of snowmobile trails increased as snow depth and penetrability off trails increased. Essentially, snow characteristics were most influential on how much time coyotes spent on snowmobile trails. In the early months of winter, snow depth was low, yet the snow column remained dry and the coyotes traveled off trails. As winter progressed and snow depth increased and snow penetrability increased, coyotes spent more travel distance on snowmobile trails. As spring approached, the snow depth remained high but penetrability decreased, hence coyotes traveled less on snowmobile trails because the snow column off trail was more supportive. Additionally, coyotes traveled closer to snowmobile trails than randomly expected and selected shallower snow when traveling off trails. Coyotes also preferred using snowmobile trails to access ungulate kills. Snow compaction from winter recreation influenced coyote movements within an area containing lynx and designated lynx habitat.

Climate change, phenology, and habitat degradation: drivers of gosling body condition and juvenile survival in lesser snow geese.

Nesting migratory geese are among the dominant herbivores in (sub) arctic environments, which have undergone unprecedented increases in temperatures and plant growing days over the last three decades. Within these regions, the Hudson Bay Lowlands are home to an overabundant breeding population of lesser snow geese that has dramatically damaged the ecosystem, with cascading effects at multiple trophic levels. In some areas the overabundance of geese has led to a drastic reduction in available forage. In addition, warming of this region has widened the gap between goose migration timing and plant green-up, and this 'mismatch' between goose and plant phenologies could in turn affect gosling development. The dual effects of climate change and habitat quality on gosling body condition and juvenile survival are not known, but are critical for predicting population growth and related degradation of (sub) arctic ecosystems. To address these issues, we used information on female goslings marked and measured between 1978 and 2005 (4125 individuals). Goslings that developed within and near the traditional center of the breeding colony experienced the effects of long-term habitat degradation: body condition and juvenile survival declined over time. In newly colonized areas, however, we observed the opposite pattern (increase in body condition and juvenile survival). In addition, warmer than average winters and summers resulted in lower gosling body condition and first-year survival. Too few plant 'growing days' in the spring relative to hatch led to similar results. Our assessment indicates that geese are recovering from habitat degradation by moving to newly colonized locales. However, a warmer climate could negatively affect snow goose populations in the long-run, but it will depend on which seasons warm the fastest. These antagonistic mechanisms will require further study to help predict snow goose population dynamics and manage the trophic cascade they induce.

Drivers of age-specific survival in a long-lived seabird: contributions of observed and hidden sources of heterogeneity.

1. We assessed the relative influence of variability in recruitment age, dynamic reproductive investment (time-specific reproductive states) and frailty (unobserved differences in survival abilities across individuals) on survival in the black-legged kittiwake. Furthermore, we examined whether observed variability in survival trajectories was best explained by immediate reproductive investment, cumulative investment, or both. 2. Individuals that delayed recruitment (≥ age 7) suffered a higher mortality risk than early recruits (age 3), especially later in life, suggesting that recruitment age may be an indicator of individual quality. Although recruitment age helped explain variation in survival, time-varying reproductive investment had a more substantial influence. 3. The dichotomy of attempting to breed or not explained variability in survival across life better than other parameterizations of reproductive states such as clutch size, brood size or breeding success. In the kittiwake, the sinequanon condition to initiate reproduction is to hold a nest site, which is considered a very competitive activity. This might explain why attempting to breed is the key level of investment that affects survival, independent of the outcome (failure or success). 4. Interestingly, the more individuals cumulate reproductive attempts over life, the lower their mortality risk, indicating that breeding experience may be a good indicator of parental quality as well. In contrast, attempting to breed at time t increased the risk of mortality between t and t + 1. We thus detected an immediate trade-off between attempting to breed and survival in this population; however, the earlier individuals recruited, and the more breeding experience they accumulated, the smaller the cost. 5. Lastly, unobserved heterogeneity across individuals improved model fit more (1·3 times) than fixed and dynamic sources of observed heterogeneity in reproductive investment, demonstrating that it is critical to account for both sources of individual heterogeneity when studying survival trajectories. Only after simultaneously accounting for both sources of heterogeneity were we able to detect the 'cost' of immediate reproductive investment on survival and the 'benefit' of cumulative breeding attempts (experience), a proxy to individual quality.

Consequences of recruitment decisions and heterogeneity on age-specific breeding success in a long-lived seabird.

An individual's age at first reproduction and investment in successive reproductive attempts are involved in mechanisms that can impede somatic repair, resulting in a decline in reproductive abilities with age (reproductive senescence). We used long-term data from the Black-legged Kittiwake, a long-lived seabird, to address the relationship between recruitment age, age-specific breeding success (BS), and reproductive senescence, while accounting for breeding experience and temporal variation in BS. We first detected late-life improvement in BS across all recruitment groups, which we recognized as "within-generation selection" or the selective disappearance of "frail" phenotypes. When such heterogeneity was accurately accounted for, we showed that all individuals suffered reproductive senescence. We first highlighted how different combinations of pre- and post-recruitment experience across recruitment groups resulted in maximal BS at intermediate ages. BS increased in early recruits as they gained post-recruitment experience, whereas late recruits gained pre-recruitment experience that led to high BS at recruitment. Only individuals recruiting at intermediate ages balanced their pre- and post-recruitment experience. Consistent with the "cumulative reproductive cost hypothesis," we also observed a faster decline in BS in early recruits at advanced ages, whereas individuals delaying recruitment experienced the slowest decline in BS with age. Early recruits, however, reached the highest levels of BS at intermediate ages, sensus stricto (10-13 years old), whereas individuals delaying recruitment experienced the lowest at similar ages. These divergent trajectories may reflect a "delayed trade-off" balancing a maximization of midlife BS against reproductive senescence at advanced ages. Additionally, annual variation in BS had a greater effect on individuals early in life, suggesting that experienced individuals were able to buffer out the effects of temporal variation on BS, which can ultimately improve fitness in stochastic environments. Our findings stress that (1) both observed and unobserved heterogeneity are important in detecting within-individual senescence, and (2) short-term trade-offs may be rare in long-lived species; thus, cumulated reproductive costs should be invoked as an alternative mechanism underlying reproductive senescence.