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Phenotypic plasticity and climate change: Can polar bears respond to longer Arctic summers with an adaptive fast?

Plasticity in the physiological and behavioural responses of animals to prolonged food shortages may determine the persistence of species under climate warming. This is particularly applicable for species that can “adaptively fast” by conserving protein to protect organ function while catabolizing endogenous tissues. Some Ursids, including polar bears ( Ursus maritimus ), adaptively fast during winter hibernation—and it has been suggested that polar bears also employ this strategy during summer. We captured 57 adult female polar bears in the Southern Beaufort Sea (SBS) during summer 2008 and 2009 and measured blood variables that indicate feeding, regular fasting, and adaptive fasting. We also assessed tissue δ13C and δ15N to infer diet, and body condition via mass and length. We found that bears on shore maintained lipid and protein stores by scavenging on bowhead whale ( Balaena mysticetus ) carcasses from human harvest, while those that followed the retreating sea ice beyond the continental shelf were food deprived. They had low ratios of blood urea to creatinine (U:C), normally associated with adaptive fasting. However, they also exhibited low albumin and glucose (indicative of protein loss) and elevated alanine aminotransferase and ghrelin (which fall during adaptive fasting). Thus, the ~ 70% of the SBS subpopulation that spends summer on the ice experiences more of a regular, rather than adaptive, fast. This fast will lengthen as summer ice declines. The resulting protein loss prior to winter could be a mechanism driving the reported correlation between summer ice and polar bear reproduction and survival in the SBS.

Oecologia

Forecasting selenium discharges to the San Francisco Bay-Delta Estuary: Ecological effects of a proposed San Luis Drain extension

Selenium discharges to the San Francisco Bay-Delta Estuary (Bay-Delta) could change significantly if federal and state agencies (1) approve an extension of the San Luis Drain to convey agricultural drainage from the western San Joaquin Valley to the North Bay (Suisun Bay, Carquinez Strait, and San Pablo Bay); (2) allow changes in flow patterns of the lower San Joaquin River and Bay-Delta while using an existing portion of the San Luis Drain to convey agricultural drainage to a tributary of the San Joaquin River; or (3) revise selenium criteria for the protection of aquatic life or issue criteria for the protection of wildlife. Understanding the biotransfer of selenium is essential to evaluating effects of selenium on Bay-Delta ecosystems. Confusion about selenium threats to fish and wildlife stem from (1) monitoring programs that do not address specific protocols necessary for an element that bioaccumulates; and (2) failure to consider the full complexity of the processes that result in selenium toxicity. Past studies show that predators are more at risk from selenium contamination than their prey, making it difficult to use traditional methods to predict risk from environmental concentrations alone. This report presents an approach to conceptualize and model the fate and effects of selenium under various load scenarios from the San Joaquin Valley. For each potential load, progressive forecasts show resulting (1) water-column concentration; (2) speciation; (3) transformation to particulate form; (4) particulate concentration; (5) bioaccumulation by invertebrates; (6) trophic transfer to predators; and (7) effects on those predators. Enough is known to establish a first-order understanding of relevant conditions, biological response, and ecological risks should selenium be discharged directly into the North Bay through a conveyance such as a proposed extension of the San Luis Drain. The approach presented here, the Bay-Delta selenium model, determines the mass, fate, and effects of selenium released to the Bay-Delta through use of (1) historical land-use, drainage, alluvial-fill, and runoff databases; (2) existing knowledge concerning biogeochemical reactions and physiological parameters of selenium (e.g., speciation, partitioning between dissolved and particulate forms, and bivalve assimilation efficiency); and (3) site-specific data mainly from 1986 to 1996 for clams and bottom-feeding fish and birds. Selenium load scenarios consider effluents from North Bay oil refineries and discharges of agricultural drainage from the San Joaquin Valley to enable calculation of (a) a composite freshwater endmember selenium concentration at the head of the estuary; and (b) a selenium concentration at a selected seawater location (Carquinez Strait) as a foundation for modeling. Analysis of selenium effects also takes into account the mode of conveyance for agricultural drainage (i.e., the San Luis Drain or San Joaquin River); and flows of the Sacramento River and San Joaquin River on a seasonal or monthly basis. Load scenarios for San Joaquin Valley mirror predictions made since 1955 of a worsening salt (and by inference, selenium) build-up exacerbated by an arid climate and massive irrigation. The reservoir of selenium in the San Joaquin Valley is sufficient to provide loading at an annual rate of approximately 42,500 pounds of selenium to a Bay-Delta disposal point for 63 to 304 years at the lower range of projections presented here, even if influx of selenium from the California Coast Ranges could be curtailed. Disposal of wastewaters on an annual basis outside of the San Joaquin Valley may slow the degradation of valley resources, but drainage alone cannot alleviate the salt and selenium build-up in the San Joaquin Valley, at least within a century. Load scenarios also show the different proportions of selenium loading to the Bay-Delta. Oil refinery loads from 1986 to 1992 ranged from 8.5 to 20 pounds of selenium per day; with treatment and cleanup, loads decreased to 3.0 pounds of selenium per day in 1999. In contrast, San Joaquin Valley agricultural drainage loads disposed of in a San Luis Drain extension could range from 45 to 117 pounds of selenium per day across a set of historical and future conditions. Components of this valley-wide load include five source subareas (i.e., Grassland, Westlands, Tulare, Kern, and Northern) defined by water and drainage management. Loads vary per subarea mainly because of proximity of the subarea to geologic sources of selenium and irrigation history. Loads from the Sacramento River, depending on flow conditions, range from 0.8 to 10 pounds of selenium per day. Loads from the San Joaquin River vary depending on restoration and flow conditions, which are considered. A consistent picture of ecological risk emerges under modeled selenium discharges from a proposed San Luis Drain extension. The threat to the estuary is greatest during low flow seasons and critically dry years. Where selenium undergoes reactions typical of low flow or longer residence time, highly problematic bioaccumulation in prey (food) is forecast. Surf scoter, greater and lesser scaup, and white sturgeon appear to be most at risk because these Bay-Delta predators feed on deposit and filter-feeding bivalves. Recent findings add Sacramento splittail and Dungeness crab to that list. During the low flow season of critically dry years, forecasted selenium concentrations in water, particulate matter, prey (diet), and predator tissue exceed guidelines with a high certainty of producing adverse effects under the most likely load scenario from a proposed San Luis Drain extension. High flows afford some protection under certain conditions in modeled San Joaquin River scenarios. However, meeting a combined goal of releasing a specific load during maximum flows and keeping selenium concentrations in the river below a certain objective to protect against bioaccumulation may not always be attainable. Management of the San Joaquin River on a constant concentration basis also could create problematic bioaccumulation during a wet year, especially during the low flow season, because high flows translate to high loads that are not always offset by seasonal river inflows. Prior to refinery cleanup, selenium contamination was sufficient to threaten reproduction in key species within the Bay-Delta ecosystems and human health advisories were posted based on selenium concentrations in tissues of diving ducks. During this time, selenium concentrations in the Bay-Delta were well below the most stringent recommended water quality criterion [1 microgram per liter (1 µg/L)]. Enhanced biogeochemical transformations to bioavailable particulate selenium and efficient bioaccumulation by bivalves characterized the system. If these biogeochemical conditions continue to prevail and agricultural selenium sources replace or exceed refinery sources, ecological forecasts suggest the risk of adverse effects will be difficult to eliminate under an out-of-valley resolution to the selenium problem. The Bay-Delta selenium model presented here is a systematic approach for conducting forecasts of the ecological effects from selenium on aquatic food webs. It is a new tool that links and models the major processes leading from loads through consumer organisms to predators. It also is a feasible approach for site-specific analysis and could provide a framework for developing new protective selenium foodweb guidelines and predator criteria. Model components that help ensure understanding ecosystems and the basis of environmental protection are (1) contaminant concentrations and speciation in sources, such as particulate material, that most influence bioavailability; (2) bioaccumulation models that calculate concentrations in diet, specifically in bivalves of the Bay-Delta that act as sensitive indicators of selenium contamination; (3) food-web type that determines what animals are threatened and when; and (4) multiple media concentrations (water, particulate material, and tissue of prey and predators) that, in-combination, determine risk or hazard.

California

Individual variation in temporal dynamics of post-release habitat selection

Translocated animals undergo a phase of behavioral adjustment after being released in a novel environment, initially prioritizing exploration and gradually shifting toward resource exploitation. This transition has been termed post-release behavioral modification. Post-release behavioral modification may also manifest as changes in habitat selection through time, and these temporal dynamics may differ between individuals. We aimed to evaluate how post-release behavioral modification is reflected in temporal dynamics of habitat selection and its variability across individuals using a population of translocated female greater sage-grouse as a case study. Sage-grouse were translocated from Wyoming to North Dakota (USA) during the summers of 2018–2020. We analyzed individual habitat selection as a function of sagebrush cover, herbaceous cover, slope, and distance to roads. Herbaceous cover is a key foraging resource for sage-grouse during summer; thus, we expected a shift from exploration to exploitation to manifest as temporally-varying selection for herbaceous cover. For each individual sage-grouse ( N = 26), we tested two competing models: a null model with no time-dependence and a model with time-dependent selection for herbaceous cover. We performed model selection at the individual level using an information-theoretic approach. Time-dependence was supported for five individuals, unsupported for seven, and the two models were indistinguishable based on AIC c for the remaining fourteen. We found no association between the top-ranked model and individual reproductive status (brood-rearing or not). We showed that temporal dynamics of post-release habitat selection may emerge in some individuals but not in others, and that failing to account for time-dependence may hinder the detection of steady-state habitat selection patterns. These findings demonstrate the need to consider both temporal dynamics and individual variability in habitat selection when conducting post-release monitoring to inform translocation protocols.

Frontiers in Conservation Science

Macroecological patterns of sexual size dimorphism in turtles of the world

Sexual size dimorphism (SSD) is a well-documented phenomenon in both plants and animals; however, the ecological and evolutionary mechanisms that drive and maintain SSD patterns across geographic space at regional and global scales are understudied, especially for reptiles. Our goal was to examine geographic variation of turtle SSD and to explore ecological and environmental correlates using phylogenetic comparative methods. We use published body size data on 135 species from nine turtle families to examine how geographic patterns and the evolution of SSD are influenced by habitat specialization, climate (annual mean temperature and annual precipitation) and climate variability, latitude, or a combination of these predictor variables. We found that geographic variation, magnitude and direction of turtle SSD are best explained by habitat association, annual temperature variance and annual precipitation. Use of semi-aquatic and terrestrial habitats was associated with male-biased SSD, whereas use of aquatic habitat was associated with female-biased SSD. Our results also suggest that greater temperature variability is associated with female-biased SSD. In contrast, wetter climates are associated with male-biased SSD compared with arid climates that are associated with female-biased SSD. We also show support for a global latitudinal trend in SSD, with females being larger than males towards the poles, especially in the families Emydidae and Geoemydidae. Estimates of phylogenetic signal for both SSD and habitat type indicate that closely related species occupy similar habitats and exhibit similar direction and magnitude of SSD. These global patterns of SSD may arise from sex-specific reproductive behaviour, fecundity and sex-specific responses to environmental factors that differ among habitats and vary systematically across latitude. Thus, this study adds to our current understanding that while SSD can vary dramatically across and within turtle species under phylogenetic constraints, it may be driven, maintained and exaggerated by habitat type, climate and geographic location.

Journal of Evolutionary Biology

A dynamic population model to investigate effects of climate and climate-independent factors on the lifecycle of the tick Amblyomma americanum (Acari: Ixodidae)

The lone star tick, Amblyomma americanum , is a disease vector of significance for human and animal health throughout much of the eastern United States. To model the potential effects of climate change on this tick, a better understanding is needed of the relative roles of temperature-dependent and temperature-independent (day-length-dependent behavioral or morphogenetic diapause) processes acting on the tick lifecycle. In this study, we explored the roles of these processes by simulating seasonal activity patterns using models with site-specific temperature and day-length-dependent processes. We first modeled the transitions from engorged larvae to feeding nymphs, engorged nymphs to feeding adults, and engorged adult females to feeding larvae. The simulated seasonal patterns were compared against field observations at three locations in United States. Simulations suggested that 1) during the larva-to-nymph transition, some larvae undergo no diapause while others undergo morphogenetic diapause of engorged larvae; 2) molted adults undergo behavioral diapause during the transition from nymph-to-adult; and 3) there is no diapause during the adult-to-larva transition. A model constructed to simulate the full lifecycle of A. americanum successfully predicted observed tick activity at the three U.S. study locations. Some differences between observed and simulated seasonality patterns were observed, however, identifying the need for research to refine some model parameters. In simulations run using temperature data for Montreal, deterministic die-out of A. americanum populations did not occur, suggesting the possibility that current climate in parts of southern Canada is suitable for survival and reproduction of this tick.

Journal of Medical Entomology

Offspring sex ratios are male-biased reflecting sex-biased dispersal in Idaho, USA, wolves.

Offspring sex ratios can vary widely across species, and the reasons for such variation have long intrigued ecologists. For group-living animals, predicting offspring sex ratios as a function of group and environmental characteristics can be challenging. Additionally, mortality of group members can upend traditional theory used to explain offspring sex ratios observed in populations. Gray wolves ( Canis lupus ) in Idaho, USA, are an excellent study species for asking questions about offspring sex ratios given their group-living behavior and persistent exposure to human-caused mortality. I hypothesized that offspring sex ratios would be influenced by the characteristics of individuals, groups, and populations. I generated genotypes for 419 adult and 400 pup wolves during 2008–2018. There was a significant male-bias in litters of wolf pups with nearly 12% more male pups born than females. The individual, group, and population variables I considered did not have significant associations with offspring sex ratios. Local resource competition helped explain offspring sex ratios in wolves in my study system, but not local resource enhancement theory. Although female helpers have been shown to help slightly more than males, offspring sex ratios did not favor the helping sex suggesting that the overall benefit of female helpers may have been negligible in wolf groups during my study. Three wolf groups consistently overproduced males, the dispersing sex, suggesting that habitat quality was poor in their territories. The male-biased offspring sex ratios observed throughout this population reflect sex-biased dispersal in wolves in Idaho. Such a pattern suggests breeding females may be reducing local resource competition (e.g., mates and successful reproduction) by producing more males than females.

Idaho

A core stochastic population projection model for Florida manatees (Trichechus manatus latirostris)

A stochastic, stage-based population model was developed to describe the life history and forecast the population dynamics of the Florida manatee (Trichechus manatus latirostris) in four separate regions of Florida. This population model includes annual variability in survival and reproductive rates, demographic stochasticity, effects of changes in warm-water capacity, and catastrophes. Further, the model explicitly accounts for uncertainty in parameter estimates. This model is meant to serve as a flexible tool for use in assessments relevant to management decision making, and was used in the State of Florida's recent biological status review. The parameter estimates and model structure described herein reflect our understanding of manatee demography at the time that this status review was completed. In the Northwest and Upper St. Johns regions, the model predicts that the populations will increase over time until warm-water capacity is reached, at which point growth will taper off. In the Atlantic region, the model predicts a stable or slightly increasing population over the next decade or so, and then a decrease as industrial warm-water capacity is lost. In the Southwest region, the model predicts a decline over time, driven by high annual mortality in the short-term and exacerbated by loss of industrial warm-water winter refuges over the next 40 years. Statewide, the likelihood of a 50% or greater decline in three manatee generations was 12%; the likelihood of a 20% or greater decline in two generations was 56%. These declines are largely driven by the anticipated loss of warm-water capacity, especially in the Atlantic and Southwest regions. The estimates of probability of extinction within 100 years were 11.9% for the Southwest region, 0.6% for the Northwest, 0.04% for the Atlantic, and <0.02% for the Upper St. Johns. The estimated probability that the statewide population will fall below 1000 animals within 100 years was 2.3%. Thus, while the estimated probability of extinction is low, the model predicts that current and emerging threats are likely to result in a long-term decline in the statewide population and a change in the regional distribution of manatees. Analyses of sensitivity and variance contribution highlight the importance of reducing uncertainty in some life-history parameters, particularly adult survival, temporal variance of adult survival, and long-term warm-water capacity. This core biological model is expected to evolve over time, as better information becomes available about manatees and their habitat, and as new assessment needs arise. We anticipate that this core model will be customized for other state and federal assessments in the near future.

Open-File Report

Mismatches between breeding phenology and resource abundance of resident alpine ptarmigan negatively affect chick survival

1. Phenological mismatches – defined here as the difference in reproductive timing of an individual relative to the availability of its food resources – occur in many avian species. Mistiming breeding activities in environments with constrained breeding windows may have severe fitness costs due to reduced opportunities for repeated breeding attempts. Therefore, species occurring in alpine environments may be particularly vulnerable. 2. We studied fitness consequences of timing of breeding in an alpine-endemic species, the white-tailed ptarmigan ( Lagopus leucura ), to investigate its influence on chick survival. We estimated phenological mismatch by measuring plant and arthropods used by ptarmigan in relation to their timing of breeding. 3. We monitored 120 nests and 67 broods over a three-year period (2013–2015) at three alpine study sites in the Rocky Mountains of Colorado. During this same period we actively monitored food resource abundance in brood-use areas to develop year and site specific resource phenology curves. We developed several mismatch indices from these curves that were then fit as covariates in mark-recapture chick survival models. 4. A correlation analysis between seasonal changes in arthropod and food plant abundance indicated that a normalized difference vegetation index (NDVI) was likely the best predictor for food available to hens and chicks. A survival model that included an interaction between NDVI mismatch and chick age received strong support and indicated young chicks were more susceptible to mismatch than older chicks. 5. We provide evidence that individual females of a resident alpine species can be negatively affected by phenological mismatch. Our study focused on individual females and did not examine if phenological mismatch was present at the population level. Future work in animal populations occurring in mountain systems focusing on a combination of both individual- and population- level metrics of mismatch will be beneficial.

Ecology and Evolution

Growth rate variation in Brown Treesnakes (Boiga irregularis): An invasive species of conservation concern

Somatic growth rate is a fundamental trait that influences metabolism, lifespan and reproductive maturity and is critical for understanding population dynamics and informing management actions. Brown Treesnakes ( Boiga irregularis ) introduced to Guam are highly invasive and can reproduce year-round without discrete cohorts. We compared snake size trajectories described by the conventionally used von Bertalanffy growth function versus the Gompertz model. Using quantile regression with a regularized effect for individual snakes we modeled growth rates of 270 marked, wild snakes as a function of size. The Gompertz model explained more of the variation in growth and rendered more realistic predictions of asymptotic sizes than did the von Bertalanffy model. With the Gompertz model, growth rates were 1.05–1.16× faster in males than in females. Females reached asymptotic sizes at shorter snout-vent lengths than males. Growth rate was positively correlated with amount of precipitation, and modeling wet-dry seasonality on Guam as a sinusoidal function identified a growth peak in September—October. Effects of seasonality and precipitation, however, were minor compared to individual and sex related differences in size-adjusted growth rates. We estimated that the 50th (and 5th, 95th) growth-rate percentile males in our study population become sexually mature at an age of 33 (∞, 15) months, while females mature at 41 (∞, 18) months, where ∞ indicates that the slowest growing snakes never reach maturity. However, 50% of the snakes mature at a size below the median, and age at maturity may be as low as 10.4 (males) and 13.7 (females) months for average-sized hatchlings that grow fast. Our results have implications for the timing of management options for this species and our approach can be broadly applied to animals where repeated growth data are obtained and age is unknown.

Ecology and Evolution

Translocation and early post-release demography of endangered Laysan teal

In an attempt to reduce the high extinction risk inherent to small island populations, we translocated wild Laysan teal Anas laysanensis to a portion of its presumed prehistoric range. Most avian translocations lack the strategic post-release monitoring needed to assess early population establishment or failure. Therefore, we monitored the survival and reproduction of all founders, and their first-generation offspring using radio telemetry for 2 years after the first release. Forty-two Laysan teal were sourced directly from the only extant population on Laysan Island and transported 2 days by ship to Midway Atoll. All birds survived the translocation with nutritional and veterinary support, and spent between 4 and 14 days in captivity. Post-release survival of 42 founders was 0.857 (95% CI 0.86-0.99) during 2004-2006 or annualized 0.92 (95% CI 0.83-0.98). Seventeen of 18 founding hens attempted nesting in the first two breeding seasons. Fledgling success was 0.57 (95% CI 0.55-0.60) in 2005 and 0.63 (95% CI 0.62-0.64) in 2006. The effective founding female population (Ne) was 13. We applied these initial demographic rates to model population growth. The nascent population size increased to >100 after only 2 years post-release (?? = 1.73). If this growth rate continues, the size of the Midway population could surpass the source population before 2010. ?? 2008 The Authors. Journal compilation ?? 2008 The Zoological Society of London.

Animal Conservation

Sources of variation in maternal allocation in a long-lived mammal

1. Life history theory predicts allocation of energy to reproduction varies with maternal age, but additional maternal features may be important to the allocation of energy to reproduction. We aimed to characterize age‐specific variation in maternal allocation and assess the relationship between maternal allocation and other static and dynamic maternal features. Mass measurements of 531 mothers and pups were used with Bayesian hierarchical models to explain the relationship between diverse maternal attributes and both the proportion of mass allocated by Weddell seal mothers, and the efficiency of mass transfer from mother to pup during lactation as well as the weaning mass of pups. Our results demonstrated that maternal mass was strongly and positively associated with the relative reserves allocated by a mother and a pup's weaning mass but that the efficiency of mass transfer declines with maternal parturition mass. Birthdate was positively associated with proportion mass allocation and pup weaning mass, but mass transfer efficiency was predicted to be highest at the mean birthdate. The relative allocation of maternal reserves declined with maternal age but the efficiency of mass transfer to pups increases, suggestive of selective disappearance of poor‐quality mothers. These findings highlight the importance of considering multiple maternal features when assessing variation in maternal allocation.

Journal of Animal Ecology

Life-stage-specific physiology defines invasion extent of a riverine fish

Many ecologists have called for mechanism-based investigations to identify the underlying controls on species distributions. Understanding these controls can be especially useful to construct robust predictions of how a species range may change in response to climate change or the extent to which a non-native species may spread in novel environments. Here, we link spatially intensive observations with mechanistic models to illustrate how physiology determines the upstream extent of the aquatic ectotherm smallmouth bass ( Micropterus dolomieu ) in two headwater rivers. Our results demonstrate that as temperatures become increasingly cold across a downstream to upstream gradient, food consumption in age 0 bass becomes increasingly constrained, and as a result, these fish become growth limited. Sufficient first summer growth of age 0 bass is essential for overwinter survival because young bass must persist from energy reserves accumulated during the summer, and those reserves are determined by body size. Our field data reveal the upstream extent of adult bass reproduction corresponds to a point in the downstream/upstream gradient where cold temperatures impair growth opportunities in young bass. This pattern was repeated in both study streams and explained why bass positioned nests twice as far upstream in the warm compared to the cold stream in the same basin. Placement of spawning nests by adult bass is likely subject to strong evolutionary selection in temperate systems: if bass spawn too far upstream, their young are unlikely to grow large enough to survive the winter. Consumption and growth in older bass (age 3–4) was far less sensitive to temperature. Based on these data, we suggest that temperature-sensitive age 0 bass constrain the upstream distribution limits of bass within temperate streams. In this study, we investigated how temperature-dependent physiology changed through the life history of a species and, in doing so, identified a climate-sensitive life-history stage that likely sets the distributional limits of all other life-history stages. We anticipate the framework developed here could be employed to identify how similar stage-specific environmental sensitivity determines distribution in many other ectothermic species.

Oregon

Survival of whirling disease resistant rainbow trout fry in the wild: A comparison of two strains

Introduced pathogens can affect fish populations, and three main factors affect disease occurrence: the environment, host, and pathogen. Manipulating at least one of these factors is necessary for controlling disease. Myxobolus cerebralis , the parasite responsible for salmonid whirling disease, became established in Colorado during the 1990s and caused significant declines in wild Rainbow Trout Oncorhynchus mykiss populations. Attempts to re-establish Rainbow Trout have focused on manipulating salmonid host resistance. A Rainbow Trout strain known as GR × CRR was developed for stocking in Colorado by crossing a whirling-disease-resistant strain known as the German Rainbow Trout (GR) with the Colorado River Rainbow Trout (CRR). The GR × CRR fish exhibit resistance similar to that shown by GR, and survival and reproduction were expected to be similar to those of CRR. One disadvantage of stocking GR × CRR is that outcrossing and backcrossing could decrease resistance, and laboratory studies have indicated that this can occur. A potential disadvantage of stocking pure GR is lower survival due to domestication. To compare fry survival between the strains, a field experiment was conducted in 1.6-km reaches of nine Colorado streams. Each stream was stocked in August 2014 with 5,000 GR × CRR and 5,000 GR individuals. In October 2014, April 2015, and August 2015, apparent survival was assessed. Two laboratory predation experiments were also conducted. The field experiment revealed that short-term apparent survival was influenced by stream, and growth rate was influenced by strain and stream. However, after 12 months, there was no difference in apparent survival or growth rate between the GR and GR × CRR strains. Laboratory experiments showed that survival did not differ between the strains when confronted with Brown Trout Salmo trutta predation. Our results indicate that the GR strain is a viable option for stocking in streams where M. cerebralis is enzootic. Further evaluation is needed to determine whether GR fish will survive to maturity and reproduce.

Journal of Aquatic Animal Health

Exposure to domoic acid is an ecological driver of cardiac disease in southern sea otters

Harmful algal blooms produce toxins that bioaccumulate in the food web and adversely affect humans, animals, and entire marine ecosystems. Blooms of the diatom Pseudo-nitzschia can produce domoic acid (DA), a toxin that most commonly causes neurological disease in endothermic animals, with cardiovascular effects that were first recognized in southern sea otters. Over the last 20 years, DA toxicosis has caused significant morbidity and mortality in marine mammals and seabirds along the west coast of the USA. Identifying DA exposure has been limited to toxin detection in biological fluids using biochemical assays, yet measurement of systemic toxin levels is an unreliable indicator of exposure dose or timing. Furthermore, there is little information regarding repeated DA exposure in marine wildlife. Here, the association between long-term environmental DA exposure and fatal cardiac disease was investigated in a longitudinal study of 186 free-ranging sea otters in California from 2001 – 2017, highlighting the chronic health effects of a marine toxin. A novel Bayesian spatiotemporal approach was used to characterize environmental DA exposure by combining several DA surveillance datasets and integrating this with life history data from radio-tagged otters in a time-dependent survival model. In this study, a sea otter with high DA exposure had a 1.7-fold increased hazard of fatal cardiomyopathy compared to an otter with low exposure. Otters that consumed a high proportion of crab and clam had a 2.5- and 1.2-times greater hazard of death due to cardiomyopathy than otters that consumed low proportions. Increasing age is a well-established predictor of cardiac disease, but this study is the first to identify that DA exposure affects the risk of cardiomyopathy more substantially in prime-age adults than aged adults. A 4-year-old otter with high DA exposure had 2.3 times greater risk of fatal cardiomyopathy than an otter with low exposure, while a 10-year old otter with high DA exposure had just 1.2 times greater risk. High Toxoplasma gondii titers also increased the hazard of death due to heart disease 2.4-fold. Domoic acid exposure was most detrimental for prime-age adults, whose survival and reproduction are vital for population growth, suggesting that persistent DA exposure will likely impact long-term viability of this threatened species. These results offer insight into the pervasiveness of DA in the food web and raise awareness of under-recognized chronic health effects of DA for wildlife at a time when toxic blooms are on the rise.

California

Birds choose long-term partners years before breeding

Pair bonds can provide social benefits to long-term monogamous species alongside their benefits for reproduction. However, little is known about when these bonds form, in particular how long they are present before breeding. Previous studies of pair formation in long-term monogamous birds have been rather data-limited, but for many migratory birds they report pair formation on the wintering grounds. We provide the first systematic investigation of prebreeding association patterns of long-term monogamous pairs by examining entire life histories based on tracking data of migratory whooping cranes, Grus americana. We found that a substantial portion (62%) of breeding pairs started associating at least 12 months before first breeding, with 16 of 58 breeding pairs beginning to associate over 2 years before first breeding. For most pairs, these associations with future breeding partners also became unique and distinguishable from association patterns with nonpartner individuals 12 months before first breeding. In addition, 60% of pair associations began before at least one partner had reached nominal sexual maturity. Most pairs began associating in the late spring upon arrival at the summer grounds, while associations beginning at other times of the year were rare. Patterns in the associations of pairs prior to breeding can point to the potential benefits of prebreeding relationships, for instance providing support in competitive interactions or increasing partner familiarity.

Animal Behaviour

Estimating golden-cheeked warbler immigration: Implications for the spatial scale of conservation

Understanding the factors that drive population dynamics is fundamental to species conservation and management. Since the golden-cheeked warbler S etophaga chrysoparia was first listed as endangered, much effort has taken place to monitor warbler abundance, occupancy, reproduction and survival. Yet, despite being directly related to local population dynamics, movement rates have not been estimated for the species. We used an integrated population model to investigate the relationship between immigration rate, fledging rate, survival probabilities and population growth rate for warblers in central Texas, USA. Furthermore, using a deterministic projection model, we examined the response required by vital rates to maintain a viable population across varying levels of immigration. Warbler abundance fluctuated with an overall positive trend across years. In the absence of immigration, the abundance would have decreased. However, the population could remain viable without immigration if both adult and juvenile survival increased by almost half or if juvenile survival more than doubled. We also investigated the response required by fledging rates across a range of immigration in order to maintain a viable population. Overall, we found that immigration was required to maintain warbler target populations, indicating that warbler conservation and management programs need to be implemented at larger spatial scales than current efforts to be effective. This study also demonstrates that by using limited data within integrated population models, biologists are able to monitor multiple key demographic parameters simultaneously to gauge the efficacy of strategies designed to maximize warbler viability in a changing landscape.

Animal Conservation

Machine learned daily life history classification using low frequency tracking data and automated modelling pipelines: Application to North American waterfowl

Background Identifying animal behaviors, life history states, and movement patterns is a prerequisite for many animal behavior analyses and effective management of wildlife and habitats. Most approaches classify short-term movement patterns with high frequency location or accelerometry data. However, patterns reflecting life history across longer time scales can have greater relevance to species biology or management needs, especially when available in near real-time. Given limitations in collecting and using such data to accurately classify complex behaviors in the long-term, we used hourly GPS data from 5 waterfowl species to produce daily activity classifications with machine-learned models using “automated modelling pipelines”. Methods Automated pipelines are computer-generated code that complete many tasks including feature engineering, multi-framework model development, training, validation, and hyperparameter tuning to produce daily classifications from eight activity patterns reflecting waterfowl life history or movement states. We developed several input features for modeling grouped into three broad categories, hereafter “feature sets”: GPS locations, habitat information, and movement history. Each feature set used different data sources or data collected across different time intervals to develop the “features” (independent variables) used in models. Results Automated modelling pipelines rapidly developed easily reproducible data preprocessing and analysis steps, identification and optimization of the best performing model and provided outputs for interpreting feature importance. Unequal expression of life history states caused unbalanced classes, so we evaluated feature set importance using a weighted F1-score to balance model recall and precision among individual classes. Although the best model using the least restrictive feature set (only 24 hourly relocations in a day) produced effective classifications (weighted F1 = 0.887), models using all feature sets performed substantially better (weighted F1 = 0.95), particularly for rarer but demographically more impactful life history states (i.e., nesting). Conclusions Automated pipelines generated models producing highly accurate classifications of complex daily activity patterns using relatively low frequency GPS and incorporating more classes than previous GPS studies. Near real-time classification is possible which is ideal for time-sensitive needs such as identifying reproduction. Including habitat and longer sequences of spatial information produced more accurate classifications but incurred slight delays in processing.

Movement Ecology

Fat or lean: adjustment of endogenous energy stores to predictable and unpredictable changes in allostatic load

1. The ability to store energy endogenously is an important ecological mechanism that allows animals to buffer predictable and unpredictable variation in allostatic load. The secretion of glucocorticoids, which reflects changes in allostatic load, is suggested to play a major role in the adjustment of endogenous stores to these varying conditions. 2. Although crucially important, the relationship between allostatic load and energy stores remains largely unexplored. Two contrasting hypotheses describe how stores may be adjusted: animals may use low allostatic loads to increase stores to a maximum possible (‘fat and fit’), or they can attain a lean physique due to fitness advantages of a low body mass (‘lean and fit’). 3. We compiled observational and experimental data available for a long-lived seabird to examine the relationship between glucocorticoids and stored energy at two life history stages (incubation and chick-rearing). Data were collected across multiple years and colonies in the North Pacific, thereby reflecting the wide range of environmental conditions birds' encounter in the marine environment. During experimental manipulations, allostatic load was minimized by supplementing food to free-living birds. 4. We found that the relationship between allostatic load and energy stores was clearly curvilinear at both life history stages. Observational data suggested that energy stores remained relatively stable under low allostatic load and decreased under high loads. Experimental data showed that birds did not maximize energy stores under favourable conditions but maintained energy stores below a physiologically attainable level. 5. Energy stores remained consistently lower during chick-rearing compared to incubation across the wide range of variations in allostatic load suggesting that stage-specific trade-offs limit the accumulation of energy during favourable environmental conditions. Secretion of glucocorticoids did not appear to mediate this shift in energy stores between the life history stages. 6. Overall, results of this study support the ‘lean and fit’ hypothesis. We conclude that increased energy stores may not necessarily reflect better environmental conditions experienced by individuals or predict their higher fitness. A major advantage of adopting a lean physique when environmental conditions allow may be the avoidance of additional energetic costs for moving a heavy body. In breeding seabirds, this advantage may be more important during chick-rearing. In the focal species, the secretion of glucocorticoids might be involved in regulation of energy stores within a life history stage but does not appear to mediate an adaptive shift in energy stores between the incubating and chick-rearing stages of reproduction.

Functional Ecology