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Amphibian terrestrial habitat selection and movement patterns vary with annual life-history period

Identification of essential habitat is a fundamental component of amphibian conservation; however, species with complex life histories frequently move among habitats. To better understand dynamic habitat use, we evaluated Wood Frog ( Lithobates sylvaticus (LeConte, 1825)) habitat selection and movement patterns during the spring migration and foraging periods and described the spatiotemporal variability of habitats used during all annual life-history periods. We radio-tracked 71 frogs in Maine during 2011–2013 and evaluated spring migration, foraging activity center (FAC), and within-FAC habitat selection. Telemetered frogs spent the greatest percentage of each field season in hibernacula (≥54.4%), followed by FACs (≥25.5%), migration habitat (≥16.9%), and breeding sites (≥4.5%). FACs ranged 49 – 1 335 m 2 (568.0 ± 493.4 m 2 ) and annual home ranges spanned 1 413 – 32 165 m 2 (11 780.6 ± 12 506.1 m 2 ). During spring migration, Wood Frogs exhibited different movement patterns (e.g., turn angles), selected different habitat features, and selected habitat features less consistently than while occupying FACs, indicating that the migration and foraging periods are ecologically distinct. Habitat-use studies that do not discriminate among annual life-history periods may obscure true ecological relationships and fail to identify essential habitat necessary for sustaining amphibian populations.

Maine

Factors affecting gray wolf (Canis lupus) encounter rate with elk (Cervus elaphus) in Yellowstone National Park

Despite encounter rates being a key component of kill rate, few studies of large carnivore predation have quantified encounter rates with prey, the factors that influence them, and the relationship between encounter rate and kill rate. The study’s primary motivation was to determine the relationship between prey density and encounter rate in understanding the mechanism behind the functional response. Elk (Cervus elaphus Linnaeus, 1758) population decline and variable weather in northern Yellowstone National Park provided an opportunity to examine how these factors influenced wolf (Canis lupus Linnaeus, 1758) encounter rates with elk. We explored how factors associated with wolf kill rate and encounter rate in other systems (season, elk density, elk group density, average elk group size, snow depth, wolf pack size, and territory size) influenced wolf–elk encounter rate in Yellowstone National Park. Elk density was the only factor significantly correlated with wolf–elk encounter rate, and we found a nonlinear density-dependent relationship that may be a mechanism for a functional response in this system. Encounter rate was correlated with number of elk killed during early winter but not late winter. Weak effects of snow depth and elk group size on encounter rate suggest that these factors influence kill rate via hunting success because kill rate is the product of hunting success and encounter rate.

Wyoming

Extreme climatic variability during migration invokes physiological and dietary plasticity among spring migrating ducks

Environmental stochasticity encountered during migration can have negative consequences for individuals and population demographics through direct reductions in survival or cross-seasonal impacts. We took advantage of substantial interannual variation in spring migration conditions over a 4 year field study to examine physiological and dietary variation among two species of migrant ducks. We collected female Lesser Scaup ( Aythya affinis (Eyton, 1838)) and Blue-winged Teal ( Spatula discors (Linnaeus, 1766)) during spring migration and measured lipid and protein reserves, an index of recent lipid metabolism based on concentrations of lipid metabolites in plasma, and diets. We documented systematic interannual variation among these metrics in both species, contrasting primarily the warmest, earliest spring and the coldest, latest spring. Lesser Scaup had reduced lipid and protein reserves and consumed less energy-rich prey during the coldest and latest spring but showed no interannual variation in the index of lipid metabolism. Blue-winged Teal similarly had reduced protein reserves in the cold, late spring but maintained constant lipid reserves among years, likely facilitated by increased consumption of energy-rich seeds reflected in diets and lipid metabolism. Our results reveal impacts of environmental stochasticity on migrants and suggest that recruitment may be impacted by variable conditions encountered during migration during extreme weather events.

South Dakota

Annual cycle of White-winged Scoters (Melanitta fusca) in eastern North America: Migratory phenology, population delineation, and connectivity

Understanding full annual cycle movements of long-distance migrants is essential for delineating populations, assessing connectivity, evaluating crossover effects between life stages, and informing management strategies for vulnerable or declining species. We used implanted satellite transmitters to track up to 2 years of annual cycle movements of 52 adult female White-winged Scoters ( Melanitta fusca (Linnaeus, 1758)) captured in the eastern United States and Canada. We used these data to document annual cycle phenology; delineate migration routes; identify primary areas used during winter, stopover, breeding, and molt; and assess the strength of migratory connectivity and spatial population structure. Most White-winged Scoters wintered along the Atlantic coast from Nova Scotia to southern New England, with some on Lake Ontario. White-winged Scoters followed four migration routes to breeding areas from Quebec to the Northwest Territories. Principal postbreeding molting areas were in James Bay and the St. Lawrence River estuary. Migration phenology was synchronous regardless of winter or breeding origin. Cluster analyses delineated two primary breeding areas: one molting area and one wintering area. White-winged Scoters demonstrated overall weak to moderate connectivity among life stages, with molting to wintering connectivity the strongest. Thus, White-winged Scoters that winter in eastern North America appear to constitute a single continuous population.

Canadian Journal of Zoology

Relatedness within and among Myotis septentrionalis colonies at a local scale

Abstract: We assessed parentage within and among maternity colonies of the northern long-eared bat (Myotis septentrionalis Troessart 1897) in north-central Kentucky from 2011–2013 to better understand colony social structure, formation, and membership dynamics. We intensively sampled colonies in close and remote (> 10 km) spatial proximity both before and after targeted day-roost removal. Colonies were not necessarily comprised of closely related individuals, but natal philopatry was common. Adjacent colonies often contained maternally related individuals, indicating that some pups did disperse, albeit not far from their natal home range. Lack of apparent overlap among maternity colonies, along with no observed individual movements between colonies, suggests that colonies may be relatively closed once established in the maternity season. Whereas some young on site had been sired by males collected on site that by chance had dispersed to the same summering grounds, most had not, as would be expected since the species mates in the fall swarms near hibernacula. The number of parentages that we inferred among colonies, however, suggests that outside the maternity season, social groups may be relatively flexible and open, with individuals moving among groups close to their natal area. Analysis of microsatellite DNA data showed a low FST (= 0.011) and best fit to a model of one multilocus genotypic cluster across the study area. We observed high turnover in colony membership between years in all colonies, regardless of roost removal treatment. Our results suggest that female northern long-eared bats exhibit fidelity to a general geographic area rather than individual colonies between years, and indicate presence of a complex and dynamic social-genetic structure. Greater understanding of colony dynamics, including formation, dissolution, and dispersal patterns, may contribute to conservation and management of this threatened species.

Kentucky

Tools to understand seasonality in health: quantification of microbe loads and analyses of compositional ecoimmunological data reveal complex patterns in Mojave Desert Tortoise (Gopherus agassizii) populations

Using data from six wild Mojave Desert Tortoise ( Gopherus agassizii (Cooper, 1861)) populations, we quantified seasonal differences in immune system measurements and microbial load in the respiratory tract, pertinent to this species’ susceptibility to upper respiratory tract disease. We quantified bacteria-killing activity of blood plasma and differential leukocyte counts to detect trends in temporal variation in immune function. We used centered log-ratio (clr) transformations of leukocyte counts and stress that such transformations are necessary for compositional data. We tested animals for the potential pathogen Pasteurella testudinis Snipes and Biberstein, 1982 with a newly created quantitative polymerase chain reaction (qPCR) assay, as well as for the known respiratory pathogens Mycoplasma agassizii Brown et al., 2001 and Mycoplasma testudineum Brown et al., 2004. We found very little disease and suggest that P. testudinis is a prevalent, commensal microbe in these Mojave Desert Tortoise populations, and its quantification may be a tool to study natural fluctuations in microbe levels in Mojave Desert Tortoise respiratory tracts. Our analyses showed that both the potential for inflammatory responses and microbe levels are highest in the spring for healthy Mojave Desert Tortoises, when lymphocyte levels are lowest. The genetic and statistical tools that we used are easily applicable to other wildlife systems and provide the necessary data to quantify species-wide trends in health and test hypotheses pertinent to host–microbe dynamics.

Canadian Journal of Zoology

Testing theoretical metapopulation conditions with genotypic data from Boreal Chorus Frogs (Pseudacris maculata)

The metapopulation concept has far-reaching implications in ecology and conservation biology. Hanski’s criteria operationally define metapopulations, yet testing them is hindered by logistical and financial constraints inherent to the collection of long-term demographic data. Hence, ecologists and conservationists often assume metapopulation existence for dispersal-limited species that occupy patchy habitats. To advance understanding of metapopulation theory and improve conservation of metapopulations, we used population and landscape genetic tools to develop a methodological framework for evaluating Hanski’s criteria. We used genotypic data (11 microsatellite loci) from a purported metapopulation of Boreal Chorus Frogs ( Pseudacris maculata (Agassiz, 1850)) in Colorado, U.S.A., to test Hanski’s four criteria. We found support for each criterion: (1) significant genetic differentiation between wetlands, suggesting distinct breeding populations; (2) wetlands had small effective population sizes and recent bottlenecks, suggesting populations do not experience long-term persistence; (3) population graphs provided evidence of gene flow between patches, indicating potential for recolonization; and (4) multiscale bottleneck analyses suggest asynchrony, indicating that simultaneous extinction of all populations was unlikely. Our methodological framework provides a logistically and financially feasible alternative to long-term demographic data for identifying amphibian metapopulations.

Colorado

Rapid adoption of nestboxes by Prothonotary Warblers (Protonotaria citrea) in mesic deciduous forest

Breeding territory selection in Prothonotary Warblers ( Protonotaria citrea (Boddaert, 1783)) is thought to hinge on standing water, with a strong preference for low-lying areas prone to seasonal flooding. However, we have observed this species nesting in much drier areas than previously reported. We recently initiated a study of the Carolina Wren ( Thryothorus ludovicianus (Latham, 1790)) using wooden nest boxes, and nearly 60% of all nests produced in these boxes during the initial study year were produced by Prothonotary Warblers, despite this species being absent from our field site during the year preceding nest-box availability. Most nests were produced in dense, closed-canopy forest with a thick shrub layer >100 m from any water body. There was no difference in the mean distance from water between nests of the Prothonotary Warbler and those of the Carolina Wren, a habitat generalist that does not nest over water. We then observed a 60% increase in the number of Prothonotary Warbler nests the following year, along with significant increases in breeding productivity. Although they nested on sites that they are not thought to prefer, our observations suggest that Prothonotary Warblers may nest in drier areas than usual if appropriate nest cavities are provided.

Canadian Journal of Zoology

Latitudinal variation in snowshoe hare (Lepus americanus) body mass: A test of Bergmann’s Rule

The relationship between body size and latitude have been the focus of dozens of studies across many species. However, results of testing Bergmann’s Rule – that organisms in colder climates or at higher latitudes possess larger body sizes – have been inconsistent across studies. We investigated whether snowshoe hares (Lepus americanus) follow the Rule by investigating differences in body mass using data from six published studies and from data of 755 individual hares captured from ten populations across North America covering 26° of latitude. We also explored alternative hypotheses related to variation in hare body mass, including winter severity, length of growing season, elevation, and snow depth. We found body mass of hares varied throughout their range, but the drivers of body mass differed based on geographic location. Females in northern populations followed Bergmann’s rule, whereas males did not. In northern populations male mass was related to average snow depth. In contrast, in southern populations body mass of both sexes was related to length of the growing season. These differences likely represent variation in the drivers of selection. Specifically, in the north, a large body size is beneficial to conserve heat because of low winter temperatures, whereas in the south, it is likely due to increased food supply associated with longer growing seasons.

Canadian Journal of Zoology

Relative abundance of coyotes (Canis latrans) influences gray fox (Urocyon cinereoargenteus) occupancy across the eastern United States

Gray fox ( Urocyon cinereoargenteus (Schreber, 1775)) populations in portions of the eastern United States have experienced declines whose trajectories differ from those of other mesocarnivore populations. One hypothesis is that gray fox declines may result from interspecific interactions, particularly competition with abundant coyotes ( Canis latrans Say, 1823). Alternatively, gray foxes may respond negatively to increased urbanization and reduced forest cover. To evaluate these hypotheses, we used single-species occupancy models of camera trap data to test the effects of habitat covariates, such as the amount of urbanization and forest, on coyote and gray fox occupancy. Additionally, we test the effect of an index based on an N -mixture model of the number of coyotes at each camera trap site on gray fox occupancy. Results indicate that occupancy probabilities of coyote and gray fox relate positively to the amount of forest, but they provided no evidence urban cover impacts gray foxes. Additionally, gray fox occupancy was negatively related to the index of the number of coyotes at each site. Our models support the idea that interactions with coyotes impact gray fox occupancy across the eastern United States. These results illustrate how large-scale studies can relate mechanisms identified within specific landscapes to phenomena observed at larger scales.

Canadian Journal of Zoology

Movements of selected minnows between the lower Yellowstone River and its tributaries

Reduced population connectivity has been implicated as a cause of decreased distributions and abundances of many Great Plains fishes. However, scant empirical evidence quantifying movement and relating the contribution of spatial linkages to population abundances and resilience exists. We used otolith microchemistry analysis to characterize the movements of western silvery minnows ( Hybognathus argyritis Girard, 1856), flathead chubs ( Platygobio gracilis (Richardson, 1836)), and sand shiners ( Notropis stramineus (Cope, 1865)) between the Yellowstone River and its tributaries. Sixty-nine percent of western silvery minnows, 65% of flathead chubs, and 42% of sand shiners moved between the Yellowstone River and tributaries. Mean total number of interchanges was highest among western silvery minnows (4.8 interchanges/mover), intermediate among flathead chubs (4.3 interchanges/mover), and lowest among sand shiners (1.4 interchanges/mover; P < 0.01). Natal movements were rare, but juvenile movements were common and frequent among all three species. Movements between main-stem and tributary habitats are probably prominent facets of the life cycles of other Great Plains minnows. Therefore, connectivity among such habitats should be a high conservation priority to enhance the long-term viability of such fishes.

Great Plains, Yellowstone River

Ecological correlates of fecal corticosterone metabolites in female Greater Sage-Grouse (Centrococercus urophasianus)

Measurement of physiological responses can reveal effects of ecological conditions on an animal and correlate with demographic parameters. Ecological conditions for many animal species have deteriorated as a function of invasive plants and habitat fragmentation. Expansion of juniper (Juniperus spp.) trees and invasion of annual grasses into sagebrush (Artemisia spp.) ecosystems have contributed to habitat degradation for Greater Sage-Grouse (Centrococercus urophasianus (Bonaparte, 1827); hereafter, “Sage-Grouse”), a species of conservation concern throughout its range. We evaluated relationships between habitat use in a landscape modified by juniper expansion and annual grasses and corticosterone metabolite levels (stress responses) in feces (FCORTm) of female Sage-Grouse. We used remotely sensed data to estimate vegetation cover within hens’ home ranges and accounted for factors that influence FCORTm in other vertebrates, such as age and weather. We collected 36 fecal samples from 22 radio-collared hens during the brood-rearing season (24 May–26 July) in southwestern Idaho 2017–18. Concentrations of corticosterone increased with home range size but decreased with reproductive effort and temperature. The importance of home range size suggests that maintaining or improving habitats that promote smaller home ranges would likely facilitate a lower stress response by hens, which should benefit Sage-Grouse survival and reproduction.

Idaho

Gray wolf (Canis lupus) predation patterns following recent recolonization in a multi-predator, multi-prey system

Predator–prey interactions are among the most fundamental of ecological relationships. Recolonizing gray wolf ( Canis lupus Linnaeus, 1758) populations present new challenges for wildlife management in multi-prey, multi-carnivore systems. We documented diet composition and kill rates for wolves in a recently recolonized area over winter and summer seasons (2014–2015). Elk ( Cervus canadensis Erxleben, 1777) were the primary ungulate prey (63%) located at wolf kill sites. Deer (mule deer ( Odocoileus hemionus (Rafinesque, 1817)) and white-tailed deer ( Odocoileus virginianus (Zimmermann, 1780))) were less prevalent than elk in wolf diets, but the amount of deer in diets (40%–50%) varied by pack and season. Juvenile elk were the most prevalent class of prey in wolf diets during summer (63.3%) and winter (36.3%), with adult elk (32.5%) observed nearly as often as juveniles in winter. Kill rates varied by season, with rates 2.3 times higher in summer (mean = 3.5 ungulates/week per pack) than winter (mean = 1.5 ungulates/week per pack), consistent with increased availability and use of neonate prey. Prey biomass acquisition did not vary by pack or season (summer = 243 kg/week per pack; winter = 182 kg/week per pack). Our study quantified predation patterns for a recolonizing wolf population, and patterns that we documented were similar to other multi-prey systems in North America.

Oregon

Estimating abundance, temporary emigration and the pattern of density dependence in a cyclic snowshoe hare population in Yukon, Canada

Estimates of demographic parameters based on capture-mark-recapture (CMR) methods may be biased when some individuals in the population are temporarily unavailable for capture (temporary emigration). We estimated snowshoe hare abundance, apparent survival, and probability of temporary emigration in a population of snowshoe hares (Lepus americanus Erxleben 1777) in the Yukon using Pollock’s robust design CMR model, and population density using spatially-explicit CMR models. Survival rates strongly varied among cyclic phases, seasons, and across five population cycles. We found strong evidence that temporary emigration was Markovian (i.e., non-random), suggesting that it varied among individuals that were temporary emigrant in the previous sampling period and those that were present in the sampled area. The probability of temporary emigration for individuals that were in the study area during the previous sampling occasion (γ´´) varied among cycles. Probability that individuals that were temporarily absent from the sampled area would remain temporary emigrants (γ´) showed strongly seasonal pattern, low in winter and high during summers. Snowshoe hare population density ranged from 0.017 (0.015–0.05) hares/ha to 4.43 (3.90–5.00) hares/ha and large-scale cyclical fluctuation. Autocorrelation functions and autoregressive analyses revealed that our study population exhibited statistically significant cyclic fluctuations, with a periodicity of 9-10 years.

Yukon

The relative importance of cisco (Coregonus artedi) in the diets of common loons (Gavia immer) among a set of cisco refuge lakes in Minnesota

Common loon ( Gavia immer (Brünnich, 1764)) foraging patterns and the relative importance of cisco ( Coregonus artedi Lesueur, 1818) in the diets of loons were evaluated for the Whitefish Chain of Lakes, a set of coldwater cisco refuge lakes in Minnesota, USA. Environmental DNA metabarcoding of loon fecal samples detected 15 fish species. Yellow perch ( Perca flavescens (Mitchill, 1814)), mimic shiner ( Notropis volucellus (Cope, 1865)), and cisco were the most prominent prey detected across the study lakes. We observed a shift in fish species consumed, with increases in detections of mimic shiner and cisco DNA among loon fecal samples collected in August and September. In some locations, suitable oxythermal habitat became restricted throughout the summer, forcing cisco into surface waters, which potentially increased their vulnerability to loon predation. Conversely, large foraging aggregations of loons were observed during late summer through fall at locations with ample oxythermal habitat and abundant cisco populations. We hypothesize that cisco were sought by loons as a high-calorie prey resource prior to migration. Conservation efforts directed at preserving water quality in important cisco refuge lakes are likely to benefit common loons through enhancement of both the forage base, for resident and migrating birds, and breeding habitat suitability.

Minnesota

Structure of breeding calls in three closely related bird species (Calidris Merrem, 1804; Scolopacidae)

We studied calls of three sandpiper species to document species’ similarities and differences. We hypothesized that functionally different calls would differ in degree of divergence. We studied two nuptial calls (complex “Song” and rhythmically repeated aerial call, RRC) of males, and a general-purpose call (“alarm” trill, AT) given by parents of both sexes in the presence of humans and other potential predators, in great knot, Calidris tenuirostris (Horsfeld, 1821); surfbird, Calidris virgata (Gmelin, 1789); and red knot, Calidris canutus (Linnaeus, 1758). Calls diverged unevenly across species—RRCs and Song diverged the most and ATs the least. Vocalizations of great knot and surfbird were most similar to one another, in agreement with a recently proposed phylogeny. Despite species differences in single acoustic traits, calls were evolutionarily conservative at higher structural levels, such as rhythmic temporal delivery of RRCs and harmonic structure (e.g., the fundamental frequency was suppressed in some call types). Some acoustic qualities that differed across species were similar across call types within species (e.g., tonality in red knot calls). Trait similarity across different calls suggests that a species’ calls cannot evolve independently of one another: common mechanisms of vocal production across different calls may impede differentiation within a species’ repertoire.

Canadian Journal of Zoology

Using the electron transport system as an indicator of organismal thermal tolerance and respiratory exploitation

Freshwater ecosystems are undergoing rapid thermal shifts, making it increasingly important to understand species-specific responses to these changes. Traditional techniques for determining a species’ thermal tolerance are often lethal and time consuming. Using the enzyme activity associated with the electron transport system (ETS; hereafter referred to as enzyme assay) may provide a non-lethal, rapid, and efficient alternative to traditional techniques for some species. We used largemouth bass Micropterus salmoides (Lacepede, 1802) to test the efficacy of using an enzyme assay to determine thermal tolerance and respiratory exploitation in response to variable acclimation temperatures. Three tissue types were dissected from fish acclimated to 20, 25, or 30 °C and used in ETS assays at temperatures ranging from 7.5 to 40 °C. While there were significant differences among tissue types and acclimation temperatures, maximal enzyme activity occurred from 25.23 to 31.91 °C. Fish lost equilibrium at 39–42 °C in traditional CT max trials, significantly higher than the upper optimum range determined via enzyme assays. The ratio of enzyme activity to measured whole organism respiration rate decreased with increasing water temperature, with the largest changes occurring at the upper optimum thermal range determined by enzyme assays. Our results indicate that ETS analysis may prove useful for obtaining biologically relevant thermal tolerances.

Canadian Journal of Zoology

Effect of feeding history on metabolic rate of largemouth bass (Micropterus nigricans): implications for bioenergetics models

Metabolic rate is a key parameter in fish energy budgets that strongly influences the output of bioenergetics models. In this study, we tested the hypothesis that metabolic rate varies with growth history of age-1 largemouth bass Micropterus nigricans Cuvier, 1828. Two groups of fish were fed alternating maintenance or ad libitum rations of fathead minnow Pimephales promelas Rafinesque, 1820, so that over a 9-week period, initial and ending size of fish was similar. After 9 weeks, oxygen consumption was measured using static, closed respirometry. Although final body weight was similar between the two groups (means, 104–108 g), specific oxygen consumption for fish fed maintenance rations (0.094 mg O 2 g −2 h −1 ) was 38% less than that measured for fish fed ad libitum (0.152 mg O 2 g −2 h −1 ). Bioenergetics estimates of food consumption were similar to observed values for fish fed ad libitum (∼7% error), but for fish fed maintenance rations, the model overestimated food consumption by 65%. By accounting for changes in metabolic rate owing to reduced feeding, error in model estimates of food consumption was reduced. These findings shed new insight into factors associated with consumption-dependent error in bioenergetics models and highlight the importance of feeding history on metabolic rate of fish. Incorporating growth-dependent metabolism into bioenergetics models can improve model accuracy and allow fisheries biologists to make more informed decisions regarding fish growth and energetics.

Canadian Journal of Zoology