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Effects of hardness and alkalinity in culture and test waters on reproduction of Ceriodaphnia dubia

Ceriodaphnia dubia were cultured in four reconstituted water formulations with hardness and alkalinity concentrations ranging from soft to the moderately hard water that is required by whole-effluent toxicity (WET) testing methods for culturing test organisms. The effects of these culture formulations alone and in combination with two levels of Cl-, SO42, and HCO3- on reproduction of C. dubia were evaluated with the standard three-brood test. Reproduction was significantly reduced when test waters had lower hardness than culture waters. However, reproduction was not significantly different when animals cultured in low-hardness waters were exposed to moderately hard waters. The hardness of the culture water did not significantly affect the sensitivity of C. dubia to the three anions. Conversely, increased hardness in test waters significantly reduced the toxicities of Cl- and SO42-, with HCO3- toxicity following the same pattern. Alkalinity exhibited no consistent effect on Cl- and SO42- toxicity. The physiological stress of placing animals cultured in moderately hard water into softer test waters might contribute to marginal failures of otherwise nontoxic effluents. The standard WET protocol should be revised to allow the culture of C. dubia under lower hardness conditions to better represent local surface water chemistries.

Environmental Toxicology and Chemistry

Estimation by capture-recapture of recruitment and dispersal over several sites

Dispersal in animal populations is intimately linked with accession to reproduction, i.e. recruitment, and population regulation. Dispersal processes are thus a key component of population dynamics to the same extent as reproduction or mortality processes. Despite the growing interest in spatial aspects of population dynamics, the methodology for estimating dispersal, in particular in relation with recruitment, is limited. In many animal populations, in particular vertebrates, the impossibility of following individuals over space and time in an exhaustive way leads to the need to frame the estimation of dispersal in the context of capture-recapture methodology. We present here a class of age-dependent multistate capture-recapture models for the simultaneous estimation of natal dispersal, breeding dispersal, and age-dependent recruitment. These models are suitable for populations in which individuals are marked at birth and then recaptured over several sites. Under simple constraints, they can be used in populations where non-breeders are not observed, as is often the case with colonial waterbirds monitored on their breeding grounds. Biological questions can be addressed by comparing models differing in structure, according to the generalized linear model philosophy broadly used in capture-recapture methodology. We illustrate the potential of this approach by an analysis of recruitment and dispersal in the roseate tern Sterna dougallii .

Oikos

Modeling spatiotemporal abundance and movement dynamics using an integrated spatial capture–recapture movement model

Animal movement is a fundamental ecological process affecting the survival and reproduction of individuals, the structure of populations, and the dynamics of communities. Methods to quantify animal movement and spatiotemporal abundances, however, are generally separate and therefore omit linkages between individual-level and population-level processes. We describe an integrated spatial capture–recapture (SCR) movement model to jointly estimate (1) the number and distribution of individuals in a defined spatial region and (2) movement of those individuals through time. We applied our model to a study of polar bears ( Ursus maritimus ) in a 28,125 km 2 survey area of the eastern Chukchi Sea, USA in 2015 that incorporated capture–recapture and telemetry data. In simulation studies, the model provided unbiased estimates of movement, abundance, and detection parameters using a bivariate normal random walk and correlated random walk movement process. Our case study provided detailed evidence of directional movement persistence for both male and female bears, where individuals regularly traversed areas larger than the survey area during the 36-day study period. Scaling from individual- to population-level inferences, we found that densities varied from <0.75 bears/625 km 2 grid cell/day in nearshore cells to 1.6–2.5 bears/grid cell/day for cells surrounded by sea ice. Daily abundance estimates ranged from 53 to 69 bears, with no trend across days. The cumulative number of unique bears that used the survey area increased through time due to movements into and out of the area, resulting in an estimated 171 individuals using the survey area during the study (95% credible interval 124–250). Abundance estimates were similar to a previous multiyear integrated population model using capture–recapture and telemetry data (2008–2016; Regehr et al., Scientific Reports 8:16780, 2018). Overall, the SCR–movement model successfully quantified both individual- and population-level space use, including the effects of landscape characteristics on movement, abundance, and detection, while linking the movement and abundance processes to directly estimate density within a prescribed spatial region and temporal period. Integrated SCR–movement models provide a generalizable approach to incorporate greater movement realism into population dynamics and link movement to emergent properties including spatiotemporal densities and abundances.

eastern Chukchi Sea

The challenge and opportunity of recovering wolf populations

The gray wolf once inhabited a wide variety of habitats throughout most of the northern hemisphere north of 20°N latitude. Because the animal preyed on livestock and competed with humans for wild prey, it was extirpated from much of its range outside of wilderness areas. Environmental awareness in the late 1960s brought for the wolf legal protection, increased research, and favorable media coverage. The species has increased in both Europe and North America, is beginning to reoccupy semiwilderness and agricultural land, and is causing increased damage to livestock. Because of the wolf's high reproductive rate and long dispersal tendencies, the animal can recolonize many more areas. In most such areas control will be necessary, but the same public sentiments that promoted wolf recovery reject control. If wolf advocates could accept control by the public rather than by the government, wolves could live in far more places. Insistence on government control discourages some officials and government agencies from promoting recovery. The use of large- or small-scale zoning for wolf management may help resolve the issue. Public education is probably the most effective way to minimize the problem and maximize wolf recovery, but the effort must begin immediately.

Conservation Biology

Linking animals aloft with the terrestrial landscape

Despite using the aerosphere for many facets of their life, most flying animals (i.e., birds, bats, some insects) are still bound to terrestrial habitats for resting, feeding, and reproduction. Comprehensive broad-scale observations by weather surveillance radars of animals as they leave terrestrial habitats for migration or feeding flights can be used to map their terrestrial distributions either as point locations (e.g., communal roosts) or as continuous surface layers (e.g., animal densities in habitats across a landscape). We discuss some of the technical challenges to reducing measurement biases related to how radars sample the aerosphere and the flight behavior of animals. We highlight a recently developed methodological approach that precisely and quantitatively links the horizontal spatial structure of birds aloft to their terrestrial distributions and provides novel insights into avian ecology and conservation across broad landscapes. Specifically, we present case studies that (1) elucidate how migrating birds contend with crossing ecological barriers and extreme weather events, (2) identify important stopover areas and habitat use patterns of birds along their migration routes, and (3) assess waterfowl response to wetland habitat management and restoration. These studies aid our understanding of how anthropogenic modification of the terrestrial landscape (e.g., urbanization, habitat management), natural geographic features, and weather (e.g., hurricanes) can affect the terrestrial distributions of flying animals.

Book chapter

Davis Pond freshwater prediversion biomonitoring study: freshwater fisheries and eagles

In January 2001, the construction of the Davis Pond freshwater diversion structure was completed by the U.S. Army Corps of Engineers. The diversion of freshwater from the Mississippi River is intended to mitigate saltwater intrusion from the Gulf of Mexico and to lessen the concomitant loss of wetland areas. In addition to the freshwater inflow, Barataria Bay basin would receive nutrients, increased flows of sediments, and water-borne and sediment-bound compounds. The purpose of this biomonitoring study was, therefore, to serve as a baseline for prediversion concentrations of selected contaminants in bald eagle ( Haliaeetus leucocephalus ) nestlings (hereafter referred to as eaglets), representative freshwater fish, and bivalves. Samples were collected from January through June 2001. Two similarly designed postdiversion studies, as described in the biological monitoring program, are planned. Active bald eagle nests targeted for sampling eaglet blood (n = 6) were generally located southwest and south of the diversion structure. The designated sites for aquatic animal sampling were at Lake Salvador, at Lake Cataouatche, at Bayou Couba, and along the Mississippi River. Aquatic animals representative of eagle prey were collected. Fish were from three different trophic levels and have varying feeding strategies and life histories. These included herbivorous striped mullet ( Mugil cephalus ), omnivorous blue catfish ( Ictalurus furcatus ), and carnivorous largemouth bass ( Micropterus salmoides ). Three individuals per species were collected at each of the four sampling sites. Freshwater Atlantic rangia clams ( Rangia cuneata ) were collected at the downstream marsh sites, and zebra mussels ( Dreissena spp.) were collected on the Mississippi River. The U.S. Geological Survey (USGS) Biomonitoring of Environmental Status and Trends (BEST) protocols served as guides for fish sampling and health assessments. Fish are useful for monitoring aquatic ecosystems because they accumulate pesticides and other contaminants. Biomarker data on individual fish, generated at the USGS National Wetlands Research Center (Lafayette, La.), included percent white blood cells in whole blood, spleen weight to body weight ratio, liver weight to body weight ratio, condition factor, splenic macrophage aggregates, and liver microsomal 7-ethoxyresorufin-o-deethylase (EROD) activity. Fish age was estimated by comparing total lengths with values from the same species in the Southeast United States as determined from the literature. Contaminant analyses were coordinated by the U.S. Fish and Wildlife Service (USFWS) Analytical Control Facility (Laurel, Md.), where residues of organochlorine (OC) pesticides, total polychlorinated biphenyls (PCBs), polycyclic aromatic hydrocarbons (PAHs), aliphatic hydrocarbons (AHs), and trace elements were determined. The organic contaminant data were generated at the Mississippi State University Chemical Lab (Mississippi State, Miss.), and the inorganic contaminant data were generated by the Texas A&M University Geochemical and Environmental Research Group (College Station, Tex.). Statistical tests were performed to assess relationships among contaminants, fish age, fish species, and collection sites. Trends in interspecific differences among fish in concentrations of contaminants were noted. Striped mullet (hereafter mullet) frequently displayed the highest chemical concentrations. Levels of contaminants were generally higher in samples obtained from the Mississippi River than in those collected from the diversion area and were higher in mussels and clams (hereafter bivalves) than in fish. Because the Mississippi River sampling site for mullet and largemouth bass was downriver of the structure and south of New Orleans and the catfish site was upriver, the downriver data may not be directly reflective of the results from the receiving waters at the Davis Pond structure. Compared to the Caernarvon freshwater prediversion study in 1990 that assessed possible influx of contaminants with the freshwater diversion, contaminant levels in fishes and bivalves in this study were generally lower, yet three nontoxic inorganic elements in Davis Pond fish samples exhibited ranges of concentrations that were more than two times higher than did those from Caernarvon. Levels in bivalves were different between diversions but about equal in the numbers of trace elements showing high levels per location. Contaminant values were compared to those listed in various literature and agency sources, both regional and national, including the National Contaminant Biomonitoring Program (NCBP), in which the 85th percentile and above represents what is considered to be an elevated contaminant concentration and cause for concern. Generally, bivalves were at the high end of their ranges for both organic and inorganic contaminants. In this study, OCs were detectable in 67 percent of fish from the Mississippi River site, ranging from 0.15 to 1.09 μg/g wet weight (ww) or fresh weight (fw), and in 11 percent of the fish from the marsh sites, ranging from 0.06 to 0.612 μg/g ww. Bivalves from the Mississippi River had OC levels of 0.096 μg/g ww, whereas none were detectable in bivalves at the marsh sites. In this study, p,p ’-dichlorodiphenyldichloroethylene ( p,p’ -DDE) (a biodegradation product of DDT [dichlorodiphenyl trichloroethane]) and total PCBs were the most frequently detected OCs and were primarily from the Mississippi River. For total OC content, using adjusted least squares means, some significant interactions were noted between fish species and sites. PAHs were detected in aquatic animals at all sites (range of 0.017–17.534 μg/g ww), as were AHs (range of 0.423–4.549 μg/g ww); the highest levels of PAHs and AHs were found in bivalves from the Mississippi River. When analysis of variance (α = 0.05) was performed with data from aquatic animals, there were only two significant relationships between PAHs, AHs, and OCs between species, site, and age or the interaction among these variables. There was an interaction between fish species and n-decane (an AH) in that mullet and largemouth bass had significantly higher levels than did catfish ( P = 0.0175). When general linear means were used to investigate associations of inorganic contaminants among fish species, site, and age or any interactions among these variables, no significant results were noted for arsenic, cadmium, lead, beryllium, boron, molybdenum, or nickel. The range of mercury in fish in this study was 0.04–0.14 μg/g ww (0.14– 0.48 μg/g dry weight [dw]), with the most elevated levels detected in predatory largemouth bass at the sampling point farthest downstream from the structure and within the marsh area. Mercury was positively correlated with fish age ( P = 0.0152), where levels were estimated to increase 0.0253 parts per million (ppm) dw per year. In the Mississippi River, catfish showed significantly higher levels of mercury than did mullet or largemouth bass ( P = 0.00167). Among fish species, mullet displayed the highest levels in fish of aluminum, barium, manganese, and iron, all considered to have low toxicity in hydrologic systems. An interaction between fish and site was seen with aluminum ( P = 0.0031), where concentrations in mullet were significantly higher in the Mississippi River than at the other sites, as was also seen with barium ( P = 0.0009), chromium ( P = <0.0001), manganese ( P = 0.0004), strontium ( P = 0.0074), vanadium ( P = 0.0156), and zinc ( P = 0.0059). For iron ( P = 0.0.0001), mullet and largemouth bass at both the Mississippi River and Lake Salvador showed higher levels than did catfish, and these two species showed higher levels at two of the four sites. An interaction between fish and site was also seen with chromium ( P = <0.0001) in that concentrations in mullet were significantly higher in the Mississippi River than at the other sites, as was also seen with strontium ( P = 0.0074), vanadium ( P = 0.0156), and zinc ( P = 0.0059), metals for which deleterious effects have been demonstrated in other ecosystems. The NCBP program lists the 85th percentile for zinc at 34.2 μg/g fw (117.9 μg/g dw). In the Davis Pond prediversion biomonitoring study (hereafter the current study), one fish (MUL31RIVER, fish ID 8) showed values higher than that (125.4 μg/g dw or 37.54 μg/g ww), and the Mississippi River bivalve sample (MUSSRIVER) had a value of 140 μg/g dw (41.2 μg/g ww). In the current study, approximately 86 percent of the fish had measurable selenium levels, yet none reached the 85th percentile. The 85th percentile for selenium from the NCBP was 0.73 μg/g ww. Significantly higher levels of selenium were seen in mullet than in largemouth bass and catfish ( P = 0.0023). The NCBP 85th percentile for lead is 0.22 μg/g ww (0.76 μg/g dw). In the current study, the range of concentrations of lead was as much as 18.3 ppm dw (MUL31RIVER, fish ID 8), with the three most elevated values (range of 3.46–5.31 μg/g ww) coming from mullet from the Mississippi River. Biomarker data are measurable and directly reflect the condition of the animal, and measuring more than one biomarker in an individual increases confidence in health assessments. In the current study, biomarkers included macrophage aggregates (MAs), liver (hepatosomatic index [HSI]) and spleen (splenosomatic index [SSI]) weight to body weight ratios, percent white blood cells (WBCs) in whole blood, and condition factor. Few significant differences were noted with any of the biomarkers between sites, and there were no relationships between species and sites. For improved use of biomarker assessments, an increase in fish sample size would be useful for postdiversion sampling, as would comparisons of fish of the same sex and reproductive condition. During the current study, success for eagle nests in the diversion area and reference sites was similar as determined by numbers of nestlings fledged. When temperatures were below average during winter 2000, nests in both regions similarly failed. At each nest, the primary evidence of food items was small mammals. Eaglets (n = 6) generally appeared healthy, and whole blood concentrations of organic contaminants exceeded detection limits with three incidences of p,p’ -DDE (0.002–0.006 μg/L ww) and one incidence of oxychlordane (0.002 μg/L ww). The levels of p,p’ -DDE were well below those that have been inversely correlated with productivity and success rates of nesting bald eagles on a regional scale. The low values found in the whole blood samples for OC pesticides and PCBs were even lower when corrected for plasma volume. Aluminum values were 3.66 and 5.75 μg/L in two samples, zinc ranged from 5.21 to 6.77 μg/L ww in six samples, and silicon ranged from 1.7 to 4.6 μg/L in four samples. Selenium was detectable in each bird with the range at 0.332–0.566 μg/L ww, and strontium ranged from 0.0581 to 0.0975 μg/L ww. Mercury was detectable in blood samples from each bird and ranged from 0.0254 to 0.0845 μg/L ww, whereas lead was detectable in four samples and ranged from 0.0042 to 0.0136 μg/L ww. Although no detectable levels of total PCBs were found (also correlated with decreased reproductive productivity), 70 percent of the aquatic animals from the Mississippi River contained total PCBs (range 0.13–0.79 μg/L), whereas only about 7 percent of the aquatic animals sampled from the marsh area contained PCBs. Suggestions for postdiversion sampling include lowering the analytical detection limit for some metals, sampling aquatic animals over the course of a single season, obtaining a higher sample number of mature fish of one species (for example, blue catfish) within a range of total lengths for biomarker analyses, obtaining otoliths for estimating fish ages, assessing dioxins in eaglet blood, examining triazines in water, and obtaining all Mississippi River fish samples as close to the Davis Pond structure intake as possible. Because contaminants found in blood of eaglets reflect their prey species and because of the contaminant levels found in fish in the current study, eaglets may not be consuming primarily these species; therefore, obtaining juvenile nutria ( Myocastor coypus ) or turtle species for contaminant analyses might be considered, as well as collecting greater blood volume and using plasma to measure OCs and PCBs. Data obtained postdiversion will be compared with prediversion data to monitor changes.

Louisiana

Pyometra in a Siberian Polecat (Mustela eversmanni)

A 2-year-old Siberian polecat (Mustela eversmanni) from a breeding colony presented for ultrasound evaluation for pregnancy. It was paired with a male for 2.75 months and had remained absent of pregnancy signs when it was anesthetized and clinically evaluated. Until this time, the animal had eaten well and shown no outward signs of debility. On palpation, the animal had a fluid-filled tubular structure in the caudal abdomen, consistent in location and size with the uterus. No sign of vaginal discharge was present. Ultrasonography revealed 10 fluid-filled evaginations (approximately 12 mm in diameter) of the uterine horns. A presumptive diagnosis of a fluid-filled reproductive tract and likely reproductive failure was made in light of the animal's history, its clinical signs, and the ultrasound findings. Euthanasia was performed because the animal was nonreproductive and might yield information relevant to the breeding colony as a whole. Necropsy of the polecat revealed a distended fluctuant uterus containing mildly odiferous, thick, yellow-green, purulent material. Histopathology confirmed the diagnosis of pyometra. A pure and heavy growth of Enterococcus fecalis was cultured from the uterine contents. In light of results from routine minimal inhibitory concentration antibiotic sensitivity screening, this isolate was resistant to all antibiotics tested in the standard teaching hospital screen.

Contemporary Topics in Laboratory Animal Science

Analysis of movement recursions to detect reproductive events and estimate their fate in central place foragers

Recursive movement patterns have been used to detect behavioral structure within individual movement trajectories in the context of foraging ecology, home-ranging behavior, and predator avoidance. Some animals exhibit movement recursions to locations that are tied to reproductive functions, including nests and dens; while existing literature recognizes that, no method is currently available to explicitly target different types of revisited locations. Moreover, the temporal persistence of recursive movements to a breeding location can carry information regarding the fate of breeding attempts, but it has never been used as a metric to quantify recursive movement patterns. Here, we introduce a method to locate breeding attempts and estimate their fate from GPStracking data of central place foragers. We tested the performance of our method in three bird species differing in breeding ecology (wood stork (Mycteria americana), lesser kestrel (Falco naumanni), Mediterranean gull (Ichthyaetus melanocephalus)) and implemented it in the R package ‘nestR’. Methods: We identified breeding sites based on the analysis of recursive movements within individual tracks. Using trajectories with known breeding attempts, we estimated a set of species-specific criteria for the identification of nest sites, which we further validated using non-reproductive individuals as controls. We then estimated individual nest survival as a binary measure of reproductive fate (success, corresponding to fledging of at least one chick, or failure) from nest-site revisitation histories during breeding attempts, using a Bayesian hierarchical modeling approach that accounted for temporally variable revisitation patterns, probability of visit detection, and missing data. Results: Across the three species, positive predictive value of the nest-site detection algorithm varied between 87 and 100% and sensitivity between 88 and 92%, and we correctly estimated the fate of 86–100% breeding attempts.

Movement Ecology

Testing four hypotheses to explain partial migration: Balancing reproductive benefits with limits to fasting endurance

Seasonal migration is ubiquitous in animals, and yet its underlying cause(s) remain poorly known. Species exhibiting short-distance altitudinal migration and intraspecific variation in migratory behavior (partial or differential migration) are ideal study systems for examining the selective pressures that affect individual migratory decisions. We used an individually marked population of yellow-eyed juncos, breeding along a 1000-m elevational gradient and migrating up and down that gradient, to examine the morphological, behavioral, and reproductive traits associated with migratory behavior. We tested the four most well-known hypotheses proposed to explain partial migration: the thermal tolerance, fasting endurance, dominance, and arrival time hypotheses. Our results indicate that: (1) limits to juncos’ fasting endurance constrain their ability to overwinter at high elevations, in support of the fasting endurance hypothesis, (2) differences in body size mediate fasting ability and are associated with variation in migratory behavior and overwinter apparent survival, (3) migratory behavior interacts with reproductive success, in partial support of the arrival time hypothesis, and (4) additional mechanisms that are not captured by the four well-known hypotheses might better explain individual variation in migratory behavior. Less migratory females achieved greater nesting success the following breeding season. Among males, nesting success influenced migratory tendency the following winter. Successful males may either migrate to a more benign winter climate without paying reproductive costs, or high levels of parental effort might physiologically constrain their ability to overwinter in harsh climates.

Behavioral Ecology and Sociobiology

Informed breeding dispersal following stochastic changes to patch quality in a pond-breeding amphibian

The unidirectional movement of animals between breeding patches (i.e. breeding dispersal) has profound implications for the ecological and evolutionary dynamics of spatially structured populations. In spatiotemporally variable environments, individuals are expected to adjust their dispersal decisions according to information gathered on the environmental and/or social cues that reflect the fitness prospects in a given breeding patch (i.e. informed dispersal). A paucity of empirical work limited our understanding of the ability of animals to depart from low-quality breeding patches and settle in high-quality breeding patches. We examined the capacity of individuals to respond to stochastic changes in habitat quality via informed breeding dispersal in a pond-breeding amphibian. We conducted a 5-year (2015–2019) capture–recapture study of boreal toads Anaxyrus boreas boreas ( n = 1,100) that breed in beaver ponds in western Wyoming, USA. During early spring of 2017, an extreme flooding event destroyed several beaver dams and resulted in the loss of breeding habitat. We used multi-state models to investigate how temporal changes in pond characteristics influenced breeding dispersal, and determine whether movement decisions were in accordance with prospects for reproductive fitness. Boreal toads more often departed from low-quality breeding ponds (without successful metamorphosis) and settled in high-quality breeding ponds (with successful metamorphosis). Movement decisions were context-dependent and associated with pond characteristics altered by beaver dam destruction. Individuals were more likely to depart from shallow ponds with high vegetation cover and settle in deep ponds with low vegetation cover. The probability of metamorphosis was related to the same environmental cues, suggesting that boreal toads assess the fitness prospects of a breeding patch and adjust movement decisions accordingly (i.e. informed breeding dispersal). We demonstrated that stochastic variability in environmental conditions and habitat quality can underpin dispersal behaviour in amphibians. Our study highlighted the mechanistic linkages between habitat change, movement behaviour and prospects for reproductive performance, which is critical for understanding how wild animals respond to rapid environmental change.

Wyoming

A semelparous fish continues upstream migration when exposed to alarm cue, but adjusts movement speed and timing

Animals make trade-offs between predation risk and pursuit of opportunities such as foraging and reproduction. Trade-offs between antipredator behaviours and foraging are well suited to manipulation in laboratory and field settings and have generated a vast compendium of knowledge. However, much less is known about how animals manage trade-offs between predation risk and pursuit of reproductive opportunities in the absence of the confounding effects of foraging. In the present study, we investigated how the nonfeeding migratory life stage of sea lamprey, Petromyzon marinus , responds to odour from dead conspecifics (a cue that induces avoidance behaviours in laboratory and field studies). We released groups of PIT-tagged sea lamprey 65 m from the shore of Lake Michigan or 287 m upstream in Carp Lake River and used antennas to detect their movements in the river. As the breeding season progressed, sea lamprey initiated upstream movement earlier and were more likely to enter the river. Sea lamprey that began the night in Lake Michigan entered Carp Lake River at higher rates and accelerated upstream when exposed to high concentrations of alarm cue, consistent with animals attempting to minimize time spent in risky areas. Sea lampreys that began the night in the river delayed upstream movement when exposed to alarm cue, consistent with animals sheltering and gathering information about a source of risk. We attribute this context-specific reaction to alarm cue to differences in perceived vulnerability to predation in sheltered positions in the river versus exposed positions in the lake. Once in the river, the vast majority of sea lamprey moved upstream independent of alarm cue or Julian date. Although life-history-induced time and energy budgets place rigid constraints on the direction of migration, sea lamprey attend to predation risk by modifying movement timing and speed.

Animal Behaviour

Life‐history tradeoffs revealed by seasonal declines in reproductive traits of Arctic‐breeding shorebirds

Seasonal declines in breeding performance are widespread in wild animals, resulting from temporal changes in environmental conditions or from individual variation. Seasonal declines might drive selection for early breeding, with implications for other stages of the annual cycle. Alternatively, selection on the phenology of nonbreeding stages could constrain timing of the breeding season and lead to seasonal changes in reproductive performance. We studied 25 taxa of migratory shorebirds (including five subspecies) at 16 arctic sites in Russia, Alaska, and Canada. We investigated seasonal changes in four reproductive traits, and developed a novel Bayesian risk‐partitioning model of daily nest survival to examine seasonal trends in two causes of nest failure. We found strong seasonal declines in reproductive traits for a subset of species. The probability of laying a full four‐egg clutch declined by 8–78% in 12 of 25 taxa tested, daily nest survival rates declined by 1–12% in eight of 22 taxa, incubation duration declined by 2.0–2.5% in two of seven taxa, and mean egg volume declined by 5% in one of 15 taxa. Temporal changes were not fully explained by individual variation. Across all species, the proportion of failed nests that were depredated declined over the season from 0.98 to 0.60, while the proportion abandoned increased from 0.01 to 0.35 and drove the seasonal declines in nest survival. An increase in abandonment of late nests is consistent with a life‐history tradeoff whereby either adult mortality increased or adults deserted the breeding attempt to maximize adult survival. In turn, seasonal declines in clutch size and incubation duration might be adaptive to hasten hatching of later nests. In other species of shorebirds, we found no seasonal patterns in breeding performance, suggesting that some species are not subject to selective pressure for early breeding.

Journal of Avian Biology

Range expansion of nonindigenous caribou in the Aleutianarchipelago of Alaska

Caribou ( Rangifer tarandus ) are nonindigenous to all but the eastern-most island of the Aleutian archipelago of Alaska. In 1958–1959, caribou were intentionally introduced to Adak Island in the central archipelago, and the population has at least tripled in recent years subsequent to the closure of a naval air facility. Although dispersal of caribou to adjacent islands has been suspected, no historical documentation has occurred to date. Herein, we report consistent detections of caribou sign on the adjacent island of Kagalaska over 2 summer field seasons (2010–2011), and visual detection of caribou on that island during the summer of 2011. Ecological impacts of caribou on Kagalaska are not strongly apparent at the present time and we do not know how many animals permanently occupy the island. However, establishment of a reproductively viable resident population on Kagalaska is worrisome and could set the stage for a step-wise invasion of additional nearby islands.

Alaska

The economics of territory selection

Territorial behavior is a fundamental and conspicuous behavior within numerous species, but the mechanisms driving territory selection remain uncertain. Theory and empirical precedent indicate that many animals select territories economically to satisfy resource requirements for survival and reproduction, based on benefits of food resources and costs of competition and travel. Costs of competition may vary by competitive ability, and costs of predation risk may also drive territory selection. Habitat structure, resource requirements, conspecific density, and predator distribution and abundance are likely to further influence territorial behavior. We developed a mechanistic, spatially-explicit, individual-based model to better understand how and why animals select particular territories. The model was based on optimal selection of individual patches for inclusion in a territory according to their net value, i.e., benefits (food resources) minus costs (travel, competition, predation risk). Simulations produced predictions for what may be observed empirically if such optimization drives placement and characteristics of territories. Simulations consisted of sequential, iterative selection of territories by simulated animals that interacted to defend and maintain territories. Results explain why certain patterns in space use are commonly observed, and when and why these patterns will differ from the norm. For example, more clumped or abundant food resources are predicted to result, on average, in smaller territories with more overlap. Strongly different resource requirements for individuals or groups in a population will directly affect space use and are predicted to cause different responses under identical conditions. Territories are predicted to decrease in size with increasing population density, which can enable a population’s density of territories to change at faster rates than their spatial distribution. Due to competition, less competitive territory-holders are generally predicted to have larger territories in order to accumulate sufficient resources, which could produce an ideal despotic distribution of territories. Interestingly, territory size and overlap are predicted to show a parabolic response to increases in predator densities, and territories are predicted to be larger where predators are more clumped in distribution. Our model’s predictions are consistent with many empirical observations, providing support for optimal patch selection as a mechanism for the economical territories of animals commonly observed in nature.

Ecological Modelling

Spatially explicit dynamic N-mixture models

Knowledge of demographic parameters such as survival, reproduction, emigration, and immigration is essential to understand metapopulation dynamics. Traditionally the estimation of these demographic parameters requires intensive data from marked animals. The development of dynamic N-mixture models makes it possible to estimate demographic parameters from count data of unmarked animals, but the original dynamic N-mixture model does not distinguish emigration and immigration from survival and reproduction, limiting its ability to explain important metapopulation processes such as movement among local populations. In this study we developed a spatially explicit dynamic N-mixture model that estimates survival, reproduction, emigration, local population size, and detection probability from count data under the assumption that movement only occurs among adjacent habitat patches. Simulation studies showed that the inference of our model depends on detection probability, local population size, and the implementation of robust sampling design. Our model provides reliable estimates of survival, reproduction, and emigration when detection probability is high, regardless of local population size or the type of sampling design. When detection probability is low, however, our model only provides reliable estimates of survival, reproduction, and emigration when local population size is moderate to high and robust sampling design is used. A sensitivity analysis showed that our model is robust against the violation of the assumption that movement only occurs among adjacent habitat patches, suggesting wide applications of this model. Our model can be used to improve our understanding of metapopulation dynamics based on count data that are relatively easy to collect in many systems.

Population Ecology

Full annual cycle climate change vulnerability assessment for migratory birds

Climate change is a serious challenge faced by all plant and animal species. Climate change vulnerability assessments (CCVAs) are one method to assess risk and are increasingly used as a tool to inform management plans. Migratory animals move across regions and continents during their annual cycles where they are exposed to diverse climatic conditions. Climate change during any period and in any region of the annual cycle could influence survival, reproduction, or the cues used to optimize timing of migration. Therefore, CCVAs for migratory animals best estimate risk when they include climate exposure during the entire annual cycle. We developed a CCVA incorporating the full annual cycle and applied this method to 46 species of migratory birds breeding in the Upper Midwest and Great Lakes (UMGL) region of the United States. Our methodology included background risk, climate change exposure × climate sensitivity, adaptive capacity to climate change, and indirect effects of climate change. We compiled information about migratory connectivity between breeding and stationary non-breeding areas using literature searches and U.S. Geological Survey banding and re-encounter data. Climate change exposure (temperature and moisture) was assessed using UMGL breeding season climate and winter climate from non-breeding regions for each species. Where possible, we focused on non-breeding regions known to be linked through migratory connectivity. We ranked 10 species as highly vulnerable to climate change and two as having low vulnerability. The remaining 34 species were ranked as moderately vulnerable. In general, including non-breeding data provided more robust results that were highly individualistic by species. Two species were found to be highly vulnerable throughout their annual cycle. Projected drying will have the greatest effect during the non-breeding season for species overwintering in Mexico and the Caribbean. Projected temperature increases will have the greatest effect during the breeding season in UMGL as well as during the non-breeding season for species overwintering in South America. We provide a model for adaptive management of migratory animals in the face of projected climate change, including identification of priority species, research needs, and regions within non-breeding ranges for potential conservation partnerships.

Ecological Applications

Southern sea otter range expansion and habitat use in the Santa Barbara Channel, California

The re-colonization of the Santa Barbara channel by sea otters brings these ESA-listed marine mammals closer to active oil and gas production facilities, shipping lanes and naturally occurring oil and gas seeps. However, the degree to which sea otters may actually be affected by human-caused oil spills or exposure to natural oil seeps is currently unknown. Between 2012 and 2014, the U.S. Geological Survey and collaborating agencies conducted a telemetry-based study of sea otters in Santa Barbara channel, in order to provide critical information for resource managers (specifically the Bureau of Ocean Energy Management, henceforth BOEM, and the U.S. Fish and Wildlife Service, henceforth USFWS) about the spatial ecology, population status, and potential population threats to sea otters in Santa Barbara Channel, with particular reference to exposure to manmade structures and sources of oil and natural gas. Analysis of spatial monitoring data using a Bayesian-based synoptic model allowed for description of sea otter home ranges, identification of hot-spots of use, and insights into habitat selection behavior by male and female sea otters. Important findings included the deeper modal depth preferred by males versus females, strong preferences by both sexes for areas with persistent kelp canopy, and greater use of soft-sediment areas by males. The synoptic model also provided the ability to predict population-level density distribution for each sex in new habitats: by calculating the value of these probability density distributions at the known locations of natural seeps, we were able to identify those seeps with higher potential for sea otter encounters. The relative probability of occurrence at locations near to some seeps was sufficiently high (about 1% likelihood of occurrence for some of our study animals) that one would anticipate occasional encounters. Data on male and female survival, reproductive success, activity budgets, and body condition all indicated that sea otters in Santa Barbara Channel are not resource limited, and thus we would expect to see continued strong population growth in this area. However, the principal cause of death for study animals was lethal bites by white sharks, suggesting that shark bite mortality represents the single biggest threat to continued population growth in the Santa Barbara Channel.

California

Social composition of soft‐release groups is correlated with survival of translocated gopher tortoises

The social structure of translocated animal populations can have important effects on the survival and reproduction of translocated individuals for both solitary and social species. The gopher tortoise ( Gopherus polyphemus ) is a reptile of conservation concern that is currently experiencing high levels of mitigation translocation in Florida, USA. Individuals live in aggregations of burrows with frequent agonistic, courtship, and burrow-sharing interactions between residents. Given that exposure to many unfamiliar individuals may increase the frequency of aggressive interactions and social stress following translocation, we predicted that tortoises with greater numbers of familiar individuals co-translocated from the same origin site would have higher survival after translocation. To test this, we updated a recently published survival analysis of 2,822 translocated tortoises and 502 identified carcasses from a translocation site in the western Florida panhandle from 2006–2022. After controlling for simultaneous effects of soft-release enclosure identity, release season, release density, region of origin, sex, and size, adding the number of potentially familiar individuals improved model fit and showed increasing the number of familiars reduced the probability of being found dead. This effect was modulated by release density, being apparent only when density was high, suggesting a role for social interactions. This effect was also present only in the first few years after release, prior to the removal of soft-release enclosures preventing dispersal, and was similar in magnitude to previously identified effects of density, release season, and region of origin. We suggest that this effect may result from reduced aggressive interactions or social stress for tortoises with a greater number of familiar individuals in their release enclosures but cannot rule out the possibility of reduced novel pathogen exposure for individuals released with a greater number of individuals from the same source site or other factors that may be confounded with the size of translocated groups. Designing and implementing mitigation translocations to account for social composition of gopher tortoise groups could improve survival in release enclosures.

Florida