Causes and consequences of winter mortality in fishes

Winter mortality has been documented in a large number of freshwater fish populations, and a smaller, but increasing, number of marine and estuarine fishes. The impacted populations include a number of important North American and European resource species, yet the sources of winter mortality remain unidentified in most populations where it has been documented. Among the potential sources, thermal stress and starvation have received the most research attention. Other sources including predation and pathogens have significant impacts but have received insufficient attention to date. Designs of more recent laboratory experiments have reflected recognition of the potential for interactions among these co-occurring stressors. Geographic patterns in winter mortality are, in some cases, linked to latitudinal clines in winter severity and variability. However, for many freshwater species in particular, the effects of local community structure (predators and prey) may overwhelm latitudinal patterns. Marine (and estuarine) systems differ from freshwater systems in several aspects important to overwintering fishes, the most important being the lack of isolating barriers in the ocean. While open population boundaries allow fish to adopt migration strategies minimizing exposure to thermal stresses, they may retard rates of evolution to local environments. Geographic patterns in the occurrence and causes of winter mortality are ultimately determined by the interaction of regional and local factors. Winter mortality impacts population dynamics through episodic depressions in stock size and regulation of annual cohort strength. While the former tends to act in a density-independent manner, the latter can be density dependent, as most sources of mortality tend to select against the smallest members of the cohort and population. Most stock assessment and management regimes have yet to explicitly incorporate the variability in winter mortality. Potential management responses include postponement of cohort evaluation (to after first winter of life), harvest restrictions following mortality events and habitat enhancement. Future research should place more emphasis on the ecological aspects of winter mortality including the influences of food-web structure on starvation and predation. Beyond illuminating an understudied life-history phase, studies of overwintering ecology are integral to contemporary issues in fisheries ecology including ecosystem management, habitat evaluation, and impacts of climate change.     

Effects of black bear relocation on elk calf recruitment at Great Smoky Mountains National Park

Previous research from 2001 to 2006 on an experimentally released elk (Cervus elaphus) population at Great Smoky Mountains National Park (GSMNP or Park) indicated that calf recruitment (i.e., calves reaching 1 yr of age per adult female elk) was low (0.306, total SE = 0.090) resulting in low or negative population growth (λ = 0.996, 95% CI = 0.945–1.047). Black bear (Ursus americanus) predation was the primary calf mortality factor. From 2006 to 2008, we trapped and relocated 49 bears (30 of which were radiocollared) from the primary calving areas in the Park and radiomonitored 67 (28 M:39 F) adult elk and 42 calves to compare vital rates and population growth with the earlier study. A model with annual calf recruitment rate correlating with the number of bears relocated each year was supported (ΔAIC_c = 0.000; β = 0.070, 95% CI = 0.028–0.112) and a model with annual calf recruitment differing from before to during bear relocation revealed an increase to 0.544 (total SE = 0.098; β = −1.092, 95% CI = −1.180 to −0.375). Using vital rates and estimates of process standard errors observed during our study, 25-yr simulations maintained a mean positive growth rate in 100% of the stochastic trials with λ averaging 1.118 (95% CI = 1.096–1.140), an increase compared with rates before bear relocation. A life table response experiment revealed that increases in population growth were mostly (67.1%) due to changes in calf recruitment. We speculate that behavioral adaptation of the elk since release also contributed to the observed increases in recruitment and population growth. Our results suggest that managers interested in elk reintroduction within bear range should consider bear relocation as a temporary means of increasing calf recruitment. © 2011 The Wildlife Society.     

Influence of body mass and environmental conditions on winter mortality risk of a northern ungulate: Evidence for a late‐winter survival bottleneck

1. A relationship between winter weather and survival of northern ungulates has long been established, yet the possible roles of biological (e.g., nutritional status) and environmental (e.g., weather) conditions make it important to determine which potential limiting factors are most influential.
2. Our objective was to examine the potential effects of individual (body mass and age) and extrinsic (winter severity and snowmelt conditions) factors on the magnitude and timing of mortality for adult (>2.5 years old) female white‐tailed deer (Odocoileus virginianus [Zimmerman, 1780]) during February–May in the Upper Peninsula of Michigan, USA.
3. One hundred and fifty deer were captured and monitored during 2009–2015 in two areas with varying snowfall. February–May survival ranged from 0.24 to 0.89 (mean = 0.69) across years. Mortality risk increased 1.9% with each unit increase in cumulative winter severity index, decreased 8.2% with each cumulative snow‐free day, and decreased 4.3% with each kg increase in body mass. Age and weekly snow depth did not influence weekly deer survival. Predation, primarily from coyote (Canis latrans [Say, 1823]) and wolves (Canis lupus [L., 1758]), accounted for 78% of known‐cause mortalities.
4. Our results suggest that cumulative winter severity, and possibly to a lesser degree deer condition entering winter, impacted deer winter survival. However, the timing of spring snowmelt appeared to be the most influential factor determining late‐winter mortality of deer in our study. This supports the hypothesis that nutrition and energetic demands from weather conditions are both important to northern ungulate winter ecology. Under this model, a delay of several weeks in the timing of spring snowmelt could exert a large influence on deer survival, resulting in a survival bottleneck.

     

A review of vital rates and cause‐specific mortality of elk Cervus elaphus populations in eastern North America

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Consumptive and non‐consumptive effects of wolf spiders on cucumber beetles and cucumber plant damage

Biological control research often focuses on the ability of predators to reduce pest densities and protect crops through consumption. Less studied is their ability to protect crops by altering pest behaviour (non‐consumptive effects). Lab experiments were conducted to test predation rates of striped cucumber beetles (Acalymma vittatum; Coleoptera: Chrysomelidae) and spotted cucumber beetles (Diabrotica undecimpunctata howardi; Coleoptera: Chrysomelidae) by large (>10 mm) wolf spiders (Araneae: Lycosidae). Field experiments were conducted to examine how the physical presence and/or cues of spiders impact the behaviour and mortality of A. vittatum (specialist) and D. undecimpunctata (generalist) cucumber beetles as well as growth and damage of cucumber plants (Cucumis sativus; Cucurbitaceae). A. vittatum and D. undecimpunctata adults were added to caged cucumber plants without a spider, with spider cues only (spider removed before beetle inclusion), with spider only (spider introduced to plants immediately before beetle inclusion), and with spiders and their cues present (spiders introduced 24 hr in advance of beetle inclusion). A. vittatum responded to spider cues primarily by emigrating from plants. Contrarily, D. undecimpunctata did not display obvious responses, such as reduced feeding or increased emigration, to spider foraging and/or cues. Actively foraging lycosids increased A. vittatum mortality and reduced densities of D. undecimpunctata in the field when cucumber plants were flowering. This study highlights how non‐consumptive and consumptive effects can play a role in modifying pest populations, and how these effects can vary across species and plant growth stages.     

Effects of supplemental feeding and aggregation on fecal glucocorticoid metabolite concentrations in elk

Habitat modifications and supplemental feeding artificially aggregate some wildlife populations, with potential impacts upon contact and parasite transmission rates. Less well recognized, however, is how increased aggregation may affect wildlife physiology. Crowding has been shown to induce stress responses, and increased glucocorticoid (GC) concentrations can reduce immune function and increase disease susceptibility. We investigated the effects of supplemental feeding and the aggregation that it induces on behavior and fecal glucocorticoid metabolite concentrations (fGCM) in elk (Cervus elaphus) using observational and experimental approaches. We first compared fGCM levels of elk on supplemental feedgrounds to neighboring elk populations wintering in native habitats using data from 2003 to 2008. We then experimentally manipulated the distribution of supplemental food on feedgrounds to investigate whether more widely distributed food would result in lower rates of aggression and stress hormone levels. Contrary to some expectations that fed elk may be less stressed than unfed elk during the winter, we found that elk on feedgrounds had fecal GC levels at least 31% higher than non-feedground populations. Within feedgrounds, fGCM levels were strongly correlated with local measures of elk density (r² = 0.81). Dispersing feed more broadly, however, did not have a detectable effect on fGCM levels or aggression rates. Our results suggest that increases in aggregation associated with winter feedgrounds affects elk physiology, and the resulting increases in fGCM levels are not likely to be mitigated by management efforts that distribute the feed more widely. Additional research is needed to assess whether these increases in fGCMs directly alter parasite transmission and disease dynamics. © 2011 The Wildlife Society.     

Abiotic,bottom-up,and top-down influences on recruitment of Rocky Mountain elk in Oregon: A retrospective analysis

Understanding the relative effects of the many factors that may influence recruitment of ungulates is fundamental to managing their populations. Over the last 4 decades, average recruitment in some populations of elk (Cervus elaphus) in Oregon, USA declined from >50 to <20 juveniles per 100 females, and several competing hypotheses address these declines. We developed a priori models and constructed covariates spanning 1977–2005 from hunter-killed elk, elk population estimates, cougar harvest, and weather statistics to evaluate abiotic, bottom-up, and top-down factors that may explain annual variation and long-term trends of pregnancy, juveniles-at-heel in late autumn, and recruitment of juvenile elk in spring. In models of pregnancy status, August precipitation, age, and cougar index had positive effects, whereas previous year (t − 1) winter severity or winter precipitation_(t−1) and elk density had negative effects. In models of juvenile-at-heel in late autumn, August precipitation, August precipitation_(t−1), cougar index × elk density_(t−1), and age had positive effects, whereas cougar index, elk density_(t−1), and winter precipitation_(t−1) had negative effects. Juvenile recruitment was best explained by positive effects of August precipitation_(t−1), lactation rate, and cougar index × elk density_(t−1) and negative effects of cougar index and elk density_(t−1). Winter severity, precipitation, and temperature were not significant in explaining variation in elk recruitment. Annual variation in pregnancy, juvenile-at-heel, and recruitment was most influenced by August precipitation, whereas long-term trends in recruitment were most influenced by cougar densities with relatively weak effects of elk density. These results provide insight into causes of year-to-year and long-term trends of elk recruitment and provide a basis for more rigorous evaluation of factors affecting recruitment of elk. © 2012 The Wildlife Society.     

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

ABSTRACT Prey behavioral responses to predation risk in wolf-ungulate-plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry-collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf- and human-predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf- and human-predation risk were similar; however, responses to human-predation risk were stronger than responses to wolf-predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public-land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.     

Roadway mortality of barn owls in Idaho,USA

We examined the temporal, spatial, and demographic factors that influenced roadway mortality of barn owls (Tyto alba) along a 248-km stretch of Interstate 84 in southern Idaho using systematic road surveys. Counts of dead animals from surveys can be underestimated because of sampling biases; therefore, we also conducted experiments to assess the effects of search and removal bias on the estimates of roadway mortality of owls. We conducted surveys every 2 weeks over a 2-year period and detected 812 dead barn owls (unadjusted mortality rate of 1.64 owls/km/yr). After adjusting this estimate for search and removal bias, we documented mortality rates of up to 5.99 owls/km/year. Owl mortality was not random in relation to sex, age class, or location along the highway. Females and juveniles, which represent individuals more likely to disperse long distances, were killed more frequently than males and adults. During the nonbreeding season, owls were killed more often near agricultural lands than in shrub-steppe, but this pattern was not apparent during the breeding season. Owls were also killed more often on portions of the roadway closer to the Snake River canyon, perhaps because of the availability of nest and roost sites. Mortality rates differed markedly between the 2 years of study, which could have been related to variability in weather and its subsequent effect on owl productivity. Our data suggest that barn owls in this region may not persist under this level of mortality without significant immigration or management. Thus, roadway management to reduce or prevent owl use of roadways, reduce rodent populations near major roads, alert motorists to the presence of owls, or otherwise reduce the chances that vehicles and owls collide would improve barn owl survival and population persistence. © 2012 The Wildlife Society.     

Bumpus in the snake den: effects of sex, size, and body condition on mortality of red-sided garter snakes

Huge breeding aggregations of red-sided garter snakes (Thamnophis sirtalis parietalis) at overwintering dens in Manitoba provide a unique opportunity to identify sources of mortality and to clarify factors that influence a snake's vulnerability to these factors. Comparisons of sexes, body sizes, and body condition of more than 1000 dead snakes versus live animals sampled at the same time reveal significant biases. Three primary sources of mortality were identified. Predation by crows, Corvus brachyrhynchos (590 snakes killed), was focussed mostly on small snakes of both sexes. Crows generally removed the snake's liver and left the carcass, but very small snakes were sometimes brought back to the nest. Suffocation beneath massive piles of other snakes within the den (301 dead animals) involved mostly small males and (to a lesser extent) large females; snakes in poor body condition were particularly vulnerable. Many emaciated snakes (n = 142, mostly females) also died without overt injuries, probably due to depleted energy reserves. These biases in vulnerability are readily interpretable from information on behavioral ecology of the snakes. For example, sex biases in mortality reflect differences in postemergence behavior and locomotor capacity, the greater attractiveness of larger females to males, and the high energy costs of reproduction for females.     

Effects of the botanical composition and weather conditions on mycotoxins in winter forage from grassland

The paper focuses on possible effects of the botanical composition and weather conditions on the extend of zearalenone and ochratoxin A concentrations in various grass swards in winter pasture systems. Zearalenone is only detectable in pure stands ofLolium perenne or inLolio-Cynosuretum plant communities, respectively. The occurrence of ochratoxin A is more frequent and less specific concerning the botanical composition. This mycotoxin was found in both,Lolium perenne andFestuca arundinacea in varying years. The incidence of ochratoxin A depends on year and is apparently related to the weather conditions in autumn and winter. There was no evidence that particular locatio8/22/2008 12:57PMns have a higher or a lower risk for high ochratoxin A or zearalenone concentrations than others. Peak values in individual swards are not intermittent over the years. Presented at the 25th Mykotoxin-Workshop in Giessen, Germany, May 19–21, 2003