Article

Motorized backcountry recreation and stress response in Mountain Caribou (Rangifer tarandus caribou)

Authors:
To read the full-text of this research, you can request a copy directly from the author.

No full-text available

Request Full-text Paper PDF

To read the full-text of this research,
you can request a copy directly from the author.

... An important challenge for SMC conservation occurs during the period of their winter range residency, when they occupy high-elevation habitats that are also ideal for helicopter-assisted downhill skiing (heli-skiing). While the effects of heli-skiing on caribou are not well known (but see Freeman, 2007, Huebel, 2012, Wilson & Wilmshurst, 2019, aerial disturbance (Harrington & Veitch, 1992;Luick et al., 1996), noise (Perra et al., 2022) and encounters with humans (Nellemann et al., 2000;Whittington & Mercer, 2004) have demonstrated the effects on caribou. For these reasons, the potential of heli-skiing activities to impact caribou recovery is substantial due to the large geographic area over which heli-skiing occurs (40 000 km 2 ) and the similarity of these activities to other known disturbances. ...
... Consequently, caribou tend to occupy lowproductivity landscapes where predator densities and the risk of predation are reduced (Johnson et al., 2002), but energetic costs can be high (Fancy & White, 1987). Our results indicate that partial refugia afforded by low-productivity areas are being further constrained by back-country recreation, likely not only limited to heli-skiing (Freeman 2007, Seip, Johnson, & Watts, 2007. Given our findings that caribou ranged more widely in the absence of heli-skiing, it is possible that the smaller home ranges to which they are restricted during years of normal heli-ski operations are also affecting resource availability and carrying capacities. ...
... The impacts of this constraint on caribou fitness are currently unknown, but smaller restricted home ranges have been associated with lower survival of adult female caribou (MacNearney et al., 2016;Williams et al., 2021). Furthermore, the energetic demands of a deep snow environment compounded by a low nutrition winter diet (Rominger, Robbins, & Evans, 1996), with avoidance or flight responses to disturbance (Webster, 1997;Freeman, 2007;Mac-Nearney et al., 2016;Lesmerises et al., 2018), suggest the potential for caribou to experience resource deficits when their ranges are artificially contracted. This potential restriction of resources could cause reductions in body condition or survival. ...
Article
Full-text available
The long‐term conservation of species at risk relies on numerous, and often concurrent, management actions to support their recovery. Generally, these actions are habitat‐based while others are focused on a species' position within its ecological community. Less studied are the impacts from human presence, despite evidence that human activity may reduce the area functionally available for occupancy or resource acquisition. In the winter of 2020/2021, COVID‐19‐related travel restrictions led to a reduction in helicopter‐assisted back‐country skiing (heli‐skiing). We examined how these reductions in heli‐skiing (termed the anthropause) affected the movement ecology and resource selection of southern mountain caribou (Rangifer tarandus caribou) as compared to two prior years (2018/2019 and 2019/2020) and the following year when heli‐skiing resumed (2021/2022). We found that home‐range size was on average 80–120% larger during the anthropause than in years of normal heli‐ski operations. Movement rates also varied among periods, with movement during the anthropause (11.9 km²/day) being higher than in 2019/2020 (7.8 km²/day) and 2021/2022 (8.7 km²/day), though similar to 2018/2019 (12.2 km²/day). Resource selection among periods did not differ, with caribou consistently selecting old forests, high elevations and gentle terrain. These results suggest that back‐country recreation, specifically heli‐skiing, may be limiting access to resources for southern mountain caribou. This limitation arises through reduced home‐range size within suitable late‐winter habitat, relative to when heli‐skiing is reduced or not occurring – consistent with Encounter Theory. While the demographic effects of reduced home‐range size and movement were not examined here, reduced access to resources likely compounds other stressors known to affect population viability of caribou. The results of this study demonstrate the impact that recreation can have on wildlife and highlight the need to consider heli‐skiing and other forms of recreation when developing recovery plans.
... Caribou generally avoid anthropogenic disturbance (Dyer et al., 2001a;Johnson et al., 2015Johnson et al., , 2020a, which results in reduced available habitat, and altered movements and energetic budgets (Bradshaw et al., 1998;Polfus et al., 2011). Caribou also physically avoid noise and human activity, including motorized and non-motorized recreation (Webster, 1997;McKay, 2007;Freeman, 2008;Lesmerises et al., 2018), aircraft (Harrington and Veitch, 1991;Stinchcomb et al., 2020), and areas with high human activity like tourist resorts, mines, active roads and pipelines (Nellemann et al., 2000(Nellemann et al., , 2001Boulanger et al., 2012). Avoidance behaviour of caribou to disturbance is compounded by sensitivity to noise and human activity, and as such the influence of anthropogenic disturbances like timber harvest, roads, seismic lines, pipelines and wellsites extend far beyond the direct footprint of the development and can vary in intensity throughout the life cycle of a disturbance (Dyer et al., 2001a;Boulanger et al., 2012;Johnson et al., 2015). ...
... Combined with the acceleration of habitat loss in Alberta and British Columbia caribou ranges (Nagy-Reis et al., 2021), caribou may increasingly become concentrated in marginal habitat (Sawyer et al., 2009;MacNearney et al., 2016), with reduced access to forage (Nellemann and Cameron, 1996), greater predation risk (Wittmer et al., 2007), and in mountainous terrain, increased exposure to avalanches (Hebblewhite et al., 2009) and reductions in survival rates (Williams et al., 2021). In addition, increased encounters with humans and decreases in home range size because of anthropogenic disturbance can result in increased stress (Freeman, 2008;Ewacha et al., 2017), increased energetic expenditure, reduced body mass (Bradshaw et al., 1998), and potentially reduced calving rates (Cameron and Ver Hoef, 1994). Our study focused on central mountain caribou ranges during winter; we could not evaluate the response of caribou to wellsites during the rest of the annual period due to the migration of caribou away from disturbed winter range towards calving and summer range in protected areas where wellsites were absent. ...
Article
Full-text available
Woodland caribou (Rangifer tarandus) are threatened in Alberta in part due to the development of oil and gas resources. To inform best management practices for caribou, we assessed how proximity to wellsites influenced caribou habitat selection, and whether habitat selection varied across wellsite activity phases (i.e., drilling, producing, and inactive). We used location data from 37 GPS-collared caribou monitored between 2007 and 2013 in west-central Alberta to model habitat selection. Our results suggest the influence of wellsites on caribou habitat selection are temporally dynamic. The largest impacts occur when human activity at wellsites is greatest, however wellsites continue to influence caribou habitat selection after human activity ceases. Caribou avoided wellsites, and avoidance increased relative to the degree of activity at the nearest wellsite. During early winter, caribou avoided wellsites in the drilling phase more than inactive and producing wellsites. During late winter, caribou avoided wellsites in producing phases more than inactive wellsites. Caribou may benefit from management practices that include i) seasonal timing restrictions on drilling, ii) reductions of human activity at wellsites, whether in duration or intensity, iii) land-use planning to coordinate the placement of wellsites to minimize impacts to caribou and their habitat, and iv) prompt and effective restoration of wellsites to match original habitat conditions once production has stopped.
... She found that moose had higher fecal GC levels in areas with snowmobile activity than in areas where snowmobiles were not present. Similarly, Freeman (2008) found that mountain caribou had higher stress levels in snowmobile use areas compared to non-use areas. These hormones can affect an animal's reproductive and territorial behavior, immune function, foraging efficiency, glucose metabolism, and locomotion, all of which help an individual cope with unpredictable situations (Lynn et al. 2010). ...
... Stress in these animals is also dependent on a number of factors. Several studies have reported the benefits of fecal GC analysis as a non-invasive measure of the effects of human disturbance on wildlife (Tomeo 2000, Creel et al. 2002, Möstl and Palme 2002, Freeman 2008, Sherriff et al. 2011, Wasser et al. 2011). However, a few publications have raised questions on the validity of fecal GC metabolites as an effective metric of wildlife hormonal responses to specific stimuli due to the variety of factors that influence fecal GC levels (Millspaugh and Washburn 2004, Touma and Palme 2005, Goyman 2012 ...
Thesis
Full-text available
Snowmobiling is a popular winter activity in northern regions of North America. Although snowmobiles are important utility vehicles and serve as a means of outdoor recreation, their activity is known to affect plants and animals. These effects have been a growing concern over the past 20 years as a result of increased snowmobile activity into once inaccessible natural areas. Minimizing the impacts of snowmobiles on biota, preserving the quality and character of wilderness areas, and providing adequate access to snowmobilers for traditional activities has been a challenge for public land managers in Alaska. To address the effects that snowmobiles have on ecological systems at site-specific and landscape-level scales, I conducted a study in the Kenai National Wildlife Refuge, a boreal ecosystem located in southcentral Alaska, to determine 1) the response of woody wetland plants to snowmobile traverses at varying snow depths, 2) the temporal and spatial variation of a winter soundscape with emphasis on anthrophony, in general, and snowmobile noise, specifically, 3) the effects of snowmobile noise on wilderness character and naturalness, and 4) the spatial and physiological response of moose to snowmobile activity and noise. I used a combination of traditional experimental designs and statistics, machine learning, and spatially-explicit predictive modeling to assess the effects snowmobile activity has on these four components. I found that snowmobile activity reduced the number of living stems and inhibited the growth of woody wetland plants by direct contact with protruding vegetation above the snow and indirectly from snow compaction. Snowmobile noise was not a large contributor of noise to the soundscape but was pervasive in remote natural areas. Snowmobile noise affected a significant area of Congressionally-designated wilderness altering the naturalness and character of the wilderness soundscape. Moose exhibited a distinct spatial partitioning and avoidance from snowmobile activity and developed areas (i.e., oil and gas compressors) at the landscape-level but at a site-specific scale snowmobile traffic and noise had no apparent affect on the stress hormone levels of moose that were selecting habitats close to snowmobile trails. I detected these impacts at both site-specific locations and across large spatial scales indicating that snowmobile effects are more than just localized disturbances. Based on these findings, I conclude that snowmobile noise and activity is an additional and unnatural forcing function on a boreal ecosystem already stressed by the harsh environmental conditions of winter.
... She found that moose had higher fecal GC levels in areas with snowmobile activity than in areas where snowmobiles were not present. Similarly, Freeman (2008) found that mountain caribou had higher stress levels in snowmobile use areas compared to non-use areas. These hormones can affect an animal's reproductive and territorial behavior, immune function, foraging efficiency, glucose metabolism, and locomotion, all of which help an individual cope with unpredictable situations (Lynn et al. 2010). ...
... Stress in these animals is also dependent on a number of factors. Several studies have reported the benefits of fecal GC analysis as a non-invasive measure of the effects of human disturbance on wildlife (Tomeo 2000, Creel et al. 2002, Möstl and Palme 2002, Freeman 2008, Sherriff et al. 2011, Wasser et al. 2011). However, a few publications have raised questions on the validity of fecal GC metabolites as an effective metric of wildlife hormonal responses to specific stimuli due to the variety of factors that influence fecal GC levels (Millspaugh and Washburn 2004, Touma and Palme 2005, Goyman 2012 ...
... She found that moose had higher fecal GC levels in areas with snowmobile activity than in areas where snowmobiles were not present. Similarly, Freeman (2008) found that mountain caribou had higher stress levels in snowmobile use areas compared to non-use areas. These hormones can affect an animal's reproductive and territorial behavior, immune function, foraging efficiency, glucose metabolism, and locomotion, all of which help an individual cope with unpredictable situations (Lynn et al. 2010). ...
... Stress in these animals is also dependent on a number of factors. Several studies have reported the benefits of fecal GC analysis as a non-invasive measure of the effects of human disturbance on wildlife (Tomeo 2000, Creel et al. 2002, Möstl and Palme 2002, Freeman 2008, Sherriff et al. 2011, Wasser et al. 2011). However, a few publications have raised questions on the validity of fecal GC metabolites as an effective metric of wildlife hormonal responses to specific stimuli due to the variety of factors that influence fecal GC levels (Millspaugh and Washburn 2004, Touma and Palme 2005, Goyman 2012 ...
Conference Paper
Full-text available
Sound is an intrinsic component of ecosystems and studies have shown that sound plays a significant role in how plants and wildlife interact with their surroundings. The same is true for human-wildlife interactions. Soundscapes vary by day, by season, and across space. Winter in northern latitudes possesses unique soundscape attributes because of a substantial decrease in wildlife vocalizations (biophony), an increase in wind events (geophony), and encroaching noise from winter recreation like snowmobiling (anthrophony). We introduce a fourth soundscape component, silence, as an additional attribute of winter soundscapes. Our objectives were to quantify and visualize the temporal and spatial variation of these four soundscape components in a winter landscape. We sampled 62 locations across the 805,000 ha of Kenai National Wildlife Refuge, Alaska, USA between December 2011 and April 2012. We recorded ambient sounds and quantified the power spectral density in spectrograms at 1 kHz frequency intervals using the Remote Environmental Assessment Laboratory (www.real.msu.edu). We identified sounds from 67,461 recordings and visualized the temporal variation of all soundscape components. We generated predictive spatial models of each soundscape component using machine learning (TreeNet). Silence was the most prevalent record, occurring predominantly at night. Anthrophony, biophony, and geophony were all more prevalent during the day. Anthrophony and biophony had similar temporal patterns over monthly time frames. Geophony was highest in February and January was the quietest month of the season. Spatially, distance to urban interface and rivers were the most common predictors of biophony and anthrophony. Geophony's top predictors were distance to forest and urban interface, and elevation. Distance to rivers, shrubland, and barren land were the top three most important predictors of silence. Our results reveal how winter's biophony, anthrophony, geophony, and silence are arranged over space and time. We also provide evidence of anthrophony's further encroachment into remote wilderness.
... As discussed above, the increased dose predictably enhanced adrenal sensitivity in females but not in males. This dimorphism is supported by another study in reindeer where a sexual difference was observed in fecal GC concentrations using a smaller dose of ACTH (1 IU/kg; [35]). A sexual dimorphism in HPA axis responsiveness is well described in rodents, with males exhibiting lower adrenocortical responses than females [32]. ...
... This explanation is a possibility in this study because GC concentrations prior to challenge were elevated in males relative to females in both ACTH trials. However, in a previous study, male reindeer did not elevate fecal GCs concentrations in response to a lower dose of ACTH (1 IU/kg) and pre-injection concen- trations were not abnormally elevated [35]. These findings suggest that insensitivity to ACTH in male reindeer is consistent with previous studies and not necessarily confounded by high pre-injection concentrations. ...
Article
Full-text available
Climate change and industrial development are contributing to synchronous declines in Rangifer populations across the Arctic. Chronic stress has been implicated as a proximate factor associated with decline in free-ranging populations, but its role in Rangifer is unspecified. Analysis of glucocorticosteroid (GC) concentration in feces, and more recently in hair, is a non-invasive method for monitoring stress in wildlife. Adrenocorticotropic hormone (ACTH) released from the pituitary gland stimulates GC release from the adrenals and can be administered to reflect adrenal activation. In this study, we assessed concentrations of GC metabolites in feces and cortisol in hair of Alaskan caribou (Rangifer tarandus granti) and reindeer (R. t. tarandus) following ACTH treatment. We predicted that ACTH challenge would increase concentrations of fecal GCs, but not hair cortisol because steroid deposited into the hair shaft occurs over an extended period of time (months) and is likely insensitive to acute adrenal stimulation. Adult caribou (n=10; mean age, 6.5 years old) exhibited a peak increase in fecal GCs 8h following a 2 IU/kg dose of ACTH compared to pre-injection concentrations. In contrast, sub-adult reindeer (n=10, 0.8 years old) elicited a diminished response to the same dose. Quadrupling the dose (8 IU/kg) prolonged the fecal GC response in female reindeer, but male reindeer were unresponsive. Hair cortisol was unaffected by a single ACTH challenge. Further investigation is required to ascertain whether subspecific differences in adrenal sensitivity are attributed to age or sex differences, or historical selective pressures from semi-domestication and/or sedentary life cycle in reindeer.
... It also relied on Wilson contradicting the results of previous work on the responses of SMC to helicopters and heli-skiers. In Wilson and Wilmshurst (2019), the authors describe how woodland caribou respond to human disturbances through elevated stress hormones (Freeman, 2008) and abandonment of habitat (Lesmerises et al., 2018), further acknowledging that there are energetic costs to disturbances that they avoid. ...
... The ranges of these ONCP subpopulations during the study years had a spatial extent of industrial impacts that was at least 50% less than that found in Klinse-Za, providing an opportunity to contrast these animals' health metrics with those of caribou exposed to lower levels of anthropogenic disturbance. Because at least some of the health metrics we evaluated in this study have established direct and indirect links with anthropogenic disturbance (e.g., nutrition and range displacement (van Beeck Calkoen et al. 2021)), human activity, and physiological stress (Freeman 2008), we felt that having a comparison group sampled under lower levels of anthropogenic disturbance was important. Given that there are currently no established values for optimal parameter ranges for most caribou health metrics, we compared the health metrics of penned caribou to surrounding subpopulations, including the ONCP subpopulations as well as previously published data from caribou living in non-mountainous boreal landscapes in BC (i.e., "boreal caribou") (Bondo et al. 2019). ...
Article
Full-text available
The health of wildlife plays a crucial role in population demography by connecting habitat and physiology. Southern mountain caribou, a population of woodland caribou (Rangifer tarandus caribou (Gmelin, 1978)) found in the mountains of southwest Canada, are facing significant threats. We evaluated the health of the Klinse-Za subpopulation within the central group of southern mountain caribou, which is part of an Indigenous-led conservation initiative aimed at enhancing caribou population growth through seasonal maternal penning. We collected health metrics from 46 female Klinse-Za caribou between 2014 and 2021. The health metrics included trace minerals, cortisol, biomarkers for inflammation, and pathogen prevalence. We compared these health metrics between penned and non-penned animals, reproductive and non-reproductive females, and nearby subpopulations. We provide correlative evidence linking reproductive success to trace nutrients but find no evidence for relationships with stress, exposure to pathogens, or biomarkers of inflammation. Based on the health metrics considered, Klinse-Za caribou were generally healthy relative to neighboring subpopulations and repeat capture for penning did not appear to create accumulated health issues. Penned caribou had lower fecal cortisol levels and inflammation markers compared to free-ranging animals. This work provides a baseline assessment of southern mountain caribou health and provides guidance on maternal penning activities in support of caribou recovery.
... Recent studies have measured the stress response of caribou in a variety of ways, including assessing levels of stress hormones in hair and feces (Wingfield et al. 1998) and changes in behavior (Freeman 2008;Wasser et al. 2011;Witter et al. 2012;Carlsson et al. 2016). There have been many studies of the behavior of barren-ground caribou, with a particular emphasis on the disturbance responses of caribou to insects during summer (Morschel and Klein 1997;Colman et al. 2001Colman et al. , 2003Hagemoen and Reimers 2002;Witter et al. 2012). ...
Article
Full-text available
Recent and dramatic declines of barren-ground caribou (Rangifer tarandus groenlandicus) in northern Canada have been associated with the effects of climate change, natural population fluctuations, and human disturbance. Past research has revealed that caribou were displaced by industrial activities. These broad-scale distributional changes imply fine-scale behavioral and stress responses. In the central Northwest Territories, Canada, diamond mines and their supporting infrastructure, including winter roads, lie across much of the winter range of a number of populations of caribou. We investigated the fine-scale, mechanistic effect of these disturbance features on the behavior and stress physiology of caribou adjacent to an ice road used to transport industrial materials. We modeled the relationship between the prevalence of observed behavioral category as well as glucocorticoids extracted from fecal pellets and a variety of environmental and disturbance variables, including the distance to the ice road. Proximity to the road was associated with a decrease in time spent foraging and an increase in walking behavior. We did not find a relationship between levels of glucocorticoids in fecal pellets and proximity to the road. These results contribute to efforts to understand and manage the effect of industrial disturbance, including ice roads, on barren-ground caribou.
... I disse områdene kan det også forventes økt stress og lavere kondisjonsparametere enn i mer urørte områder. Slike undersøkelser av stress er ikke undersøkt i Norge for villrein, men for caribou i Nord Amerika er det vist at forstyrrelser øker stressnivå, med konsekvenser for fysiologi og ernaering saerlig for drektige simler og i ressursbegrensede områder (Freeman 2008, Wasser m.fl. 2011, Joly m.fl. ...
Technical Report
Full-text available
[Title in English: Statistical models of cumulative impactrs of human activities on wild reindeer areas: Identifying important grazing areas and scenario analyses for impact assessment and spatial planing] Summary: Abstract Wild reindeer are threatened both nationally and internationally, and several populations have declined sharply or become extinct in recent decades. Several factors are responsible for these trends, but it is well documented that infrastructure development and disturbance by humans can cause habitat degradation and fragmentation and can have a major negative impact on reindeer populations. This led to a large amount of research aimed at understanding and quantifying anthropogenic disturbance on reindeer and supporting sustainable land planning. In the last decade we have developed statistical methods and a software (ConScape) to quantify the functionality of reindeer habitats, to identify movement corridors, and to measure the degree to which these are affected by the piecemeal development and "cumulative impact" of different infrastructure and human activities. The models are based on more than 3 million GPS-positions from the largest wild reindeer management areas, and on ca. 350 environmental layers describing the landscape (topography, vegetation, roads, hydropower, trails, tourist volume, private cabins, tourist cabins, etc.), climate, and, in some areas, local knowledge. Based on these models, several maps have been produced to describe statistically how wild reindeer perceive resources and barriers on a local scale, and which areas and corridors are most functional in the entire landscape. The latter is particularly important, as reindeer perceive the landscape as a continuous network of functional areas that they can access through movement corridors. On the other side, reindeer also perceive the cumulative impact of different infrastructures and human activities and respond by avoiding or decreasing the use of some areas due to disturbance and/or barriers to movements. All maps, a description of methods, and the reference list to scientific and popular publications are available in the Web App: https://www.nina.no/Naturmangfold/Hjortedyr/reindeermapsnorway. In addition, we developed a simulation tool to guide sustainable land planning and impact assessment through scenario analyses. The tool has already been used to predict the expected effect of 80 mitigation measures, suggested by boards of local experts to minimize cumulative impacts on reindeer habitat functionality and movement corridors in several areas in Norway (see Appendix 1 and Web-App). The scenarios that can be tested to include a combination of the removal/closure/relocation of existing infrastructures, changes in their intensity of use, the construction of new infrastructure (e.g. cottage, road, hiking and skiing trial, bridge over water magazines), and climate change. The statistical approach, software, maps, web-app and scenario analyses were developed within the Norwegian Research Council projects led by NINA “RenewableReindeer”, «ProdChange» and “OneImpact”, and in the related project "Scenario analyse Øyulvsbu og formidling". Comparison between statistical and expert-based maps was funded by the project from the Norwegian Environment Agency «OneImpact og kvalitetsnorm for villrein». These projects were or are supported by several partners and funding sources including the Norwegian Environment Agency, the Directorate of Water and Energy, the Hydropower company Sira Kvina, the wild reindeer project in Setesdal, the Wild reindeer centre / wild reindeer council, Siri Bøthun nature management Norwegian University of Life Sciences NMBU, and several international collaboration partners (Universite Catholique Louvain, Julia Computing Inc, Sveriges landbruksuniversitet SLU, University of Guelp, Canada, University of Alberta, Canada, The Nature Conservancy, Universidad Politecnica de Madrid, University of Glasgow). Parallel to this analytical work, management measures have been undertaken in Norway to counter the degradation of wild reindeer areas. In 2020 the government adopted the “Quality Standards for wild reindeer in Norway” (Lovdata 2020), to meet both international obligations and national objectives for the conservation of viable populations within ecologically functioning habitats. Every fourth year, each of the 23 Norwegian sub-populations is classified into good (green), medium (yellow) or poor (red) quality, based on 3 sub-standards: 1) population conditions; 2) lichens; 3) human impact on habitats. Sub-standard 3 is therefore an important tool to ensure sustainable management of wild reindeer habitats. Sub-standard 3 aims to identify critical declines in habitat quality and connectivity in each wild reindeer area, so that the cause of the decline can be addressed, and the status of the area restored to acceptable levels (yellow) as soon as possible. In the longer term, the aim is that all national wild reindeer areas should be of good quality (green). The procedure thus needs to robustly identify areas where habitat loss and fragmentation increased above a critical threshold, and to provide information on the relative contribution of the responsible infrastructure and human activities. In the current Quality Standards, sub-standard 3 is implemented based on expert-based assessments of changes in available wild reindeer observations, that are assumed to reflect changes in human impact. The process involves the delineation of polygons representing reindeer seasonal ranges, corridors, and a set of focal areas where socio-ecological challenges related to human activities have been identified. The reduction in use of focal areas is assessed by the experts in the last decade compared to the previous four decades. The classification is then conducted by assessing the proportion of habitat lost within all focal areas, as compared to the habitat available within the seasonal range. In this report we provide an overview of the statistical approach, and we compare the performance of what we for simplicity call the “statistical maps” to the “expert-based polygons” developed in sub-standard 3, using both visual and quantitative approaches. We discuss the lessons learned by comparing the two approaches, and how these can help achieving the sustainability goals for the management of wild reindeer areas. This was the main focus of the project "OneImpact and quality standard for wild reindeer" funded by the Norwegian Environment Agency. As a proof of concepts, we also perform a preliminary statistical classification of the state of the wild reindeer areas following Delnorm 3 (“Sub-standard 3”). Last, we show how statistical approaches and simulations can help identifying the most effective among 76 measures suggested to mitigate cumulative impacts from anthropogenic activities in Setesdal, Nordfjella and Snøhetta (Appendix 1). Research on cumulative impacts of infrastructure and human activities Reindeer areas are not threatened by a single anthropogenic factor such as a hydropower reservoir, a road, a tourist resort or a cabin village. Reindeer is a migratory species that require wide and well-connected ranges, whose functionality is threatened by the combination of all these factors together. The simultaneous presence of several sources of disturbance spread across the landscape creates a series of obstacles to their free movements that can force reindeer to use only parts of their originally large ranges, and abandon pastures used in the past. Infrastructures and human activities that, individually, may have only little effect on reindeer, can together cause a large cumulative impact. This depends on how strong the impact of each of these activities are, how often or how many of these infrastructures occur on the landscape, how far their effect can be perceived by reindeer, and on their position with respect to important reindeer grazing areas and corridors. Statistical models are the most efficient way to understand complex interactions involving many variables at high resolution, over large areas. In the past decade we built data infrastructure, methods and software and we used them to quantify the effect of more than 200 data layers describing vegetation, topography, climate, infrastructures (e.g. roads, hydropower, cabins, trails) and human activities (e.g. tourist volume) on more than 400 GPS-monitored reindeer. The approach builds upon two statistical analyses: one quantifies habitat quality or habitat loss (probability to use an area, taking into account vegetation, topography, climate, infrastructures and human activities); the other quantifies fragmentation, or barriers to movements. These analyses offer robust estimates of the magnitude of the effect of each infrastructure and human activity, and of the distance at which such effects can be perceived by reindeer, for each 100 m pixel in Norway. We also synthesised in different ways human impact in each 100 m pixel in Norway (current “human footprint”; “natural potential” etc), and we ranked the main factors causing habitat loss in each reindeer area. Last, we took a “bird-eye-view” of the landscape and developed new models showing all movement corridors and the most functional areas. Models can be used also to predict beyond the data (i.e. extrapolation) in areas where no GPS data are available, or under past or future scenario of infrastructure development (e.g. build roads, cabins, hydropower), mitigation measures (e.g. move a cabin, reduce or re-direct tourism), or climate change (e.g. changes in snow). Comparing statistical and expert-based approaches Statistical maps and expert-based polygons of seasonal ranges generally agree on the location of suitable reindeer pastures, though meaningful comparisons are prevented by the size of the polygons, that at times extend to the entire management area. However, statistical models perform significantly better than the polygons in discriminating between areas used or not used by reindeer, according to both GPS data and to reindeer observations. This is partly because the polygons are delineated by hand, and necessarily include “obviously” unsuitable areas such as hydropower reservoir, roads, cabins etc. We show that this risks to underestimate habitat loss in the classification of Sub-standard 3. Statistical models use data on infrastructure and human activities to quantify anthropogenic impacts directly and in real-time (as soon as new developments are updated in the data). Models can also be used to prevent impacts by testing the effect of land development plans, before infrastructures are built. The expert-based approach on the contrary is set to detect habitat loss or fragmentation only following a decline in reindeer observations in an area. This risks to delay or prevent the detection of human impacts, especially in fragmented reindeer areas where human impact is widespread, and reindeer have no access to refuge areas. In such areas, reindeer have to stay, and might suffer higher stress levels, with consequences for individual conditions. Statistical models provide high-resolution estimates of cumulative impacts, for every 100 m of the landscape. This implies that all sources of disturbance are quantified, including both highintensity ones (e.g. high-traffic roads, railway), or sources of disturbance that may be of lowerintensity, but diffuse and widespread in the landscape (e.g. network of hiking trails, private cabins). This allows avoiding the risk of failing to address cumulative impacts of low-intensity, diffuse sources of disturbance. A simulation showed that the same amount of disturbance spread across the entire wild reindeer area or concentrated within a specific location can lead to a vastly different classification following Sub-standard 3, as the procedure currently does not account for diffuse, low-intensity sources of disturbance. Another advantage of using high-resolution model estimates is that there is no need to subjectively outline several polygons (i.e. one for each focal areas, seasonal range, corridor and influence area - for each season). Using simulations, we showed that the process of outlining the polygons can entirely determine the final classification of each wild reindeer area following Sub-standard 3. Hence, the final classification can be determined at the first stage of Sub-standard 3, when the polygons are drawn, irrespective from the following expert-based assessment of a possible decline in reindeer area use therein. This is because the classification depends on the proportion of the seasonal ranges covered by the focal area, and thus their respective size can fully determine the assessment. Indeed, most of the four areas we assessed could only be classified as “green” or “yellow” – not “red” – as the sum of their focal areas is too small to allow the area to be classified as red. Hence, deciding whether to include or exclude a hydropower reservoir or a glacier in the polygon may determine the final classification of that area. Statistical analysis coupled with network models can simultaneously assess habitat loss and fragmentation (corridors and functional habitat) caused by multiple sources of disturbance and taking into account also a variety of natural factors. In other words, statistical models assess Substandard 3A (on habitat loss) and 3B (on fragmentation) within the same framework, in a standardized way, within and across reindeer areas. Can statistical models support decision-making processes? The reliability of both data and models increased rapidly in recent decades, and the report shows that results are robust with respect to available reindeer data. Feedbacks from local experts are generally positive, and the results correspond fairly well with local knowledge, even in areas where GPS data are not available. “All models are wrong, but some are useful”. The question is not “are the model representing the truth?” (no, they never will), but “are they useful to achieve the goals of Sub-standard 3?”, or “can they help identifying the most efficient mitigation measures?”. In their current form, the models seemed useful to understand why reindeer use a specific area, or why they no longer use it, which human activities cause the strongest impact in different seasons, or what is the expected impact in areas with little data or local knowledge. Statistical models can also be useful to identify areas, human activities and infrastructures that would need to be prioritized for conservation, mitigation or restoration both locally and across reindeer areas. These models were not made for Sub-standard 3, but are a flexible, “living product” that can be adjusted, updated and improved, for instance by adding data on bucks, changing the seasons or, most importantly, by integrating local knowledge. The models are operational, flexible, reproducible, and could support Sub-standard 3 in a variety of ways. The report discusses in detail strengths and weaknesses of both statistical and expert-based approaches and concludes that a robust interaction between statistical models and local knowledge would be ideal for maximizing the probability of reaching the sustainability goals of Sub-standard 3, while minimizing the risks highlighted in this report.
... from Matilde Tomaselli, University of Calgary). Fecal glucocorticoid metabolite levels in elk (Creel et al., 2002) and caribou (Freeman, 2008) are also positively associated with snowmobile activity. Lower food abundance and quality (Adamczewski et al., 1992), along with increased agonistic behaviors (displacements) related to food accessibility often restricted to feeding craters (Gray, 1987), may lead to higher nutritional and social stress in the fall and winter. ...
Thesis
Full-text available
Glucocorticoids play a key role in energy regulation and are mediators of the physiological stress response in mammals. Their concentrations are commonly measured in wildlife to understand the effects of environmental changes and anthropogenic disturbances, but their use is associated with multiple challenges and there is a need for species-specific validation. Muskoxen (Ovibos moschatus) are an essential part of the Arctic ecosystem, where they have a strong economic, nutritional, and sociocultural value for Indigenous communities. Recent population declines and mortality events suggest that muskoxen may be threatened by the multiple environmental changes and associated stressors to which they are increasingly exposed. Overall, I sought to establish fecal glucocorticoid metabolites (FGM) and qiviut (woolly undercoat) cortisol as biomarkers of physiological stress in muskoxen, and to apply these tools together with Indigenous knowledge (IK) to explore potential causes and patterns of physiological stress in wild muskoxen. Through two repeated pharmacological challenges in captive muskoxen, I showed that qiviut cortisol and FGM levels accurately reflect long-term (over the period of the hair’s growth) and short-term changes in circulating cortisol, respectively. I also demonstrated that changes in circulating cortisol are not reflected in qiviut in the absence of growth and highlighted variations across body regions, significant differences in qiviut segments over time, and differences between shed and unshed qiviut. Additionally, I documented IK which provided novel insights on the potential stressors of muskoxen and their specific importance. Finally, I identified important factors influencing qiviut cortisol (sex, geographical location, season, and year), and found associations between qiviut cortisol and marrow fat and lungworm intensity. Findings were interpreted in part collaboratively with IK holders. This work has advanced our understanding of glucocorticoid deposition and stability in hair, and of the limitations and challenges associated with hair glucocorticoid interpretation. It has highlighted the multiple benefits of incorporating IK in wildlife endocrinology studies and provided a framework for doing so. Finally, identifying factors associated with qiviut cortisol is a key step to simultaneously investigating the causes and consequences, both at the individual and population levels, of physiological stress in muskoxen.
... The threat to caribou from snowmobiles and other intensive winter motorized recreation such as heliskiing in late winter has been well studied (Duchesne et al. 2000, Fancy and White 1985, Kinley 2008, Seip et al. 2007. In one study, mountain caribou exhibited increased corticosteroid levels (a well-known metabolic indicator of physiological stress) even up to 10 km from snowmobiles (Freeman 2008). ...
Article
Full-text available
In early 2014 Environment Canada released a recovery strategy for the “Southern Mountain” ecotype of woodland caribou. This strategy falls short of a recovery plan, notes the absence of detailed maps of critical habitat (which are required by the Species at Risk Act, SARA), and states that such maps “will be prepared for each local population unit as the information becomes available and will be included in a revised recovery strategy, or in one or more action plans” (Environment Canada 2014:page 40). This report was prepared to inform federal species-at-risk officials about available data, especially mapping, of critical habitat for southern mountain caribou.
... For example, roe deer from the field habitat can be also more exposed to stress, due to the lower possibility to hide or escape from danger (own observations). Numerous studies of wild living animals have shown that the noises caused by motor vehicles and some human activities can result in elevated levels of the stress hormone cortisol (Creel et al. 2002, Barja et al. 2007, Freeman 2008, Zwijacz-Kozica et al. 2013) and higher levels of such metabolites as glycogen and lipid products (Majewska et al. 1982). Cortisol is a glucocorticoid, an example of a class of compounds which allow the organism to adapt to stress and maintain homeostasis. ...
Article
Dental enamel is the hardest tissue of the mammalian body, consisting of 96–98% inorganic compound. As the dentition is functionally adapted to diet and feeding behaviour, relative differences in enamel thickness can reflect dietary adaptations. We hypothesize that differences in enamel thickness are related to adaptation for diet associated with habitat quality dwelling of European roe deer Capreolus capreolus. To test this hypothesis, 49 first permanent left lower molars were extracted from the mandible of roe deer (from Lithuania – 28 and Poland – 21 molars) inhabiting two type of habitats: field and forest. The linear thickness of total enamel (mean value of enamel thickness measured at three different points) was found to differ between the roe deer from the field and forest habitats, irrespective of age, with the animals of field ecotype tend to have thinner enamel (F(1,26) = 6.845, P = 0.025). This suggests that there is an adaptation in enamel thickness to various types of diet in the field and forest habitat. On the other hand, roe deer from the field habitat can be also more exposed to stress, due to the lower possibility to hide or are more vulnerable to potential threats. More frequent exposure to stress can significantly disrupt ameloblasts secretion and thus affect the thickness of the enamel.
... Lesmerises et al. (2018) found a strong enough individual response to skiers that habitats were abandoned by caribou. A similar individual disturbance response where snowmobiles and helicopter skiing were present resulted in elevated stress hormones in caribou as compared to areas where these activities were absent (Freeman, 2008). Threshold disturbances by people approaching on foot (skiing or hiking) suggested that reindeer (Rangifer tarandus tarandus) pay an energy price to avoid disturbance (Reimers et al., 2003). ...
Article
Full-text available
Helicopter-and snowcat-supported backcountry skiing is a unique industry that is widespread throughout southern mountain caribou habitat in British Columbia (BC). We analyzed records of helicopter and skier encounters with caribou collected by tourism operators under an agreement between the BC government and HeliCat Canada. Average reported encounter rates were low for helicopters (0.6%) and skiing groups (0.1%); however, encounters were likely underreported due to factors that affect caribou sightability. Helicopters encountered caribou more frequently than skiing groups, caribou were detected from helicopters at greater distances than by skiers. We used Bayesian network models to assess the independent contribution of different factors to the behavioural response of caribou to encounters. Encounter distance was the most important factor in both helicopter and skiing models. Larger groups of caribou responded strongly to skiers but not to helicopters, although the independent effect of this factor was small in both models. Larger helicopters elicited stronger reactions from caribou than smaller machines and were responsible for 25% of the modelled variation in caribou response. Encounters with helicopters at distances of 100-500m had a 78% probability of eliciting a concerned-to-very-alarmed response from caribou, while skiers at a similar distance had a 60% probability of eliciting the same response. The probability of concerned-to-very-alarmed responses dropped to <20% at encounter distances of >1000 m. These results indicate that initial encounter distance is the key variable to manage risk to caribou of helicopter and skiing encounters. Ongoing feedback on the effectiveness of management practices is critical to ensure the continued viability of industries operating in caribou habitat.
... Muskoxen react to snowmobile activity (McLaren and Green, 1985) and in the area around Cambridge Bay, behavioural changes, with animals having a much longer flight distance, have been observed with increased snowmobile activity (personal communication from Matilde Tomaselli, University of Calgary). Faecal glucocorticoid metabolite levels in elk (Creel et al., 2002) and caribou (Freeman, 2008) are also positively associated with snowmobile activity. Lower food abundance and quality (Adamczewski et al., 1992), along with increased agonistic behaviours (displacements) related to food accessibility often restricted to feeding craters (Gray, 1987), may lead to higher nutritional and social stress in the fall and winter. ...
Article
Full-text available
Muskoxen (Ovibos moschatus) are increasingly subject to multiple new stressors associated with unprecedented climate change and increased anthropogenic activities across much of their range. Hair may provide a measurement of stress hormones (glucocorticoids) over periods of weeks to months. We developed a reliable method to quantify cortisol in the qiviut (wooly undercoat) of muskoxen using liquid chromatography coupled to tandem mass spectrometry. We then applied this technique to determine the natural variability in qiviut cortisol levels among 150 wild muskoxen, and to assess differences between sexes, seasons and years of collection. Qiviut samples were collected from the rump of adult muskoxen by subsistence and sport hunters in seven different locations in Nunavut and the Northwest Territories between 2013 and 2016. Results showed a high inter-individual variability in qiviut cortisol concentrations, with levels ranging from 3.5 to 48.9 pg/mg (median 11.7 pg/mg). Qiviut cortisol levels were significantly higher in males than females, and varied seasonally (summer levels were significantly lower than in fall and winter), and by year (levels significantly increased from 2013 to 2015). These differences may reflect distinct environmental conditions and the diverse stressors experienced, as well as physiological and/or behavioural characteristics. Quantification of qiviut cortisol may serve as a valuable tool for monitoring health and informing conservation and management efforts.
... Backcountry recreational activity, especially in winter, has the potential to disturb and displace mountain caribou from preferred habitats and to increase their risk of predation and exposure to avalanches (Seip and Cichowski 1996, Simpson and Terry 2000, Cichowski et al. 2004, Seip et al. 2007, COSEWIC 2014, USFWS 2014. Disturbance and displacement can increase stress and energy expenditure among caribou (Reimers et al. 2003, Freeman 2008) and potentially reduce feeding and body condition, whereas hardpacked winter trails can allow wolves and other predators greater access to caribou (COSEWIC 2014). In recent decades, winter activities (e.g., snowmobiling, skiing, heli-skiing, snowshoeing) have expanded to many of the remote areas required by caribou (Seip et al. 2007). ...
Technical Report
Full-text available
Woodland caribou (Rangifer tarandus caribou), a subspecies of caribou, occur across the boreal regions of North America and are comprised of eight recognized populations. The southern mountain caribou population consists of 17 subpopulations, or herds, with the South Selkirk subpopulation being one of these. This subpopulation occurs in the southern Selkirk Mountains of southeastern British Columbia, northeastern Washington, and northern Idaho, and is the only caribou herd that ranges into the contiguous U.S. The subpopulation possibly numbered in the hundreds in the late 1800s, but decreased to an estimated 25-100 caribou between 1925 and the mid-1980s. Numbers ranged between 33 and 51 animals from 1991 to 2009 despite being supplemented with 103 caribou in two separate multi-year translocations in the late 1980s and 1990s. Most recently, the subpopulation declined rapidly from 46 to 12 caribou between 2009 and 2016. The percent of calves in the subpopulation during late winter surveys averaged 9.9% per year from 2004 to 2016, which is below the estimated 12-15% needed to maintain a stable population with high adult survival. Predation is considered the most immediate threat to the South Selkirk subpopulation. In addition, past conversion of old-growth forests to earlier successional stages has brought higher densities of deer, moose, and elk and their predators (i.e., wolves, cougars, and bears) into closer proximity to herd members, resulting in greater predation risk to caribou. Other threats are highway collisions, human disturbance associated mostly with winter backcountry recreation, small population size coupled with isolation, and climate change. For these reasons, it is recommended that woodland caribou remain a state endangered species in Washington.
... Until these issues are resolved, we recommend that hair be collected from the same site, using standardized methods, if longitudinal and between-individual comparisons are to be made. To our knowledge, this is the first longitudinal study on Rangifer to present data on both faecal cortisol and corticosterone metabolites from the same samples (Freeman, 2008; Ashley et al., 2011; Wasser et al., 2011; Macbeth, 2013; Yin et al., 2015), and it has revealed several noteworthy patterns that would have been missed in a cross-sectional study. First, analysis revealed what appears to be a 21 day cyclic pattern in faecal cortisol levels that was not present in faecal corticosterone levels. ...
Article
Full-text available
Stress hormones (glucocorticoids), incorporated into hair/fur and faeces, have been proposed as biomarkers of overall health in wildlife. Although such biomarkers may be helpful for wildlife conservation and management, their use has rarely been validated. There is a paucity of studies examining the variation of stress hormones in mammals and how they relate to other health measures, such as parasitism. Parasites are ubiquitous in wildlife and can influence the fitness of individual animals and populations. Through a longitudinal experiment using captive reindeer (Rangifer tarandus tarandus), we tested whether animals infected with Ostertagia gruehneri, a gastrointestinal nematode with negative impacts on fitness of the host, had higher stress levels compared with those that had been treated to remove infection. Faecal samples were collected weekly for 12 weeks (June–September) and hair was collected at the start and end of the study; glucocorticoids were quantified using enzyme immunoassays. Contrary to what was expected, infected reindeer had similar levels of cortisol in hair and slightly lower glucocorticoid metabolites in faeces compared with uninfected reindeer. Faecal corticosterone levels were higher than faecal cortisol levels, and only corticosterone increased significantly after a handling event. These results suggest that reindeer may use a tolerance strategy to cope with gastrointestinal nematodes and raise the question as to whether moderate infection intensities with nematodes are beneficial to the host. By removing nematodes we may have altered the gut microbiota, leading to the observed elevated faecal glucocorticoid metabolite levels in the treated reindeer. These findings demonstrate the importance of considering both cortisol and corticosterone in physiological studies, as there is mounting evidence that they may have different functionalities.
Article
Full-text available
Ecological and conservation research has focused on documenting wildlife's behavioural and demographic responses to anthropogenic disturbance. Yet, a significant gap remains in our understanding of the physiological mechanisms that underlie behavioural or demographic adaptations to human activities. Glucocorticoid concentrations (GCCs) are widely considered as relevant endpoints to assess the physiological response of wildlife to natural or anthropogenic stressors, yet recent studies show mixed outcomes concerning GCC shifts in disturbed populations. Consequently, uncertainties remain around the role of GCCs as reliable indicators of the effect of anthropogenic disturbance on organisms' physiology, likely due to the masking effect of several confounding factors. To unveil generalizable patterns of physiological effects of human disturbance on wildlife, we collected GCCs measured in conspecific populations living in undisturbed and disturbed areas from 121 studies and 114 species of terrestrial vertebrates, and assessed the roles of disturbance type and species traits in shaping GCC shifts using a phylogenetic meta‐analysis of log response ratios (lnRRΔ). We found a 36% increase in GCCs due to human disturbance in mammals (lnRRΔ = 0.304), but no evidence of a generalizable effect for birds, non‐avian reptiles and amphibians. Mammals showed similar responses to all forms of anthropogenic disturbance, while birds showed 24.6% increase in GCCs for habitat conversion only (lnRRΔ = 0.221). Furthermore, change in GCCs tended to be higher in large‐sized than in small‐sized mammals, but comparatively lower in herbivorous mammals. Finally, we found non‐migratory birds to elicit stronger responses compared with migratory birds (lnRRΔ = 0.234). Our study provides novel evidence supporting the influential roles of diet and body mass in mammals, and migratory behaviours in birds, in shaping glucocorticoid responses to human disturbance, establishing a link between magnitude of physiological responses and pace‐of‐life‐related traits. Furthermore, we found GCC trends to be coherent with known demographic and behavioural responses, suggesting that GCC shifts might be costly to sustain, thus serving as potential early‐warning signals of population decline. Finally, we found consistency in the magnitude of GCC shifts in mammals facing all types of human disturbance, highlighting that even seemingly innocuous interactions with humans may still induce substantial physiological changes. Read the free Plain Language Summary for this article on the Journal blog.
Preprint
HeliCat Canada members operating in mountain caribou range have been collecting and reporting wildlife sightings data since 2010, resulting in more than 3,750 spatially referenced sightings of animals and tracks by the end of the 2022 operating season. Mountain caribou, mountain goats, and wolverine have been the most commonly recorded species, with caribou observations generally declining over time, mountain goat sightings showing no trend, and observations of wolverines and their tracks increasing. No reaction and Unconcerned are the most common responses of caribou and mountain goats to encounters with helicopters or skiers. Behavioural responses are generally stronger at shorter encounter distances, but responses remain variable, with some animals still showing No reaction or an Unconcerned response to close encounters. Recorded responses to skiers tend to be stronger than to helicopters, particularly among caribou. The number and proportion of Alarmed or Very alarmed responses to encounters was higher in early years of monitoring but has been stable since about 2014. Evidence suggesting that HeliCat activities are causing population declines of mountain caribou or mountain goats is lacking. Opportunities to further reduce risk include: improved habitat mapping, expanded monitoring and reporting, telemetry data sharing, and passive detection of wildlife. However, the benefits of all of these approaches are also currently associated with a variety of limitations.
Technical Report
Full-text available
Effects of winter recreation on the natural environment is summarized here for use by land managers seeking to mitigate negative impacts. Studies show that winter recreation of all kinds can impact wildlife, which are particularly vulnerable to disturbance from unpredictable human activities such as off-trail snowmobiling or backcountry skiing. In addition, research has quantified ways in which over-snow vehicle (OSV) use can damage vegetation, compress soils, affect air and water quality, and disrupt natural soundscapes.
Article
Full-text available
In the face of rapid environmental change, anticipating shifts in microparasite and macroparasite dynamics, including emergence events, is an enormous challenge. We argue that immunological studies in natural populations are pivotal to meeting this challenge: Many components of environmental change – shifts in biotic assemblages, altered climate patterns, and reduced environmental predictability – may affect host immunity. We suggest that wild ungulates can serve as model systems aiding the discovery of immunological mechanisms that link environmental change with parasite transmission dynamics.Our review of eco-immunological studies in wild ungulates reveals progress in understanding how co-infections affect immunity and parasite transmission; and how environmental and genetic factors interact to shape immunity. Changes in bioavailability of micronutrients have been linked to immunity and health in wild ungulates. Although physiological stress in response to environmental change has been assessed, downstream effects on immunity have not been studied. Moreover, the taxonomic range of ungulates studied is limited to bovids (bighorn sheep, Soay sheep, chamois, musk oxen, bison, African buffalo) and a few cervids (red deer, black-tailed deer). We discuss areas where future studies in ungulates could lead to significant contributions in understanding patterns of immunity and infection in natural populations and across species.This article is protected by copyright. All rights reserved.
Article
Full-text available
Caribou in Gros Morne National Park reacted to provocation by snowmachine with significant differences in their response between years. Upon exposure to snowmachines, caribou were displaced 60 to 237 m from their initial locations. Groups with calves allowed the snowmachines to approach more closely before responding (5 to 600 m) than adult-only groups (30 to 1300 m), and their overall flight distances were less. Time spent in locomotion and overall reaction time were greater for animals engaged in the most sedentary activities (eg. standing; mean = 239 s and mean = 262 s, and lying; mean = 166 s and mean = 273 s) than for animals already engaged in more dynamic activities such as walking (mean = 118 s and mean = 133 s), running (mean = 74 s and mean = 63 s) and feeding (mean = 118 s and mean = 133 s). Annual differences in the response of adult-only groups were not due to differences in the sex ratio of these groups, but may be related to annual variation in winter weather conditions.
Article
Full-text available
Caribou (Rangifer tarandus) in British Columbia are classified into mountain, northern and boreal ecotypes based on behavioural and ecological characteristics. We recognized 12 mountain caribou herds, 27 northern caribou herds, and an area occupied by low density boreal caribou dispersed in the boreal forests of the northeast portion of the province. Abundance estimates were usually based on attempts at total counts made from the air. Trends were based on repeated population estimates or the difference between recruitment and mortality rates for each herd. In 1996 the-re were approximately 18 000 caribou in British Columbia; 2300 mountain and 15 600 northern and boreal. These estimates suggest a slight increase in the numbers of both ecotypes over the last 18 years. Fifteen percent of the herds were reportedly increasing, 10% were decreasing, 31% were stable, but for 44% of the herds the trend was unknown. Historically caribou were found throughout 8 of the 14 biogeoclimatic zones in B.C. Caribou are now rarely found in the Sub-Boreal Spruce zone, likely due ro increased predation from wolves that increased in response to increasing moo-se numbers. Ranges of several herds in the Engelmann Spruce — Subalpine Fir and Alpine Tundra zones of south-eas-tern British Columbia are also reduced relative to historic conditions, probably because of habitat loss, habitat fragmen-tation, predation and hunting. Forest harvesting represents the greatest threat to caribou habitat and current research focuses on the mitigation of forest harvesting impacts.
Article
Full-text available
To better understand the effect of winter tourism and public recreation on wild mountain reindeer (Rangifer tarandus tarandus), we compared reindeer response distances after direct provocations by skiers and snowmobiles during 3 winters in Setesdal-Ryfylke, Southern Norway. Reindeer being provoked by a snow-mobile discovered the observer at longer distances than reindeer being provoked by a skier (370 [skier] vs. 534 [snowmobile] m; P = 0.002), while total flight (756 vs. 570 m; P = 0.037) and total distance moved (970 vs. 660 m; P = 0.008) by reindeer were shorter for snowmobile than skier provocation. The fright (328 [skier] vs. 328 [snowmobile] m), flight (281 vs. 264 m), and escape (543 vs. 486 m) distances due to skier or snowmobile provocation were not different (P > 0.05). For pooled data, fright distances of reindeer were affected by 2 other independent variables. Fright distance was longer when the animals were provoked from below rather than from above (P = 0.046), while their escape distances were longer when the animals were lying rather than when grazing prior to being provoked (P< 0.05). Based on maximum and minimum distance moved for all provocations pooled, daily estimated energy expenditure of reindeer increased between 31 and 590 kJ, representing 0.2 and 2.9% of their estimated total daily energy expenditure. Overall, provocations by skiers or snowmobiles revealed similar behavioral responses. An estimated maximum rate of 3 daily encounters between reindeer and skiers or snowmobiles during winter vacation and Easter would result in moderate energy costs that should be easily compensated for and thus have no demographic consequences. Increasing snowmobile use will, however, significantly expand the area where humans are in contact with reindeer during winter and spring, a period of negative energy balance for reindeer.
Article
Full-text available
Fecal glucocorticoid assays provide a potentially useful, noninvasive means to study physiological responses of wildlife to various stressors. Consequently we quantified fecal glucocorticoid concentrations among free-ranging elk (Cervus elaphus) by subherd, sex, and season and determined their relationship to various human activities and environmental conditions. Using a validated technique, we assayed 558 fecal samples collected from 30 radiocollared elk in the Black Hills, South Dakota, from June 1995 to September 1997. Fecal glucocorticoid measures were least in winter (x̄=17.41 ng/g, SE=2.97 for bull subherds and 18.9 ng/g, SE=2.85 for cow subherds) and increased to peak concentrations in summer (x̄=33.6 ng/g, SE=3.42 for bull subherds and 34.21 ng/g, SE=3.71 for cow subherds). Vehicle use along primary roads, primary road density, and mean temperature each independently explained a significant portion of the variation in fecal glucocorticoid concentrations ( r2=0.61, F2, 557=286.13r^{2}=0.61,\ F_{2,\ 557}=286.13 , P<0.001). Annual glucocorticoid secretion also may be related to normal seasonal metabolic rhythms. Though more research is needed on fecal glucocorticoid concentrations of undisturbed elk, we hypothesize that human activities, high temperatures, or normal seasonal metabolic rhythms may have elevated summer glucocorticoid concentrations. Our findings suggested that fecal glucocorticoid assays, if coupled with population performance measures, could assess physiological effects of natural and human-induced disturbances on free-ranging elk. Also, our study illustrated the need for caution when interpreting fecal glucocorticoid measurements, because several confounding factors may influence interpretation.
Article
Full-text available
Measuring stress hormones (i.e., glucocorticoids) in large free-ranging vertebrates such as white-tailed deer (Odocoileus virginianus) is difficult, and invasive procedures (e.g., animal capture and blood collection) have been the only tools available. Non-invasive techniques, including fecal and saliva glucocorticoid measurements, are currently being developed that offer advantages over traditional techniques. Using adrenocorticotropic hormone (ACTH) challenges, we determined that fecal glucocorticoid metabolite and salivary cortisol assays measured adrenal activity (i.e., stress hormones) in white-tailed deer. For each of four ACTH trials, we housed captive deer singly for 72 hr in an outdoor pen to which they had been exposed previously. After 24 hours, we injected a total of 50 I.U. of ACTH (Cortrosyn™) intramuscularly (2 25-I.U. injections in experiments 1 and 2; 1 50-I.U. injection in experiments 3 and 4). All available fecal samples were collected every hour, homogenized, and frozen. During the last 2 experiments, we collected saliva samples at 10-min intervals for 2 hr prior to and 6 hr after the ACTH injection. Fecal glucocorticoid metabolites were extracted from fecal samples and assayed using a commercially available I125{\rm I}^{125} corticosterone radioimmunoassay. We used an enzyme immunoassay to measure salivary cortisol. A peak in fecal glucocorticoids occurred at 20-24 hr (experiments 1 and 2) and 10-13 hr (experiments 3 and 4) post-ACTH injection, physiologically validating the assay's ability to detect biologically important changes in adrenocortical activity. Salivary cortisol levels increased 30-60 min after ACTH injection, confirming the assay's ability to detect changes in cortisol concentrations. These findings demonstrated that fecal glucocorticoid and salivary cortisol assays provide an index of physiological stress in white-tailed deer. These techniques may prove useful in addressing conservation issues surrounding white-tailed deer, provided additional sampling questions are addressed.
Article
Full-text available
We synthesize findings from cooperative research on effects of petroleum development on caribou (Rangifer tarandus granti) of the Central Arctic Herd (CAH). The CAH increased from about 6000 animals in 1978 to 23 000 in 1992, declined to 18 000 by 1995, and again increased to 27 000 by 2000. Net calf production was consistent with changes in herd size. In the Kuparuk Development Area (KDA), west of Prudhoe Bay, abundance of calving caribou was less than expected within 4 km of roads and declined exponentially with road density. With increasing infrastructure, high-density calving shifted from the KDA to inland areas with lower forage biomass. During July and early August, caribou were relatively unsuccessful in crossing road/pipeline corridors in the KDA, particularly when in large, insect-harassed aggregations; and both abundance and movements of females were lower in the oil field complex at Prudhoe Bay than in other areas along the Arctic coast. Female caribou exposed to petroleum development west of the Sagavanirktok River may have consumed less forage during the calving period and experienced lower energy balance during the midsummer insect season than those under disturbance-free conditions east of the river. The probable consequences were poorer body condition at breeding and lower parturition rates for western females than for eastern females (e.g., 1988 - 94: 64% vs. 83% parturient, respectively; p = 0.003), which depressed the productivity of the herd. Assessments of cumulative effects of petroleum development on caribou must incorporate the complex interactions with a variable natural environment.
Article
Full-text available
Free-living snow buntings (Plectrophenax nivalis) and Lapland longspurs (Calcarius lapponicus) showed marked increases in circulating corticosterone up to 1 hr post-capture, indicating that their hypothalamo-pituitary-adrenal axes responded to the acute stress of handling and restraint in a manner similar to that of other vertebrates, although the magnitude of these responses was generally less. In the snow bunting there was no difference in the adrenocortical response to stress between June (breeding) and September (just prior to autumn migration). However, individual variation in the maximum corticosterone level attained during the stressful period was significantly correlated with fat score in both species. Those with greater fat depots have reduced responsiveness to stress.
Article
Full-text available
We investigated terrain preferences of caribou (Rangifer tarandus granti) in an oilfield region near Prudhoe Bay, Alaska. Under disturbance-free conditions, the distribution of calving caribou determined by aerial transect surveys was correlated with indices of terrain ruggedness based on map contours. Caribou preferred quadrats dominated by fine- textured rugged terrain, particularly when present in large clusters, and avoided quadrats with flatter terrain. Displacement of maternal females from a zone within 4 km of roads and production-related facilities reduced use of rugged terrain types in that zone by 52%; the remaining preferred terrain was scattered and less accessible. This reduction was accompanied by a 43% increase in caribou use of rugged terrain 4-10 km from surface development. Given that terrain ruggedness is positively correlated with forage quality and biomass availability, combined underuse and overuse of these important habitats may compromise summer nutrition of lactating female caribou, thereby depressing body condition and, hence, subsequent reproductive success.
Article
Full-text available
Military training exercises have increased in Alaska in recent years, and the possible effects of low-altitude overflights on wildlife such as barren-ground caribou (Rangifer tarandus) have caused concern among northern residents and resource agencies. We evaluated the effects of overflights by low-altitude, subsonic jet aircraft by U.S. Air Force (USAF) A-10, F-15, and F-16 jets on daily activity and movements of free-ranging female caribou. This study was conducted on caribou of the Delta Caribou Herd in interior Alaska during each of 3 seasons in 1991: late winter, postcalving, and insect harassment. Noise levels experienced by caribou were measured with Animal Noise Monitors (ANMs) attached to radiocollars. Caribou subjected to overflights in late winter interrupted resting bouts and consequently engaged in a greater number of resting bouts than caribou not subjected to overflights (P = 0.05). Caribou subjected to overflights during postcalving were more active (P = 0.03) and moved farther (P = 0.01) than did caribou not subjected to overflights. Caribou subjected to overflights during the insect season responded by becoming more active (P = 0.01). Responses of caribou to aircraft were mild in late winter, intermediate in the insect season, and strongest during postcalving. We conclude that females with young exhibit the most sensitive response to aircraft disturbance. Accordingly, military training exercises should be curtailed in areas where caribou are concentrated during calving and postcalving.
Article
Full-text available
Knowledge of the effect of mining developments on caribou Rangifer tarandus is fragmentary. We examined the impact of the Hope Brook gold mine, southwestern Newfoundland, on the La Poile woodland caribou herd on a section of their year-round range. We examined the impact of the mine on caribou distribution during three phases of mine activity (pre-disturbance, construction and open-pit mining and underground mine and mill operation) in five seasons (winter, late winter, pre-calving, calving and autumn). Aerial surveys were conducted on a monthly basis from September 1985 to July 1991. Following initiation of the mine construction, caribou abundance increased with distance from the mine site in all seasons, and caribou avoided areas within 4 km of the site in most seasons. Within 6 km of the mine centre, group size and the number of caribou decreased as mine activity progressed in late winter, pre-calving and calving seasons. Although the impact of the mine was most prominent in the pre-calving and calving seasons, caribou responded to mine disturbance in all seasons. This highlights the importance of evaluating the year-round impact of human-induced environmental change.
Article
Full-text available
Increasing outdoor activities by humans could negatively influence reindeer and caribou Rangifer tarandus populations. We recorded the behaviour of feral reindeer R. t. tarandus when a person directly approached them on foot or on skis in Forolhogna, Norway, during March, July and September–October 1996. The farther away the person was when first sighted, the greater the distance the reindeer group fled. The distance the reindeer moved away in response to the approaching person was greatest in July and least in September–October during autumn rut and shortly after the hunting season closed. In September–October rutting activities affected reindeer behaviour more than the disturbance caused by the directly approaching human. Both the distance at which the reindeer group responded by flight and the distance they moved away decreased with increasing group size. Upon flight, when all escape options were available, reindeer more often escaped uphill and into the wind than along level ground, downhill, down wind or crossways to the wind. All reindeer in a group moved towards the approaching human before taking final flight during 50% of 82 disturbance events, the closest approach was within 43 m in March, 24 m in July, and 13 m in September–October. No reindeer group responded by flight when the approaching human was still > 310 m away in March, > 351 m in July, and > 180 m in September–October. In relation to the current level of human activity in the area, our observations indicate no serious negative consequences for the reindeer following disturbance from a directly approaching human, not even shortly after the hunting season.
Article
Full-text available
We used census results and radiotelemetry locations of > 380 collared individuals sampled over the entire distribution of the endangered mountain ecotype of woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)) in British Columbia, Canada, to delineate population structure and document the size and trend of the identified populations. We also describe the spatial pattern of decline and the causes and timing of adult mortality and provide estimates of vital rates necessary to develop a population viability analysis. Our results indicate that the abundance of mountain caribou in British Columbia is declining. We found adult female annual survival rates below annual survival rates commonly reported for large ungulates. The major proximate cause of population decline appears to be predation on adult caribou. Spatial patterns of population dynamics revealed a continuous range contraction and an increasing fragmentation of mountain caribou into smaller, isolated subpopulations. The population fragmenta
Article
Full-text available
Mountain goat (Oreamnos americanus) responses to helicopter traffic were investigated at Caw Ridge (Alberta) from June to August 1995. A population of 109 marked individuals inhabited the ridge during the study. As measured by their overt responses, mountain goats were disturbed by 58% of the flights and were more adversely affected when heli- copters flew within 500 m. Eighty-five percent of flights within 500 m caused the goats to move >100 m; 9% of the flights >1,500 m away caused the goats to move similar dis- tances. Helicopter visibility and height above ground, number of goats in the group, group type (bachelor or nursery), and behavior of groups just prior to helicopter flights did not appear to influence reactions of goats to helicopters. Helicopter flights caused the disintegration of social groups on 25 occasions and resulted in 1 case of severe injury to an adult female. Based on these observations, restriction of helicopter flights within 2 km of alpine areas and cliffs that support mounta
Article
Full-text available
Potential avoidance by wild reindeer (Rangifer tarandus tarandus) of high-altitude tourist resorts during winter was investigated in and near Rondane National Park in Norway. Distribution of reindeer was mapped using systematic snowmobile surveys during 1991-96 and compared with results from investigations of snow and vegetation characteristics. Maternal reindeer avoided a 10 km zone around the resort. Cows and calves increased in density from 0.6 ± 0.6 reindeer km-2 at 5-10 km from the resort to 7.6 ± 2.2 reindeer km-2 at 15-25 km from the resort. Bulls and yearlings were more tolerant, constituting nearly 92% of all observed animals 5-10 km from the resort. Nearly all animals avoided the zone within 5 km of the resort. There were no significant differences in distribution of lichen heath, hardness of snow, integrated ram hardness index (IRH) values, or snow depths on ridges with increasing distance from the resort. Available biomass of lichens was ca. 1200 g m-2 0-5 km from the resort and decreased to a low of ca. 250 g m-2 at 15-25 km distance, a pattern that probably reflects overgrazing as a result of avoiding the tourist resort. Such avoidance implies reduced forage intake during winter, substantial reduction in available habitat, and lower productivity of the herd. The results suggest that avoidance by wild animals of sources of anthropogenic disturbance may involve long-term impacts, such as reductions in carrying capacity, that are more serious than those expected from direct physiological stress.
Article
Full-text available
We investigated the effects of hydroelectric development on the movements and space-use of caribou (Rangifer tarandus caribou) in west-central Newfoundland, Canada. We compared patterns of range use, site fidelity, and timing of migration before, during, and after project construction. Coincidental with the first year of project construction, caribou were less likely to be found within 3 km of the site; this persisted at least 2 years after construction was completed. Relative timing of migration was individual-specific; the rank order of spring arrival on, and autumn departure from, the calving and summer grounds tended to be consistent year-to-year. This is the first report of such individual-specific consistency in migration for a non-avian species. This predictability disappeared during development: the year-to-year consistency of fall and spring migration among individuals was apparent before and after construction, but not during construction. Variation in calving site fidelity was correlated to year-to-year differences in snowfall. We conclude that the development caused a disruption of migrational timing during construction and longer-term diminished use of the range surrounding the project site. Long-term studies of individually marked animals can aid in environmental assessments for migratory animals.
Article
Full-text available
Mountain sheep Ovis canadensis respond dramatically to helicopter disturbance. Significantly more animals abandoned sampling blocks and moved farther during helicopter surveys than on nonsurvey days throughout the year. Likewise, mountain sheep changed the vegetation type they occurred in more often after than before helicopter surveys; however, this difference was only significant during spring. Mountain sheep did not habituate or become sensitized to repeated helicopter overflights: time since capture was not related to their movements. The negative influence of the helicopter was extreme and may override variables that might otherwise be correlated with movement patterns of mountain sheep: this outcome also may hold for other ungulates. Further, sampling with helicopters may result in the violation of fundamental assumptions of population estimators routinely employed in conservation efforts for large mammals. The consequences of disturbing mountain sheep, such as altering use of habitat, increasing susceptibility to predation, or increasing nutritional stress, need additional study. These factors all have ramifications for the conservation of mountain sheep and other large mammals disturbed by helicopter sampling.
Article
Full-text available
Field endocrinology techniques allow the collection of samples (i.e., blood, urine, feces, tissues) from free-living animals for analysis of hormones, receptors, enzymes, etc. These data reveal mechanisms by which individuals respond to environmental challenges, breed, migrate and regulate all aspects of their life cycles. Field endocrinology techniques can also be used to address many issues in conservation biology. We briefly review past and current ways in which endocrine methods are used to monitor threatened species, identify potential stressors and record responses to environmental disturbance. We then focus on one important aspect of conservation: how free-living populations respond to human disturbance, particularly in relation to ecotourism. Breeding adult Magellanic penguins, Spheniscus magellanicus, appear to habituate well to tourists, and breed in an area where about 70,000 people visit during the season. Baseline levels of corticosterone return to normal after exposure of naïve birds to humans. However, penguin chicks appear to show a heightened adrenocortical response to handling stress in nests exposed to tourists, compared to chicks living in areas isolated from human intrusions. Given that developmental exposure to stress can have profound influences on how individuals cope with stress as adults, this potential effect of tourists on chicks could have long-term consequences. This field endocrine approach identified a stressor not observed through monitoring behavior alone.
Article
We investigated possible avoidance behavior of calving semi-domesticated reindeer (Rangifer tarandus tarandus) near recreational cabins, roads, and power transmission lines in Repparfjord Valley, northern Norway. The distribution, sex, and general age composition of the reindeer were mapped during the 1998 and 1999 calving seasons (n = 776 and n = 678, respectively) using systematic snowmobile and ski surveys. Mean reindeer density within preferred habitat was 78% lower in the area <4 km from the tourist resort compared to the area >4 km from the resort (1.47 vs. 6.68 reindeer/km2, respectively). Mean reindeer density by the power line corridor without traffic was 73% lower in the area <4 km from the power line compared to areas >4 km from the power line for comparable habitat. Areas <4 km from anthropogenic structures were avoided despite low levels of human traffic and a high proportion of preferred habitat. Within snow-free sites available for grazing, no significant differences occurred in phenological development of cotton-grass (Eriophorum spp.), in live:dead ratio of plant material, or in graminoid biomass with distance to the tourist resort. Almost 74% of all available forage was located within the avoided 0-4-km zones from the resort or the separate power line. Our results suggest that power lines, even without human traffic, may result in substantial reductions in the use of foraging areas. The combined actions of power lines, roads, and cabins may increase potential avoidance, thereby increasing the use of remaining undisturbed grazing grounds. Possible increased competition for high-quality forage may affect lactation, body condition, and, hence, reproductive success in the long term.
Article
Aerial surveys were conducted annually in June 1978-87 near Prudhoe Bay, Alaska, to determine changes in the distribution of calving caribou (Rangifer tarandus granfi) that accompanied petroleum-related development. With construction of an oil field access road through a calving concentration area, mean caribou density (no./km2) decreased from 1.41 to 0.31 (P = 0.05) within 1 km and increased from 1.41 to 4.53 (P = 0.04) 5- 6 km from the road. Concurrently, relative caribou use of the adjacent area declined (P < 0.02), apparently in response to increasing surface develop- ment. We suggest that perturbed distribution associated with roads reduced the capacity of the nearby area to sustain parturient females and that insufficient spacing of roads may have depressed overall calving activity. Use of traditional calving grounds and of certain areas therein appears to favor calf survival, principally through lower predation risk and improved foraging conditions. Given the possible loss of those habitats through dis- placement and the crucial importance of the reproductive process, a cautious approach to petroleum development on the Arctic Slope is warranted.
Article
We investigated changes in distribution and terrain use of calving barren-ground caribou (Rangifer tarandus granti) with increasing density of roads in the Kuparuk Development Area, an oil-field region near Prudhoe Bay, Alaska. In June of 1987-1992, caribou density, as determined by aerial surveys, was inversely related to road density, declining by 63% at >0.0-0.3 km road/km(2) and by 86% at >0.6-0.9 km road/km(2). The latter road density virtually excluded cow-calf pairs. Effects of avoidance were most apparent in preferred rugged terrain, comprising important habitats for foraging during the calving period. Our results show that (i) females and calves are far more sensitive to surface development than adult males and yearlings, (ii) the greatest incremental impacts are attributable to initial construction of roads and related facilities, and (iii) the extent of avoidance greatly exceeds the physical "footprint" of an oil-field complex. A disproportionate reduction in use of foraging habitats within the Kuparuk Development Area, combined with decreasing tolerance of the expanding industrial complex, may explain the recent displacement of some calving activity to areas farther inland, and, in part, lower fecundity. Possible consequences include heightened competition for forage, increased risk of predation, and lower productivity of the herd.
Article
The short-term impacts on caribou (Rangifer turandus) of low-level jet fighter training activity at Canadian Forces Base Goose Bay (Labrador) were investigated during the 1986-88 training seasons (April-October). Visual observations of low-level (30 m agl) jet overpasses indi- cated an initial stade response but otherwise brief overt reaction by woodland caribou on late-winter alpine tundra habitat. Between 1986 and 1988, daily effects of jet overflights were monitored on 10 caribou equipped with satellite-tracked radiocollars, which provided daily indices of activity and movement. Half the animals were exposed to jet overflights; the other 5 caribou were avoided during training exercises and therefore served as con- trol animals. In 1988, the control caribou were from a population that had never been overflown. Level of exposure to low-level flying within the exposed population did not significantly affect daily activity levels or distance travelled, although comparison with the unexposed population did suggest potential effects. The results indicate that significant impacts of low-level overflights can be minimized through a program of avoidance.
Article
Effects on woodland caribou (Rangifer tarandus caribou) of low-level military jet training at Canadian Forces Base - Goose Bay (Labrador) were studied during the 1986-88 training seasons. Calf survival was periodically monitored during 1987 and 1988 in a sample of 15 females wearing satellite-tracked radiocollars. During 1987, each female's exposure to low-level overflights was experimentally manipulated on a daily basis. In 1988, daily exposure was determined by analyzing jet flight tracks following the low-level flying season. Calf survival was monitored by survey flights every 3-4 weeks. A calf survival index, the number of survey periods (maximum = 4) that a cow was accompanied by a calf, was negatively correlated with the female's exposure to low-level jet overflights during the calving and immediate post-calving period and again during the period of insect harassment during summer. No significant relationship between calf survival and exposure to low-level flying was seen during the pre-calving period, during the late post-calving period prior to insect harassment, and during fall. In view of the continued depression of popula- tion growth in the woodland caribou population within the low-level training area, jets should avoid overflying woodland caribou calving range at least during the last week of May and the first three weeks of June.
Article
We investigated changes in distribution and terrain use of calving barren-ground caribou (Rangifer tarandus granti) with increasing density of roads in the Kuparuk Development Area, an oil-field region near Prudhoe Bay, Alaska. In June of 1987-1992, caribou density, as determined by aerial surveys, was inversely related to road density, declining by 63% at >0.0-0.3 km road/km2 and by 86% at >0.6-0.9 km road/km2. The latter road density virtually excluded cow-calf pairs. Effects of avoidance were most apparent in preferred rugged terrain, comprising important habitats for foraging during the calving period. Our results show that (i) females and calves are far more sensitive to surface development than adult males and yearlings, (ii) the greatest incremental impacts are attributable to initial construction of roads and related facilities, and (iii) the extent of avoidance greatly exceeds the physical "footprint" of an oil-field complex. A disproportionate reduction in use of foraging habitats within the Kuparuk Development Area, combined with decreasing tolerance of the expanding industrial complex, may explain the recent displacement of some calving activity to areas farther inland, and, in part, lower fecundity. Possible consequences include heightened competition for forage, increased risk of predation, and lower productivity of the herd.
Article
We evaluated elk (Cervus elaphus) reproductive success following removal of human disturbance during calving season, by comparing data from 2 segregated groups of free-ranging elk (control and treatment) from 1 pre-disturbance year, 2 disturbance years, and 2 post-disturbance years. Treatment-group elk were subjected to simulated recreational activity during calving season in disturbance years but not in pre- or post-disturbance years. Control animals experienced only ambient levels of disturbance throughout the study, and their calf/cow proportions (proportion of marked adult female elk maintaining a calf in Jul and Aug) were similar throughout the 5 years. We observed reduced productivity of treatment-group elk compared to controls during disturbance years after adjusting for nontreatment year differences. We hypothesized that productivity would return to, or potentially exceed, pre-disturbance levels following removal of disturbance. Productivity rebounded following release from disturbance, and full recovery was achieved by the second post-disturbance year. However, we did not observe productivity in excess of pre-disturbance years, as might be expected if release from energetic demands of maintaining a calf in 1 year increases probability of maintaining a calf in the following year. Our results are consistent with hypotheses that human-induced disturbance during parturition periods can reduce reproductive success and that removal of disturbance can allow productivity to recover to pre-disturbance levels. Managers of wildlife and wildlife habitat should consider potential impacts of human-induced disturbance on wildlife populations. Wildlife populations, depressed by human-induced disturbance during the neonatal period, may have the ability to rebound if the disturbance is removed.
Article
Seasonal variation, sex differences, and invasive sample collection may confound glucocorticoid measures as indices of stress. We investigated the effects of sex and season on glucocorticoid production on a non-invasive basis by measuring concentrations of cortisol metabolites in feces of undisturbed red deer (Cervus elaphus). Although feces can be collected easily, assignment to individuals is difficult. Anonymous fecal samples may cause overrepresentation of particular individuals thus introducing a source of error when estimating mean hormone levels within a population. We therefore examined the effects of collecting anonymous fecal samples on mean fecal cortisol metabolite levels. Neither sex nor sample collection mode significantly affected mean fecal cortisol metabolite concentrations in the studied population of red deer. Fecal glucocorticoid excretion varied seasonally with a peak during December and January. Out of several potential predictor variables investigated, minimum ambient temperature and snow proved to be the only factors exerting a significant effect on fecal glucocorticoid excretion. We suggest that high winter glucocorticoid levels may act via catabolic function during adaptation of deer to cold winter month when resources are limited.
Article
The responses of a herd of migratory woodland caribou (Rangifer tarandus caribou) to timber harvesting that fragmented about 11% of their winter range in west-central Alberta were examined in this study. From 1981 to 1996, 45 caribou were radio-collared and monitored during the initiation and completion of first-pass timber harvest (50% removal). Variables examined were home-range size, daily movement rates, and distance to the nearest cut block for radio-collared individuals. Daily movement rates and individual winter range sizes decreased as timber harvesting progressed. Caribou avoided using recently fragmented areas by an average of 1.2 km. If fragmentation of the winter range continues through timber harvesting and other industrial activities, the "spacing out" antipredator strategy used by caribou may be compromised. Based on these findings, timber-harvesting strategies are recommended that (i) ensure an adequate area of usable habitat to support the current population, (ii) minimize the amount of fragmented area, and (iii) in the short term avoid presently defined core use areas.
Article
Restricting human activity in elk (Cervus elaphus) calving areas during calving season can be controversial because of increasing human uses of elk habitat, and little evidence exists to evaluate impacts of these activities on elk populations. We evaluated effects of human-induced disturbance on reproductive success of radiocollared adult female elk using a control-treatment study in central Colorado. Data were collected during 1 pretreatment year and 2 treatment years. Treatment elk were repeatedly approached and displaced by study personnel throughout a 3-4-week period of peak calving during both treatment years, while control elk did not receive treatment. We observed elk on alpine summer ranges in July and August on both areas to estimate the proportion of marked cows maintaining a calf. Calf/cow proportions for the control area remained stable, but those for the treatment area declined each year. Average number of disturbances/elk/year effectively modeled variation in calf/cow proportions, supporting treatment as the cause of declining calf/cow proportions. Average decrease in calf/cow proportion in the treatment group was 0.225. Modeling indicated that estimated annual population growth on both study areas was 7% without treatment application given that existing human activities cause some unknown level of calving-season disturbance. With an average of 10 disturbances/cow above ambient levels, our model projected no growth. Our results support maintaining disturbance-free areas for elk during parturitional periods.
Article
Telemetered heart rates (HR) and behavioral responses of mountain sheep (Ovis canadensis canadensis) reacting to human disturbance in the Sheep River Wildlife Sanctuary, southwestern Alberta, were recorded. Cardiac and behavioral responses of sheep (4 ewes, 1 ram) to an approaching human were greatest when the person was accompanied by a dog or approached sheep from over a ridge. Reactions to road traffic were minimal as only 8.8% of vehicle passes elicited HR responses. No reactions to helicopters or fixed-wing aircraft were observed at distances exceeding 400 m from sheep. Responses to disturbance were detected using HR telemetry that were not evident from behavioral cues alone. However, mean duration of the HR response (138.6 sec) was not greater (P > 0.05) than mean period of the behavioral reaction when sheep were alert or withdrawing from harassing stimuli (112.4 sec). Use of HR telemetry in harassment research is discussed.
Article
Wilson and Hamilton. 2003. Cumulative effects of habitat change and recreation on mountain caribou ii Executive Summary We used a cumulative effects analysis to examine the effects of logging and/or burning and commercial backcountry ski use on mountain caribou habitat and habitat use in the Central Selkirk Mountains. We used forest inventory databases to infer logging and/or burning activity in caribou range during each decade 1960-2000. Skier-runs were summarized by month-year within zones defined by Canadian Mountain Holidays (CMH) for their Galena and Kootenay tenure areas. We also examined snowmobile use by interviewing users in Nakusp and Trout Lake. Areas and linear features (i.e., logging roads) used by snowmobilers were mapped and classified according to estimated use. Industrial forestry tenures cover 81% of the range of the Central Selkirks mountain caribou. Logging and/or burning disturbed >340 km 2 within the 6090 km 2 caribou range between 1960 and 2000. Caribou used these areas less than expected during the late-winter season, based on telemetry locations collected during 1992-2001. There was little or no net loss of high and very high-suitability habitat in the caribou range because lower quality habitats were recruited into higher quality habitats as forests aged from 1960 to 2000; however, this was partly an artefact of the forest age class breaks used in the suitability model. Some snowmobiling areas were located in important caribou habitat areas, as indicated by a proposed forestry zonation strategy for mountain caribou. There was no evidence that caribou used these areas less often than expected, based on univariate tests of our limited data.
Article
The effect of human activities on animal populations is widely debated, particularly since a recent decision by the U.S. Department of the Interior to ban snowmobiles from national parks. Immunoassays of fecal glucocorticoid levels provide a sensitive and noninvasive method of measuring the physiological stress responses of wildlife to disturbances. We tested for associations between snowmobile activity and glucocorticoid levels in an elk (Cervus elaphus) population in Yellowstone National Park and wolf (Canis lupus) populations in Yellowstone, Voyageurs, and Isle Royale national parks. For wolves, comparisons among populations and years showed that fecal glucocorticoid levels were higher in areas and times of heavy snowmobile use. For elk, day-to-day variation in fecal glucocorticoid levels paralleled variation in the number of snowmobiles after we controlled for the effects of weather and age. Also for elk, glucocorticoid concentrations were higher in response to snowmobiles than to wheeled vehicles after we controlled for the effects of age, weather, and number of vehicles. Despite these stress responses, there was no evidence that current levels of snowmobile activity are affecting the population dynamics of either species in these locations.
Article
Mountain caribou are an ecotype of woodland caribou (Rangifer tarandus caribou) that live in subalpine forests in southeastern British Columbia, Canada, extending into northern Idaho and Washington, USA. These caribou are listed as Threatened in Canada, Endangered in the United States, and are the subject of recovery planning efforts in both countries. Many areas of mountain caribou winter habitat experience intensive use by recreational snowmobilers. During 4 surveys, we recorded caribou on all 4 census blocks with little or no snowmobile activity (x̄ density = 0.41 caribou/km2), but during 3 of 4 years, we observed no caribou on the census block with intensive snowmobile activity. The year we observed caribou on the snowmobile block, most were using areas inaccessible to snowmobiles. We used a Resource Selection Function (RSF) based on radiotelemetry data for the area to compare habitat quality among the different census blocks. The absence of caribou from the intensive snowmobile area during most years could not be explained by differences in habitat quality. The RSF predicted that the intensive snowmobile area could support 53-96 caribou (95% CI). We conclude that intensive snowmobiling has displaced caribou from an area of suitable habitat. We recommend that snowmobile activity be restricted from all or most high-quality mountain caribou habitat as part of the recovery planning process.
Article
Short-term behavioural responses of 101 groups of Svalbard reindeer Rangifer tarandus platyrhynchus to direct provocation by a snowmobile were recorded in April (late winter) 1987. The median size of groups was 3·3 animals. The first visible responses of reindeer to an approaching snowmobile usually involved independent behaviour by different individuals in a group. Flight, by contrast, was a co-ordinated group response. Group median response distances were: minimum reaction distance, 640 m; disturbance distance, 410 m; distance at initial flight, 80 m; and distance of flight, 160 m. Group median response times were: total running time, 22 s; total location time, 38 s; maximum duration of disturbance, 193 s. Energy and time budget models indicate that one median flight response can cause an increase in a reindeer's daily energy expenditure of approximately 0·4% and a loss of daily grazing time also of 0·4%. The study failed to detect any way in which the current level of snowmobile traffic might substantially reduce the physical wellbeing of Svalbard reindeer.
Article
Noninvasive fecal glucocorticoid analysis has tremendous potential as a means of assessing stress associated with environmental disturbance in wildlife. However, interspecific variation in excreted glucocorticoid metabolites requires careful selection of the antibody used in their quantification. We compared four antibodies for detecting the major fecal cortisol metabolites in yellow baboons following 3H cortisol administration, ACTH challenge, and HPLC separation of fecal glucocorticoid metabolites. The most effective antibody (ICN corticosterone RIA; Cat. No. 07-120102) demonstrated relatively high cross-reactivities to the major cortisol metabolites present in feces during peak excretion, following both radiolabel infusion and ACTH challenge. This same antibody also detected increased fecal glucocorticoid metabolites after ACTH administration in the African elephant, black rhinoceros, Roosevelt elk, gerenuk, scimitar-horned oryx, Alaskan sea otter, Malayan sun bear, cheetah, clouded leopard, longtailed macaque, and northern spotted owl. Results suggest that (1) fecal glucocorticoid assays reliably detect endogenous changes in adrenal activity of a diverse array of species and (2) where comparisons were made, the ICN corticosterone antibody generally was superior to other antibodies for measuring glucocorticoid metabolites in feces.
Article
We assessed the impact of ecotourist visits during winter on woodland caribou Rangifer tarandus caribou time budgets in the Charlevoix Biosphere Reserve, Canada. We compared the behaviour of caribou during and after ecotourist visits with their behaviour during days without visits. In the presence of ecotourists, caribou increased time spent vigilant and standing, mostly at the expense of time spent resting and foraging. After visits, caribou tended to rest more than during control days. Caribou reduced time spent foraging during ecotourist visits as the number of observers increased. The impact of ecotourists appeared to decrease as winter progressed. Visits were short (=39.3 min) and caribou never left their winter quarters because of human presence. However, caribou abandoned their wintering area twice in response to wolf presence. Although winter is a difficult period of the year for caribou, our results suggest that with proper precautions caribou in Charlevoix can tolerate ecotourist visits.
Article
High rates of behavioural disruption caused by human activities could jeopardize the body condition and reproductive success of wildlife. I exposed Dall's sheep (Ovis dalli dalli) of the Yukon Territory to experimental overflights by a fixed-wing aircraft and a helicopter. Aircraft approaches that were more direct (as determined by the aircraft's elevation and horizontal distance from sheep) were more likely to elicit fleeing or to disrupt resting. Latency to resume feeding or resting after fixed-wing overflights was longer during more direct approaches. During indirect approaches by helicopters, sheep far from rocky slopes were much more likely to flee than sheep on rocky slopes. Sheep did not flee while nearby helicopters flew along the opposite side of a ridge, presumably because the obstructive cover buffered disturbing stimuli. Results provide preliminary parameters for predicting energetic and fitness costs incurred as a function of overflight rates, and can help mitigate disturbance by guiding temporal and spatial restrictions to aircraft.
Article
In most cooperatively breeding birds and mammals, reproductive rates are lower for social subordinates than for dominants, and it is common for reproduction in subordinates to be completely suppressed. Early research conducted in captivity showed that losing fights can increase glucocorticoid (GC) secretion, a general response to stress. Because GCs can suppress reproduction, it has been widely argued that chronic stress might underlie reproductive suppression of social subordinates in cooperative breeders. Contradicting this hypothesis, recent studies of cooperative breeders in the wild show that dominant individuals have elevated GCs more often than do subordinates. The findings that elevated GCs can be a consequence of subordination or a cost of dominance complicate the conventional view of social stress, with broad ramifications for the evolution of dominance and reproductive suppression.
Article
Southern Norway holds the last remaining population of wild reindeer (Rangifer tarandus tarandus) in Europe. Roads, railroads, and power lines have fragmented the original population into 26 separate herds. The reindeer populations are regulated directly according to availability of winter forage. These winter ranges, generally at lower elevations, are however, often subjected to development. Effects of infrastructure and associated human activity on the distribution of wild reindeer during winter were investigated in the Nordfjella mountain region of western Norway. Systematic aerial surveys of the distribution of ca. 2500 reindeer were conducted during late winter 1986–1998. Areas within 2.5 km from power lines were used less than available in 6 of the 8 sampling years, and areas beyond this zone more than expected. Density of reindeer was 79% lower within 2.5 km from power lines compared with background areas, and increased with increasing distance from infrastructure for comparable habitat. Available forage in terms of lichen cover declined 15–30-fold with distance, and was lowest in the undisturbed areas with the highest density of grazing animals. Areas within 5 km from resorts or from roads and power lines in combination were avoided in all years. Continued construction of roads, power lines and cabin resorts endanger these available winter ranges, and hence the long term survival of the population.
Article
The endangered, endemic Yellow-eyed penguin (Megadyptes antipodes) is one of the flagship species for New Zealand's wildlife tourism, and recently concern has been raised that tourism-related pressures may be becoming too great. We compared two neighbouring breeding areas exposed to different levels of human disturbance. Penguins at the site exposed to unregulated tourism showed significantly lower breeding success and fledging weights than those in an area visited infrequently for monitoring purposes only. High parental baseline corticosterone concentrations correlated with lower fledgling weights at both sites. Stress-induced corticosterone concentrations were significantly higher at the tourist-exposed site, suggesting birds have been sensitized by frequent disturbance. Consequences are likely to include reduced juvenile survival and recruitment to the tourist site, while the changed hormonal stress responses may ultimately have an effect on adult fitness and survival. For maintenance of attractive Yellow-eyed penguin-viewing destinations we recommend that tourists stay out of breeding areas and disturbance at penguin landing beaches is reduced.