Calving rate,calf survival rate,and habitat selection of forest-dwelling caribou in a highly managed landscape

Logging negatively affects the threatened forest-dwelling caribou (Rangifer tarandus caribou) through its positive effects on large predator populations. As recruitment is a key component of caribou population growth rate, we assessed calving rates of females and calf survival rates during the most critical period for calf survival, the calving period. We also identified causes of calf mortality and investigated the influence of predation risk, food availability, and human disturbance on habitat selection of females during the calving period at both the home-range and forest stand scales. We hypothesized that caribou should display habitat selection patterns to reduce predation risk at both scales. Using telemetry, we followed 22 females and their calves from 2004 to 2007 in a highly managed study area in Québec, Canada. Most females (78.5 ± 0.05 [SE]) gave birth each year, but only 46.3 ± 8.0% of the calves survived during the first 50 days following birth, and 57.3 ± 14.9% of them died from black bear (Ursus americanus) predation. At the home-range scale, caribou selected calving areas located at upper slope positions and avoided high road density areas. Surprisingly, they also selected the forested habitat type having the lowest lateral cover (mixed and deciduous stands) while avoiding the highest cover (regenerating conifer stands). At the forest stand scale, caribou selected areas located at relatively high elevations and with a lower basal area of black spruce trees. The selection of upper slope positions likely favored spatial segregation between calving females and wolves (Canis lupus) but not black bear. Our results suggest that calving females used areas from which they could visually detect approaching predators. While wolf avoidance appeared to be effective in a highly managed landscape, caribou did not appear to have adjusted their predator avoidance strategy to the recent increase in black bear abundance, who have benefited from increased food abundance. This situation requires focused attention from wildlife managers as logging activities are progressing towards the north within the core of forest-dwelling caribou range. © 2011 The Wildlife Society.     

Linking moose habitat selection to limiting factors

It has been suggested that patterns of habitat selection of animals across spatial scales should reflect the factors limiting individual fitness in a hierarchical fashion. Animals should thus select habitats that permit avoidance of the most important limiting factor at large spatial scales while the influence of less important factors should only be evident at fine scales. We tested this hypothesis by investigating moose Alces alces habitat selection using GPS telemetry in an area where the main factors limiting moose numbers were likely (in order of decreasing importance) predation risk, food availability and snow. At the landscape scale, we predicted that moose would prefer areas where the likelihood of encountering wolves was low or areas where habitats providing protection from predation were dominant. At the home‐range scale, we predicted that moose selection would be driven by food availability and snow depth. Wolf territories were delineated using telemetry locations and the study area was divided into 3 sectors that differed in terms of annual snowfall. Vegetation surveys yielded 6 habitat categories that differed with respect to food availability, and shelter from predation or snow. Our results broadly supported the hypothesis because moose reacted to several factors at each scale. At the landscape scale, moose were spatially segregated from wolves by avoiding areas receiving the lowest snowfall, but they also preferentially established their home range in areas where shelter from snow bordered habitat types providing abundant food. At the home‐range scale, moose also traded off food availability with avoidance of deep snow and predation risk. During winter, moose increased use of stands providing shelter from snow along edges with stands providing abundant food. Habitat selection patterns of females with calves differed from that of solitary moose, the former being associated primarily with habitats providing protection from predation. Animals should attempt to minimize detrimental effects of the main limiting factors when possible at the large scale. However, when the risk associated with several potential limiting factors varies with scale, we should expect animals to make trade‐offs among these.     

Woodland caribou habitat selection patterns in relation to predation risk and forage abundance depend on reproductive state

The ideal free distribution assumes that animals select habitats that are beneficial to their fitness. When the needs of dependent offspring differ from those of the parent, ideal habitat selection patterns could vary with the presence or absence of offspring. We test whether habitat selection depends on reproductive state due to top‐down or bottom‐up influences on the fitness of woodland caribou (Rangifer tarandus caribou), a threatened, wide‐ranging herbivore. We combined established methods of fitting resource and step selection functions derived from locations of collared animals in Ontario with newer techniques, including identifying calf status from video collar footage and seasonal habitat selection analysis through latent selection difference functions. We found that females with calves avoided predation risk and proximity to roads more strongly than females without calves within their seasonal ranges. At the local scale, females with calves avoided predation more strongly than females without calves. Females with calves increased predation avoidance but not selection for food availability upon calving, whereas females without calves increased selection for food availability across the same season. These behavioral responses suggest that habitat selection by woodland caribou is influenced by reproductive state, such that females with calves at heel use habitat selection to offset the increased vulnerability of their offspring to predation risk.     

Hierarchical habitat selection by woodland caribou: its relationship to limiting factors

Habitat selection is a hierarchical process that may yield various patterns depending on the scales of investigation. We employed satellite radio‐telemetry to examine patterns of habitat selection by female woodland caribou in central Saskatchewan at both coarse (seasonal range) and fine (daily area) scales. At each scale, we converted spatial data describing compositions of available and used habitat to standardised resource selection indices and examined them with multivariate analyses of variance. Seasonal ranges generally showed preferential inclusion of peatlands and black spruce dominated stands relative to recently disturbed stands and early seral stage forests. In all populations, caribou preferred peatlands and black spruce forests to all other habitat types at the daily area scale, in general, these patterns may reveal the effective avoidance of wolves, the primary factor limiting caribou throughout the boreal forest. In three populations where seasonal ranges showed the selective inclusion of either young jack pine stands or clearcuts along with peatlands and black spruce forests, we found a relative avoidance of the clearcuts and young jack pine stands at the daily area scale. As all caribou populations in the area are thought to be relics of a once more continuous distribution, the seasonal range selection by animals in disturbed areas may better describe historic rather than current habitat selection. We found inter‐annual variation in selection at the coarser spatial scale in one population, and inter‐seasonal variation in selection at the finer spatial scale in three populations, indicating that the relative grains of the spatial and temporal scales coincide. We were better able to explain the seasonal variations in finer scale selection by considering available forage, a factor less likely than predation to limit woodland caribou populations. The data agree with the theory that the spatial and temporal hierarchy of habitat selection reflects the hierarchy of factors potentially limiting individual fitness.     

Habitat selection by a large herbivore at multiple spatial and temporal scales is primarily governed by food resources

Habitat selection can be considered as a hierarchical process in which animals satisfy their habitat requirements at different ecological scales. Theory predicts that spatial and temporal scales should co‐vary in most ecological processes and that the most limiting factors should drive habitat selection at coarse ecological scales, but be less influential at finer scales. Using detailed location data on roe deer Capreolus capreolus inhabiting the Bavarian Forest National Park, Germany, we investigated habitat selection at several spatial and temporal scales. We tested 1) whether time‐varying patterns were governed by factors reported as having the largest effects on fitness, 2) whether the trade‐off between forage and predation risks differed among spatial and temporal scales and 3) if spatial and temporal scales are positively associated. We analysed the variation in habitat selection within the landscape and within home ranges at monthly intervals, with respect to land‐cover type and proxys of food and cover over seasonal and diurnal temporal scales. The fine‐scale temporal variation follows a nycthemeral cycle linked to diurnal variation in human disturbance. The large‐scale variation matches seasonal plant phenology, suggesting food resources being a greater limiting factor than lynx predation risk. The trade‐off between selection for food and cover was similar on seasonal and diurnal scale. Habitat selection at the different scales may be the consequence of the temporal variation and predictability of the limiting factors as much as its association with fitness. The landscape of fear might have less importance at the studied scale of habitat selection than generally accepted because of the predator hunting strategy. Finally, seasonal variation in habitat selection was similar at the large and small spatial scales, which may arise because of the marked philopatry of roe deer. The difference is supposed to be greater for wider ranging herbivores.     

Consequences of migratory strategy on habitat selection by mule deer

Ungulate behavior is often characterized as balancing selection for forage and avoidance of predation risk. Within partially migratory ungulate populations, this balancing occurs across multiple spatial scales, potentially resulting in different exposure to costs and benefits between migrants and residents. We assessed how availability and selection of forage and risk from predators varied between summer ranges of migrant and resident mule deer (Odocoileus hemionus; a species in which individual migratory strategies are generally fixed for life) in 3 study areas in western Montana, USA, during summers 2017–2019. We hypothesized that mule deer would face a tradeoff between selecting forage and avoiding predation risk, and that migration and residency would pose contrasting availability of forage and risk at a broad (summer range) spatial scale. We hypothesized deer exposed to lower forage at a given spatial scale would compensate for reduced availability by increasing selection of forage at the cost of reduced avoidance of predators, a mechanism whereby migrants and residents could potentially achieve similar exposure to forage despite disparate availability. We compared the availability of forage (kcal/m²) and predation risk from wolves (Canis lupus) and mountain lions (Puma concolor) between summer ranges of each migratory strategy, then assessed how selection for those factors at the home range (second order) and within-home range (third order) scales varied using resource selection functions (RSFs). As forage availability increased among mule deer summer ranges and individual home ranges, selection for forage decreased at the second-order (P = 0.052) and third-order (P = 0.081) scales, respectively, but avoidance of predators varied weakly. In 1 study area, summer range of residents contained lower forage and higher risk than summer range of migrants, but residents compensated for this disadvantage through stronger selection of forage and avoidance of risk at finer spatial scales. In the other 2 study areas, summer range of migrants contained lower forage and higher risk than residents, but migrants did not compensate through stronger selection for beneficial resources. The majority of mule deer in our study system were migratory, though the benefits of migration were unclear, suggesting partial migration may persist in populations even when exposure to forage and predation risk appears unequal between strategies.     

Mothers' Movements: Shifts in Calving Area Selection by Partially Migratory Elk

Loss of migratory behavior or shifts in migratory ranges are growing concerns to wildlife managers. How ungulates prioritize safety from predators at the expense of high-quality foraging opportunities during calving may be key to understanding these shifts and long-term reproductive success. We compared trade-offs in selection for forage and predation risk by elk (Cervus canadensis) following 3 migratory tactics (western and eastern migration and resident) during 2 time periods in a declining (by almost 70% from 2002–2016), partially migratory elk population adjacent to Banff National Park in Alberta, Canada. We hypothesized that regardless of migratory tactic, maternal elk would show stronger trade-offs between high-quality foraging associated with higher predation risk and forage resources of lower-quality yet lower risk on calving ranges than on ranges used during summer because of vulnerability of their offspring. Additionally, we hypothesized these trade-offs would occur at high (2002–2006) and low (2013–2016) elk population sizes. We used a machine-learning algorithm to predict dates of parturition based on global positioning system (GPS) movements of elk equipped with vaginal implants (n = 60) and predictions were within 1.43 ± 0.85 (SE) days of the known date. We applied the model to an additional 58 GPS-collared elk without vaginal implants. Based on changes in localized movements, we defined calving areas as the 26 days post-parturition and compared habitat characteristics of calving areas to 10 similar-sized areas centered on random locations during summer for the same individual in a latent selection framework. Across the 2 time periods, parturition occurred from 8 May–11 July with median parturition dates differing among migratory tactics and residents shifting towards an earlier parturition date in the later period. All elk, regardless of migratory tactic and time period, selected calving areas with greater forage resources than were available on areas used during summer, with no evidence for greater selection of areas that reduced predation risk at the expense of higher-quality foraging. Calving season selection for areas with abundant forage exposed western migrants to high risk of bear (Ursus spp.) predation, residents to high risk of wolf (Canis lupus) predation, and eastern migrants to low risk of bear and wolf predation. Patterns in exposure to predation risk during calving between migratory tactics were consistent with the recent decline in western migrants and increase in eastern migrants, implying that conditions on calving areas contributed to observed changes in the number of elk following these tactics. © 2021 The Wildlife Society.     

Habitat selection by an Alpine ungulate: the significance of forage characteristics varies with scale and season

Habitat selection in ungulates should ensure access to abundant forage of sufficiently high quality. Species living in rugged mountain areas have to face nutritional bottlenecks regularly and should show particularly sophisticated habitat selection behaviour. However, patterns and mechanisms of such adaptations remain little studied. We analysed habitat selection and its seasonal variability of 10 GPS‐collared red deer Cervus elaphus living in a topographically challenging landscape of the Swiss Alps. We hypothesised that resource selection by red deer was scale‐dependent and predicted that scale‐dependence would vary among seasons in relation to seasonal changes of available forage biomass and quality, which we sampled across the entire study area of 250 km². The studied population of Alpine red deer undertook altitudinal migrations and showed scale‐dependent habitat selection that was strongest in winter and declined through spring and summer. Selection occurred mostly at the larger (landscape/home‐range location) scale and less so at the smaller (within home‐range) scale. Topographic parameters were selected mainly at the landscape scale and mostly in winter. About 70% of all instances of preference for habitat parameters were associated with above‐average forage characteristics, represented mostly by higher crude protein content, in a few cases also by higher biomass or both. The overall pattern of space use by red deer characterised by migration and seasonal habitat selection was therefore closely linked to the quality of food resources, although some trade‐offs with avoiding human disturbance may also have been involved.     

Habitat structure and association with ungulates drive habitat selection and grouping behaviour of lesser rhea (Rhea pennata subsp. pennata)

Social prey species respond to predation risk by modifying habitat selection and grouping behaviour. These responses may depend on both actual predation risk (predator probability of occurrence) and/or on perceived predation risk associated with habitat structure. Other factors like food availability and co-occurrence with other species may also affect habitat selection and group formation. We analyse habitat selection and grouping behaviour (group size and cohesion) of lesser rhea (Rhea pennata subsp. pennata), a ratite endemic of South America inhabiting steppe shrublands and grasslands, in relation to actual (puma probability of occurrence) and perceived (habitat structure: openness, visibility) predation risk, co-occurrence with other herbivore species and forage availability in the Chilean Patagonia. We used data from 9 sampling seasons in 5 years. Results show that habitat selection, group size and cohesion in lesser rhea were mainly driven by variables associated with perceived predation risk and by co-occurrence with other herbivores both during breeding and non–breeding season. As expected, lesser rhea preferred open habitats (vegas and grasslands) that allow a behaviour of ‘watch and run’ to avoid predation and formed larger groups in them. Moreover, lesser rhea positively selected year-round habitats where livestock occur, forming large groups during non–breeding season there. Group size and co-occurrence with other herbivores significantly decreased group cohesion, suggesting a reduction of perceived predation risk. Therefore, lesser rhea seems to take advantage of forming mixed interspecific groups to reduce predation risk. These results suggest that lesser rhea habitat selection and grouping behaviour are preferentially driven by factors related to perceived predation risk than by actual predator occurrence or food availability.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

Coyote Habitat Selection and Management Implications for the Gaspésie Caribou

ABSTRACT Anthropogenic disturbances can promote establishment and growth of predator populations in areas where secondary prey can then become threatened. In this study, we investigated habitat selection of eastern coyotes (Canis latrans), a relatively new predator in the vicinity of an endangered population of caribou (Rangifer tarandus caribou). We hypothesized that coyotes in the boreal forest depend mainly on disturbed habitat, particularly that of anthropogenic origin, because these habitats provide increased food accessibility. Coyotes would likely take advantage of moose (Alces alces) carcasses, berries, and snowshoe hares (Lepus americanus) found in open habitats created by logging. To test these predictions, we described coyote diet and habitat selection at different spatial and temporal levels and then compared resource availability between habitats. To do so, we installed Global Positioning System radiocollars on 23 individual coyotes in the Gaspésie Peninsula, eastern Québec, Canada. Coyotes selected clear-cuts of 5–20 years and avoided mature coniferous forests both at the landscape and home-range levels. Clear-cuts of 5–20 years were found to contain a high availability of moose carcasses and berries, and vulnerability of snowshoe hares is known to increase in clear-cuts. The importance of these 3 food resources was confirmed by the characteristics of core areas used by coyotes and diet analysis. Moose remains were found at 45% of core areas and coyote diet comprised 51% moose on an annual basis. Anthropogenic disturbances in the boreal forest thus seem to benefit coyotes. Our results indicated that the relationship between coyotes and caribou likely involves spillover predation. This knowledge allows managers to consider spillover predation by coyotes as a possible threat for endangered caribou population when the predator depends mainly on habitat of anthropogenic origin and to suggest methods to alleviate it when developing management plans.     

Female Moose Prioritize Forage Over Mortality Risk in Harvested Landscapes

Since 2010, several moose (Alces alces) populations have declined across North America. These declines are believed to be broadly related to climate and landscape change. At the western reaches of moose continental range, in the interior of British Columbia, Canada, wildlife managers have reported widespread declines of moose populations. Disturbances to forests from a mountain pine beetle (Dendroctonum ponderosae) outbreak and associated salvage logging infrastructure in British Columbia are suspected as a mechanism manifested in moose behavior and habitat selection. We examined seasonal differences in moose habitat selection in response to landscape change from mountain pine beetle salvage logging infrastructure: dense road networks and large intensive forest harvest cutblocks. We used 157,447 global positioning system locations from 83 adult female moose from 2012 to 2016 on the Bonaparte Plateau at the southern edge of the Interior Plateau of central British Columbia to test whether increased forage availability, landscape features associated with increased mortality risk, or the cumulative effects of salvage logging best explain female moose distribution using resource selection functions in an information-theoretic framework. We tested these hypotheses across biological seasons, defined using a cluster analysis framework. The cumulative effects of forage availability and risk best predicted resource selection of female moose in all seasons; however, the covariates included in the cumulative models varied between seasons. The top forage availability model better explained moose habitat use than the top risk model in all seasons, except for the calving and fall seasons where the top risk model (distance to road) better predicted moose space use. Selection of habitat that provides forage in winter, spring, and summer suggests that moose seasonally trade predation risk for the benefits of foraging in early seral vegetation communities in highly disturbed landscapes. Our results identified the need for intensive landscape-scale management to stem moose population declines. Additional research is needed on predator densities, space use, and calf survival in relation to salvage logging infrastructure. © 2020 The Wildlife Society.     

Behavioural strategies towards human disturbances explain individual performance in woodland caribou

Behavioural strategies may have important fitness, ecological and evolutionary consequences. In woodland caribou, human disturbances are associated with higher predation risk. Between 2004 and 2011, we investigated if habitat selection strategies of female caribou towards disturbances influenced their calf’s survival in managed boreal forest with varying intensities of human disturbances. Calf survival was 53 % and 43 % after 30 and 90 days following birth, respectively, and 52 % of calves that died were killed by black bear. The probability that a female lose its calf to predation was not influenced by habitat composition of her annual home range, but decreased with an increase in proportion of open lichen woodland within her calving home range. At the local scale, females that did not lose their calf displayed stronger avoidance of high road density areas than females that lost their calf to predation. Further, females that lost their calf to predation and that had a low proportion of ≤5-year-old cutovers within their calving home range were mostly observed in areas where these young cutovers were locally absent. Also, females that lost their calf to predation and that had a high proportion of ≤5-year-old cutovers within their calving home range were mostly observed in areas with a high local density of ≤5-year-old cutovers. Our study demonstrates that we have to account for human-induced disturbances at both local and regional scales in order to further enhance effective caribou management plans. We demonstrate that disturbances not only impact spatial distribution of individuals, but also their reproductive success.     

Habitat and introduced predators influence the occupancy of small threatened macropods in subtropical Australia

Australia has had the highest rate of mammal extinctions in the past two centuries when compared to other continents. Frequently cited threats include habitat loss and fragmentation, changed fire regimes and the impact of introduced predators, namely the red fox (Vulpes vulpes) and the feral cat (Felis catus). Recent studies suggest that Australia's top predator, the dingo (Canis dingo), may have a suppressive effect on fox populations but not on cat populations. The landscape of fear hypothesis proposes that habitat used by prey species comprises high to low risk patches for foraging as determined by the presence and ubiquity of predators within the ecosystem. This results in a landscape of risky versus safe areas for prey species. We investigated the influence of habitat and its interaction with predatory mammals on the occupancy of medium‐sized mammals with a focus on threatened macropodid marsupials (the long‐nosed potoroo [Potorous tridactylous] and red‐legged pademelon [Thylogale stigmatica]). We assumed that differential use of habitats would reflect trade‐offs between food and safety. We predicted that medium‐sized mammals would prefer habitats for foraging that reduce the risk of predation but that predators would have a positive relationship with medium‐sized mammals. We variously used data from 298 camera trap sites across nine conservation reserves in subtropical Australia. Both dingoes and feral cats were broadly distributed, whilst the red fox was rare. Long‐nosed potoroos had a strong positive association with dense ground cover, consistent with using habitat complexity to escape predation. Red‐legged pademelons showed a preference for open ground cover, consistent with a reliance on rapid bounding to escape predation. Dingoes preferred areas of open ground cover whereas feral cats showed no specific habitat preference. Dingoes were positively associated with long‐nosed potoroos whilst feral cats were positively associated with red‐legged pademelons. Our study highlights the importance of habitat structure to these threatened mammals and also the need for more detailed study of their interactions with their predators.     

Elk parturition site selection at local and landscape scales

Selection of habitat components by ungulates associated with parturition sites varies among and within species depending upon vulnerability to predators, variation in local topography and climate regimes, and the length of time that the maternal–neonatal unit spends at or near the parturition location. We marked 169 parturition locations of elk (Cervus elaphus nelsoni) in western Wyoming using vaginal implant transmitters and evaluated parturition-specific habitat selection at macro- and microhabitat scales using a resource selection function modeling approach. Elk calved in a variety of habitats, yet demonstrated selection at both spatial scales. We found the strongest support for models that incorporated multiple habitat features and focused on topographical and vegetative cover types that provide physical and thermal cover at the macrohabitat scale and for visual cover models at the microhabitat scale. Models based solely on forage availability or quality were least supported at both scales, which may be indicative of a brief occupation of the parturition location or low heterogeneity in the availability of forage resources on parturition ranges. Results of early elk natural history studies may have represented a bias introduced by variable sightability and accessibility of females with calves and a lack of differentiation between calving and neonatal periods. More clearly defining calving site selection and removing biases toward more open habitats where sightability of neonates is greater may be used by wildlife or land managers to improve or protect calving habitats, which is often a stated objective of management actions. The results of this study suggest that microhabitat is more important to elk and that temporal closures over broad areas versus closures focused on specific macrohabitats may be more effective in protecting calving animals. © 2011 The Wildlife Society.     

Avoidance of roads and selection for recent cutovers by threatened caribou: fitness-rewarding or maladaptive behaviour?

The impact of anthropogenic disturbance on the fitness of prey should depend on the relative effect of human activities on different trophic levels. This verification remains rare, however, especially for large animals. We investigated the functional link between habitat selection of female caribou (Rangifer tarandus) and the survival of their calves, a fitness correlate. This top-down controlled population of the threatened forest-dwelling caribou inhabits a managed forest occupied by wolves (Canis lupus) and black bears (Ursus americanus). Sixty-one per cent of calves died from bear predation within two months following their birth. Variation in habitat selection tactics among mothers resulted in different mortality risks for their calves. When calves occupied areas with few deciduous trees, they were more likely to die from predation if the local road density was high. Although caribou are typically associated with pristine forests, females selected recent cutovers without negative impact on calf survival. This selection became detrimental, however, as regeneration took place in harvested stands owing to increased bear predation. We demonstrate that human disturbance has asymmetrical consequences on the trophic levels of a food web involving multiple large mammals, which resulted in habitat selection tactics with a greater short-term fitness payoff and, therefore, with higher evolutionary opportunity.     

Comment arising from a paper by Wittmer et al.: hypothesis testing for top-down and bottom-up effects in woodland caribou population dynamics

Conservation strategies for populations of woodland caribou Rangifer tarandus caribou frequently emphasize the importance of predator–prey relationships and the availability of lichen-rich late seral forests, yet the importance of summer diet and forage availability to woodland caribou survival is poorly understood. In a recent article, Wittmer et al. (Can J Zool 83:407–418, 2005b) concluded that woodland caribou in British Columbia were declining as a consequence of increased predation that was facilitated by habitat alteration. Their conclusion is consistent with the findings of other authors who have suggested that predation is the most important proximal factor limiting woodland caribou populations (Bergerud and Elliot in Can J Zool 64:1515–1529, 1986; Edmonds in Can J Zool 66:817–826, 1988; Rettie and Messier in Can J Zool 76:251–259, 1998; Hayes et al. in Wildl Monogr 152:1–35, 2003). Wittmer et al. (Can J Zool 83:407–418, 2005b) presented three alternative, contrasting hypotheses for caribou decline that differed in terms of predicted differences in instantaneous rates of increase, pregnancy rates, causes of mortality, and seasonal vulnerability to mortality (Table 1, p 258). These authors rejected the hypotheses that food or an interaction between food and predation was responsible for observed declines in caribou populations; however, the use of pregnancy rate, mortality season and cause of mortality to contrast the alternative hypotheses is problematic. We argue here that the data employed in their study were insufficient to properly evaluate a predation-sensitive foraging hypothesis for caribou decline. Empirical data on seasonal forage availability and quality and plane of nutrition of caribou would be required to test the competing hypotheses. We suggest that methodological limitations in studies of woodland caribou population dynamics prohibit proper evaluation of the mechanism of caribou population declines and fail to elucidate potential interactions between top-down and bottom-up effects on populations. An erratum to this article can be found at     

Mechanisms of species coexistence: a field test of theoretical models using intertidal snails

Competitor coexistence is often facilitated by spatial segregation. Traditionally, spatial segregation is predicted to occur when species differ in the habitat in which they are either superior at competing for resources or less susceptible to predation. However, predictions from a behavioural model demonstrate that spatial segregation and coexistence can also occur in the absence of such interspecific trade‐offs in competitive ability and vulnerability to predation. Unlike other models of competitor coexistence this model predicts that when species rank both habitat productivity and ‘riskinesses’ similarly, but differ slightly in their habitat‐specific vulnerabilities to predators, they will tend to segregate across habitats, with the species experiencing the higher ratio of mortality risk across the habitats occurring primarily in the safer habitat. Here, we investigate the hypothesis that intraspecific trade‐offs between resource availability and mortality risk can lead to spatial segregation of competing species by (1) documenting the spatial (i.e. intertidal) distribution of two marine snails, Littorina sitkana and L. subrotundata and (2) performing field experiments to quantify growth and mortality rates of each species at ‘low’ and ‘high’ intertidal heights. Our results indicate that both species agree on the rankings of habitat riskiness and productivity, experiencing higher predation and higher growth in low‐ than in high‐intertidal habitats. However, L. sitkana and L. subrotundata experienced differences in their habitat‐specific mortality risks and growth rates. Despite both species being similarly at risk of predation in high‐intertidal habitats (where mortality was lower), L. subrotundata was subject to significantly higher mortality than L. sitkana at the low‐intertidal height. In contrast, growth rate differences between habitats were greater for L. sitkana than for L. subrotundata. Whereas both species grew at the same rate at the high‐intertidal level (where growth was lower), L. sitkana individuals grew more rapidly than L. subrotundata snails at the low‐intertidal level. As predicted by the behavioural model, the species that experienced the higher ratio of mortality across habitats (i.e. L. subrotundata) occurred exclusively in the safer, high‐intertidal habitat. Taken together, these results provide support for the hypothesis that spatial segregation, and potentially competitor coexistence, can occur in the absence of interspecific trade‐offs in resource acquisition ability or vulnerability to predation.     

Using habitat selection theories to predict the spatiotemporal distribution of migratory birds during stopover – a case study of pink‐footed geese Anser brachyrhynchus

Understanding how animals select for habitat and foraging resources therein is a crucial component of basic and applied ecology. The selection process is typically influenced by a variety of environmental conditions including the spatial and temporal variation in the quantity and quality of food resources, predation or disturbance risks, and inter‐ and intraspecific competition. Indeed, some of the most commonly employed ecological theories used to describe how animals choose foraging sites are: nutrient intake maximisation, density‐dependent habitat selection, central‐place foraging, and predation risk effects. Even though these theories are not mutually exclusive, rarely are multiple theoretical models considered concomitantly to assess which theory, or combination thereof, best predicts observed changes in habitat selection over space and time. Here, we tested which of the above theories best‐predicted habitat selection of Svalbard‐breeding pink‐footed geese at their main spring migration stopover site in mid‐Norway by computing a series of resource selection functions (RSFs) and their predictive ability (k‐fold cross validation scores). At this stopover site geese fuel intensively as a preparation for breeding and further migration. We found that the predation risk model and a combination of the density‐dependent and central‐place foraging models best‐predicted habitat selection during stopover as geese selected for larger fields where predation risk is typically lower and selection for foraging sites changed as a function of both distance to the roost site (i.e. central‐place) and changes in local density. In contrast to many other studies, the nutritional value of the available food resources did not appear to be a major limiting factor as geese used different food resources proportional to their availability. Our study shows that in an agricultural landscape where nutritional value of food resources is homogeneously high and resource availability changes rapidly; foraging behaviour of geese is largely a tradeoff between fast refuelling and disturbance/predator avoidance.     

Spatial scales of habitat selection decisions: implications for telemetry‐based movement modelling

Movement influences a myriad of ecological processes operating at multiple spatial and temporal scales. Yet our understanding of animal movement is limited by the resolution of data that can be obtained from individuals. Traditional approaches implicitly assume that movement decisions are made at the spatial and temporal scales of observation, although this scale is typically an artifact of data‐gathering technology rather than biological realism. To address this limitation, we used telemetry‐based movement data for caribou Rangifer tarandus in Newfoundland, Canada, and compared movement decisions estimated at the temporal resolution of GPS relocations (2 h) to a novel model describing directional movement to areas reachable over an extended period. We showed that this newer model is a better predictor of movement decisions by caribou, with decisions made at the scale of ～2 km, including the strong avoidance of dense coniferous forest, an outcome not detectable at the scale of GPS relocations. These results illustrate the complexity of factors affecting animal movement decisions and the analytical challenges associated with their interpretation. Our novel modelling framework will help support increased accuracy in predictive models of animal space‐use, and thereby aid in determining biologically meaningful scales for collecting movement and habitat data.