Jodie Martin

Selecting habitat to survive: The impact of road density on survival in a large carnivore

Habitat selection studies generally assume that animals select habitat and food resources at multiple scales to maximise their fitness. However, animals sometimes prefer habitats of apparently low quality, especially when considering the costs associated with spatially heterogeneous human disturbance. We used spatial variation in human disturbance, and its consequences on lynx survival, a direct fitness component, to test the Hierarchical Habitat Selection hypothesis from a population of Eurasian lynx Lynx lynx in southern Norway. Data from 46 lynx monitored with telemetry indicated that a high proportion of forest strongly reduced the risk of mortality from legal hunting at the home range scale, while increasing road density strongly increased such risk at the finer scale within the home range. We found hierarchical effects of the impact of human disturbance, with a higher road density at a large scale reinforcing its negative impact at a fine scale. Conversely, we demonstrated that lynx shifted their habitat selection to avoid areas with the highest road densities within their home ranges, thus supporting a compensatory mechanism at fine scale enabling lynx to mitigate the impact of large-scale disturbance. Human impact, positively associated with high road accessibility, was thus a stronger driver of lynx space use at a finer scale, with home range characteristics nevertheless constraining habitat selection. Our study demonstrates the truly hierarchical nature of habitat selection, which aims at maximising fitness by selecting against limiting factors at multiple spatial scales, and indicates that scale-specific heterogeneity of the environment is driving individual spatial behaviour, by means of trade-offs across spatial scales.

Coping with spatial heterogeneity and temporal variability in resources and risks: adaptive movement behaviour by a large grazing herbivore.

Movement is a key mean for mobile species to cope with heterogeneous environments. While in herbivorous mammals large-scale migration has been widely investigated, fine-scale movement responses to local variations in resources and predation risk remain much less studied, especially in savannah environments. We developed a novel approach based on complementary movement metrics (residence time, frequency of visits and regularity of visits) to relate movement patterns of a savannah grazer, the blue wildebeest Connochaetes taurinus, to fine-scale variations in food availability, predation risk and water availability in the Kruger National Park, South Africa. Wildebeests spent more time in grazing lawns where the grass is of higher quality but shorter than in seep zones, where the grass is of lower quality but more abundant. Although the daily distances moved were longer during the wet season compared to the dry season, the daily net displacement was lower, and the residence time higher, indicating a more frequent occurrence of area-concentred searching. In contrast, during the late dry season the foraging sessions were more fragmented and wildebeests moved more frequently between foraging areas. Surprisingly, predation risk appeared to be the second factor, after water availability, influencing movement during the dry season, when resources are limiting and thus expected to influence movement more. Our approach, using complementary analyses of different movement metrics, provided an integrated view of changes in individual movement with varying environmental conditions and predation risk. It makes it possible to highlight the adaptive behavioral decisions made by wildebeest to cope with unpredictable environmental variations and provides insights for population conservation.

Identifying space use at foraging arena scale within the home ranges of large herbivores.

An intermediate spatiotemporal scale of food procurement by large herbivores is evident within annual or seasonal home ranges. It takes the form of settlement periods spanning several days or weeks during which foraging activity is confined to spatially discrete foraging arenas, separated by roaming interludes. Extended by areas occupied for other activities, these foraging arenas contribute towards generating the home range structure. We delineated and compared the foraging arenas exploited by two African large herbivores, sable antelope (a ruminant) and plains zebra (a non-ruminant), using GPS-derived movement data. We developed a novel approach to specifically delineate foraging arenas based on local change points in distance relative to adjoining clusters of locations, and compared its output with modifications of two published methods developed for home range estimation and residence time estimation respectively. We compared how these herbivore species responded to seasonal variation in food resources and how they differed in their spatial patterns of resource utilization. Sable antelope herds tended to concentrate their space use locally, while zebra herds moved more opportunistically over a wider set of foraging arenas. The amalgamated extent of the foraging arenas exploited by sable herds amounted to 12-30 km2, compared with 22-100 km2 for the zebra herds. Half-day displacement distances differed between settlement periods and roaming interludes, and zebra herds generally shifted further over 12h than sable herds. Foraging arenas of sable herds tended to be smaller than those of zebra, and were occupied for period twice as long, and hence exploited more intensively in days spent per unit area than the foraging arenas of zebra. For sable both the intensity of utilization of foraging arenas and proportion of days spent in foraging arenas relative to roaming interludes declined as food resources diminished seasonally, while zebra showed no seasonal variation in these metrics. Identifying patterns of space use at foraging arena scale helps reveal mechanisms generating the home range extent, and in turn the local population density. Thereby it helps forge links between behavioural ecology, movement ecology and population ecology.

Spatially nested niche partitioning between syntopic grazers at foraging arena scale within overlapping home ranges

Niche separation among species with similar resource requirements can be expressed at various spatiotemporal scales, from the resource components selected at feeding sites to habitat and home range occupation and ultimately geographic distribution ranges. African large herbivores present a challenge to niche theory because multiple species commonly overlap both spatially and in vegetation components consumed. Aided by GPS telemetry, we investigated the space use patterns of two large grazers that are frequently associated in mixed-species aggregations. Specifically, we compared a generalist grazer with hindgut fermentation (plains zebra) with a similar-sized grazing ruminant (blue wildebeest) in west-central Kruger National Park, South Africa. We found that herds of the two species overlapped substantially in the home ranges that they occupied, but exploited spatially distinct foraging arenas for periods lasting several days or weeks within these ranges. Moreover, wildebeest and zebra differed in duration of settlement, extent of areas occupied during settlement, consequent exploitation intensity per unit area, proportion of time spent within foraging arenas relative to roaming interludes, and movement rates while within these arenas. In particular, wildebeest herds concentrated within small areas for prolonged periods, while zebra herds used more foraging arenas but exploited them for briefer periods. Both species overlapped substantially in habitat use, although wildebeest more strongly favored gabbro uplands and sodic sites presenting short grass lawns while zebra made greater use of areas with a taller grass cover. Hence resource partitioning was expressed mainly through behavioral distinctions in patch exploitation at foraging arena scale rather than in home range or habitat separation. Although zebra may have been partially excluded from the grasslands kept short by wildebeest, these sites formed only a small part of the wider ranges utilized by zebra, thereby restricting the competitive consequences. Hence spatially nested resource partitioning of this form contributes to the coexistence of these two grazers, and may be a mechanism enabling niche separation among other species.

Habitat selectivity influences the reactive responses of African ungulates to encounters with lions

Prey mortality depends on the likelihood of encountering a predator and of being killed following such an encounter. While the proactive responses of large herbivores to the risk of predation have been well documented, the reactive responses of prey following encounters have been less well studied and restricted mostly to northern temperate ecosystems. We postulated that the reactive response should depend on the habitat selection of the prey species as well as the hunting tactic (cursorial versus stalking) of the predator. We investigated the reactive responses of a selective grazer, the blue wildebeest, Connochaetes taurinus, and a generalist grazer, the plains zebra, Equus quagga, to encounters with lions, Panthera leo, their main predator, in Kruger National Park. We used simultaneous global positioning system (GPS) locations of prey and predators to identify encounters and assess their frequency and characteristics (e.g. time of day). We compared both the immediate response (stay or flee) and subsequent movement (remain in the habitat patch or relocate) of the two ungulate species following encounters with lions. Despite being more predictably located in open short grass areas, wildebeest encountered lions no more frequently than zebra. However, being more selective for these particular habitat types, wildebeest more often stayed near the encounter site than zebra. Specifically, wildebeest were less likely to leave when located in more open areas with short grass providing both high-quality food and little concealment for stalking lions. We suggest that this is because wildebeest incur higher risks and greater feeding costs than zebra should they relocate, owing to their narrower preference for areas of short grass. Our findings show how the adaptive responses of wildebeest and zebra following encounters with their major predator depend on their habitat requirements.

Temporal shifts in landscape connectivity for an ecosystem engineer, the roe deer, across a multiple-use landscape

ContextRoutine movements of large herbivores, often considered as ecosystem engineers, impact key ecological processes. Functional landscape connectivity for such species influences the spatial distribution of associated ecological services and disservices.ObjectivesWe studied how spatio-temporal variation in the risk-resource trade-off, generated by fluctuations in human activities and environmental conditions, influences the routine movements of roe deer across a heterogeneous landscape, generating shifts in functional connectivity at daily and seasonal time scales.MethodsWe used GPS locations of 172 adult roe deer and step selection functions to infer landscape connectivity. In particular, we assessed the influence of six habitat features on fine scale movements across four biological seasons and three daily periods, based on variations in the risk-resource trade-off.ResultsThe influence of habitat features on roe deer movements was strongly dependent on proximity to refuge habitat, i.e. woodlands. Roe deer confined their movements to safe habitats during daytime and during the hunting season, when human activity is high. However, they exploited exposed open habitats more freely during night-time. Consequently, we observed marked temporal shifts in landscape connectivity, which was highest at night in summer and lowest during daytime in autumn. In particular, the onset of the autumn hunting season induced an abrupt decrease in landscape connectivity.ConclusionsHuman disturbance had a strong impact on roe deer movements, generating pronounced spatio-temporal variation in landscape connectivity. However, high connectivity at night across all seasons implies that Europe’s most abundant and widespread large herbivore potentially plays a key role in transporting ticks, seeds and nutrients among habitats.

Common drivers of seasonal movements on the migration – residency behavior continuum in a large herbivore

This study aimed to (1) identify the scale of environmental drivers of seasonal movements on the migration – residency behavior continuum in a large herbivore species and to (2) test the hypothesis that the same environmental drivers and spatio-temporal scaling should influence spatial processes in both migrants (long distance migration) and residents (short distance range shifts). We performed a comparative analysis of the influence of plant phenology and snow cover duration on seasonal movements of five partially migrating red deer populations with contrasting environmental conditions, at the seasonal range scale and at the study area scale. The five populations presented varying proportions of migrants, large gradients of migration distances and seasonal range shifts. The probability for a red deer to migrate was strongly influenced by large-scale environmental conditions, consistent with the resource heterogeneity hypothesis (high spatio-temporal scaling favors migration). Distances moved by both migrants and residents were strongly related to large-scale environmental conditions as well. We showed that similar proximal causes influenced these seasonal movements, reinforcing the idea of a continuum from migration to residency in response to seasonal environmental changes. Together, our findings suggest that global warming, by homogenizing large-scale environmental conditions, may thus decrease migratory tactics.