Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Truly sedentary? The multi-range tactic as a response to resource heterogeneity and unpredictability in a large herbivore.

Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multi-range tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment.

Country, cover or protection: what shapes the distribution of red deer and roe deer in the Bohemian Forest Ecosystem?

The Bohemian Forest Ecosystem encompasses various wildlife management systems. Two large, contiguous national parks (one in Germany and one in the Czech Republic) form the centre of the area, are surrounded by private hunting grounds, and hunting regulations in each country differ. Here we aimed at unravelling the influence of management-related and environmental factors on the distribution of red deer (Cervus elaphus) and roe deer (Capreolus capreolus) in this ecosystem. We used the standing crop method based on counts of pellet groups, with point counts every 100 m along 218 randomly distributed transects. Our analysis, which accounted for overdispersion as well as zero inflation and spatial autocorrelation, corroborated the view that both human management and the physical and biological environment drive ungulate distribution in mountainous areas in Central Europe. In contrast to our expectations, protection by national parks was the least important variable for red deer and the third important out of four variables for roe deer; protection negatively influenced roe deer distribution in both parks and positively influenced red deer distribution in Germany. Country was the most influential variable for both red and roe deer, with higher counts of pellet groups in the Czech Republic than in Germany. Elevation, which indicates increasing environmental harshness, was the second most important variable for both species. Forest cover was the least important variable for roe deer and the third important variable for red deer; the relationship for roe deer was positive and linear, and optimal forest cover for red deer was about 70% within a 500 m radius. Our results have direct implications for the future conservation management of deer in protected areas in Central Europe and show in particular that large non-intervention zones may not cause agglomerations of deer that could lead to conflicts along the border of protected, mountainous areas.

A one night stand? Reproductive excursions of female roe deer as a breeding dispersal tactic.

Breeding dispersal, defined as the net movement between successive breeding sites, remains a poorly understood and seldom reported phenomenon in mammals, despite its importance for population dynamics and genetics. In large herbivores, females may be more mobile during the breeding season, undertaking short-term trips (excursions) outside their normal home range. If fertilisation occurs, leading to gene flow of the male genome, this behaviour could be considered a form of breeding dispersal from a genetic point of view. Here, we investigated ranging behaviour of 235 adult roe deer using intensive GPS monitoring in six populations across Europe within the EURODEER initiative. We show that excursions are common from June to August among females, with 41.8% (vs. 18.1% of males) making at least one excursion. Most individuals performed only one excursion per season and departure dates for females were concentrated in time, centred on the rutting period, suggesting a link with reproduction. The distance females travelled during excursions was significantly greater than the site-specific average diameter of a male home range, while travel speed decreased once they progressed beyond this diameter, indicating search behaviour or interaction with other male(s) outside the resident male's territory. Because adults are normally highly sedentary, the potential for mating with relatives is substantial; hence, we conclude that rut excursions could be an alternative tactic enabling females to avoid mating with a closely related male. To understand better the ultimate drivers at play, it will be crucial to explore the genetic causes and consequences of this behaviour.

Seasonality, weather and climate affect home range size in roe deer across a wide latitudinal gradient within Europe.

1. Because many large mammal species have wide geographical ranges, spatially distant populations may be confronted with different sets of environmental conditions. Investigating how home range (HR) size varies across environmental gradients should yield a better understanding of the factors affecting large mammal ecology. 2. We evaluated how HR size of a large herbivore, the roe deer (Capreolus capreolus), varies in relation to seasonality, latitude (climate), weather, plant productivity and landscape features across its geographical range in Western Europe. As roe deer are income breeders, expected to adjust HR size continuously to temporal variation in food resources and energetic requirements, our baseline prediction was for HR size to decrease with proxies of resource availability. 3. We used GPS locations of roe deer collected from seven study sites (EURODEER collaborative project) to estimate fixed-kernel HR size at weekly and monthly temporal scales. We performed an unusually comprehensive analysis of variation in HR size among and within populations over time across the geographical range of a single species using generalized additive mixed models and linear mixed models, respectively. 4. Among populations, HR size decreased with increasing values for proxies of forage abundance, but increased with increases in seasonality, stochastic variation of temperature, latitude and snow cover. Within populations, roe deer HR size varied over time in relation to seasonality and proxies of forage abundance in a consistent way across the seven populations. Thus, our findings were broadly consistent across the distributional range of this species, demonstrating a strong and ubiquitous link between the amplitude and timing of environmental seasonality and HR size at the continental scale. 5. Overall, the variability in average HR size of roe deer across Europe reflects the interaction among local weather, climate and seasonality, providing valuable insight into the limiting factors affecting this large herbivore under contrasting conditions. The complexity of the relationships suggests that predicting ranging behaviour of large herbivores in relation to current and future climate change will require detailed knowledge not only about predicted increases in temperature, but also how this interacts with factors such as day length and climate predictability.

Estimation of the probability of fighting in fallow deer (Dama dama) during the rut.

Fighting between males is a frequent component of the rutting behavior of Cervidae. Frequent conflicts are exhausting; fighting may be risky and can lead to serious injuries or even death. We focused on the process of assessment of the opponent's fighting ability and escalation of the combat, estimating the probability of fighting based on the encounter components such as groaning and parallel walk. In this study, we observed the agonistic behavior of fallow deer bucks (Dama dama) during the rut over four seasons. During this time, we recorded 205 encounters between bucks. Non-contact display, which allows contestants to assess their opponents fighting ability, occurred in 83% of the encounters. The highest predicted probability of a fight was found when both of the males vocalized and turned into the parallel walk. The chance of a clear outcome decreased when the males were fighting in comparison to when they did not fight. The initiator of the competitive encounter won 41% of the cases, while the attacked buck won 23% of the encounters. If the contestants avoided fighting, however, the initiator won 78% of encounters. Therefore, the initiator was more successful when no fight occurred compared to when the encounters escalated into fighting. In most cases where ritualized behavior occurred, one of the opponents left after vocalization or parallel walk occurred. Thus, vocalization and parallel walk increased the probability for a clear outcome. The probability of a fight was lowest in situations where the males displayed asymmetric behavior. Increased symmetry of the contestants' behavior was strongly correlated with a higher probability of a fight. Thus, these results indicate that fallow deer bucks use efficient tactic during the rut, which, in turn, minimizes the chance of injury while fighting during the breeding season.