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Funding: This article is a contribution of the COVID-19 Bio-Logging Initiative, which is funded in part by the Gordon and Betty Moore Foundation (GBMF9881) and the National Geographic Society (NGS-82515R-20) (both grants to C.R.). COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals’ 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide. Postprint

Competitively dominant carnivore species can limit the population sizes and alter the behavior of inferior competitors. Established mechanisms that enable carnivore coexistence include spatial and temporal avoidance of dominant predator species by subordinates, and dietary niche separation. However, spatial heterogeneity across landscapes could provide inferior competitors with refuges in the form of areas with lower competitor density and/or locations that provide concealment from competitors. Here, we combine temporally overlapping telemetry data from dominant lions (Panthera leo) and subordinate African wild dogs (Lycaon pictus) with high-resolution remote sensing in an integrated step selection analysis to investigate how fine-scaled landscape heterogeneity might facilitate carnivore coexistence in South Africa?s Hluhluwe-iMfolozi Park, where both predators occur at exceptionally high densities. We ask whether the primary lion-avoidance strategy of wild dogs is spatial avoidance of lions or areas frequented by lions, or if wild dogs selectively use landscape features to avoid detection by lions. Within this framework, we also test whether wild dogs rely on proactive or reactive responses to lion risk. In contrast to previous studies finding strong spatial avoidance of lions by wild dogs, we found that the primary wild dog lion-avoidance strategy was to select landscape features that aid in avoidance of lion detection. This habitat selection was routinely used by wild dogs, and especially when in areas and during times of high lion-encounter risk, suggesting a proactive response to lion risk. Our findings suggest that spatial landscape heterogeneity could represent an alternative mechanism for carnivore coexistence, especially as ever-shrinking carnivore ranges force inferior competitors into increased contact with dominant species.

Aim: The recent recovery of large carnivores in Europe has been explained as resulting from a decrease in human persecution driven by widespread rural land abandonment, paralleled by forest cover increase and the consequent increase in availability of shelter and prey. We investigated whether land cover and human population density changes are related to the relative probability of occurrence of three European large carnivores: the grey wolf (Canis lupus), the Eurasian lynx (Lynx lynx) and the brown bear (Ursus arctos).Location: Europe, west of 64 degrees longitude.Methods: We fitted multi-temporal species distribution models using >50,000 occurrence points with time series of land cover, landscape configuration, protected areas, hunting regulations and human population density covering a 24-year period (1992-2015). Within the temporal window considered, we then predicted changes in habitat suitability for large carnivores throughout Europe.Results: Between 1992 and 2015, the habitat suitability for the three species increased in Eastern Europe, the Balkans, North-West Iberian Peninsula and Northern Scandinavia, but showed mixed trends in Western and Southern Europe. These trends were primarily associated with increases in forest cover and decreases in human population density, and, additionally, with decreases in the cover of mosaics of cropland and natural vegetation.Main conclusions: Recent land cover and human population changes appear to have altered the habitat suitability pattern for large carnivores in Europe, whereas protection level did not play a role. While projected changes largely match the observed recovery of large carnivore populations, we found mismatches with the recent expansion of wolves in Central and Southern Europe, where factors not included in our models may have played a dominant role. This suggests that large carnivores' co-existence with humans in European landscapes is not limited by habitat availability, but other factors such as favourable human tolerance and policy.

Additional file 1. Chapter 1: Study area summary statistics. Chapter 2: MigrateR parameters. Chapter 3: Generation of random steps and use of step length as a variable in the model. Chapter 4: Sensitivity analysis for number of available steps. Chapter 5: Definition of the variable ���Time of day���. Chapter 6: Collinearity of model predictors. Chapter 7: Model coefficients for steps at 3- and 5-hour fix intervals. Chapter 8: R code used to fit random slopes models.

We present methodological advances to a recently developed framework to study sequential habitat use by animals using a visually-explicit and tree-based Sequence Analysis Method (SAM), derived from molecular biology and more recently used in time geography. Habitat use sequences are expressed as annotations obtained by intersecting GPS movement trajectories with environmental layers. Here, we develop IM-SAM, where we use the individual reference area of use as the reference spatial context. To assess IM-SAM’s applicability, we investigated the sequential use of open and closed habitats across multiple European roe deer populations ranging in landscapes with contrasting structure. Starting from simulated sequences based on a mechanistic movement model, we found that different sequential patterns of habitat use were distinguished as separate, robust clusters, with less variable cluster size when habitats were present in equal proportions within the individual reference area of use. Application on real roe deer sequences showed that our approach effectively captured variation in spatio-temporal patterns of sequential habitat use, and provided evidence for important behavioral processes, such as day-night habitat alternation. By characterizing sequential habitat use patterns of animals, we may better evaluate the temporal trade-offs in animal habitat use and how they are affected by changes in landscapes. Sequence dissimilarity; dendrogram; ungulates; spatio-temporal habitat use; mechanistic movement model