Considering habitat conversion and fragmentation in characterisation factors for land-use impacts on vertebrate species richness.

Human land use is one of the primary threats to terrestrial species richness and is considered a priority for meeting global sustainability and biodiversity targets. Decision-support tools, such as life cycle assessment (LCA), are widely used for developing strategies to achieve such objectives. Currently available life cycle impact assessment (LCIA) methods apply the countryside species-area relationship (c-SAR) to quantify habitat conversion impacts on species richness. However, additional effects of habitat fragmentation are yet ignored in these assessments. We use the species-habitat relationship (SHR), an adaptation of the c-SAR that considers both habitat conversion and fragmentation effects, to develop a new set of land-use characterisation factors for 702 terrestrial ecoregions (in 238 countries), four land-use types (urban, cropland, pasture, and forestry), and four taxonomic groups (amphibians, birds, mammals, and reptiles; plus the aggregate of these vertebrate groups). The SHR generally predicts higher per-area impacts of land-use than the impacts estimated by the c-SAR (a median relative difference of +9%), indicating that land-use impacts may be systematically underestimated when ignoring fragmentation effects. Whereas per-area impacts of land-use on regional species richness are highest in temperate regions, reflecting the diminished extent of natural habitat, per-area impacts of land-use on global species richness are highest in the subtropics, reflecting the importance of tropical regions and islands to global vertebrate species diversity. The large variety in magnitude of land-use impacts across the world's regions emphasizes the importance of regionalised assessments. The set of characterisation factors proposed here can be readily used in environmental decision-making.

Paint it black: Efficacy of increased wind turbine rotor blade visibility to reduce avian fatalities.

As wind energy deployment increases and larger wind-power plants are considered, bird fatalities through collision with moving turbine rotor blades are expected to increase. However, few (cost-) effective deterrent or mitigation measures have so far been developed to reduce the risk of collision. Provision of "passive" visual cues may enhance the visibility of the rotor blades enabling birds to take evasive action in due time. Laboratory experiments have indicated that painting one of three rotor blades black minimizes motion smear (Hodos 2003, Minimization of motion smear: Reducing avian collisions with wind turbines). We tested the hypothesis that painting would increase the visibility of the blades, and that this would reduce fatality rates in situ, at the Smøla wind-power plant in Norway, using a Before-After-Control-Impact approach employing fatality searches. The annual fatality rate was significantly reduced at the turbines with a painted blade by over 70%, relative to the neighboring control (i.e., unpainted) turbines. The treatment had the largest effect on reduction of raptor fatalities; no white-tailed eagle carcasses were recorded after painting. Applying contrast painting to the rotor blades significantly reduced the collision risk for a range of birds. Painting the rotor blades at operational turbines was, however, resource demanding given that they had to be painted while in-place. However, if implemented before construction, this cost will be minimized. It is recommended to repeat this experiment at other sites to ensure that the outcomes are generic at various settings.

A dead giveaway: Foraging vultures and other avian scavengers respond to auditory cues.

Carrion represents an unpredictable and widely distributed primary food source for vultures and other avian scavengers. Avian scavengers in African savanna ecosystems are reported to rely exclusively on visual stimuli to locate carcasses. However, carnivores' predation of large mammalian herbivores and subsequent competition for access to the carcass can result in considerable noise, often audible over long distances and for prolonged periods. Vultures and other avian scavengers may therefore detect and respond to these auditory cues, as do the mammalian carnivores alongside which vultures have coevolved, but this has not been investigated to date. Working in the Serengeti ecosystem, Tanzania, we used diurnal auditory broadcasts to simulate predation and competitive carnivore feeding interactions. Based on the current understanding of avian scavenger ecology, we hypothesized that avian responses to call-in stations would be evoked exclusively by visual, rather than auditory, cues. We therefore predicted that (a) the arrival of avian scavengers at call-in stations should be preceded and facilitated by mammalian carnivores and that (b) the arrival of avian scavengers would be positively correlated with the number of mammalian scavengers present, which would increase detectability. We recorded 482 birds during 122 separate playback events. In 22% of these instances, avian scavengers arrived first, ruling out responses based exclusively on visual observations of mammalian carnivores, thereby contradicting our first prediction. Furthermore, the first avian arrivals at survey sessions were inversely related to the number of hyenas and jackals present, contradicting our second prediction. Since no bait or carcasses were used during the experiments, these responses are indicative of the birds' ability to detect and respond to audio stimuli. Our findings challenge the current consensus of sensory perception and foraging in these species and provide evidence that avian scavengers have the ability to use sound to locate food resources.

Effect of tower base painting on willow ptarmigan collision rates with wind turbines.

Birds colliding with turbine rotor blades is a well-known negative consequence of wind-power plants. However, there has been far less attention to the risk of birds colliding with the turbine towers, and how to mitigate this risk.Based on data from the Smøla wind-power plant in Central Norway, it seems highly likely that willow ptarmigan (the only gallinaceous species found on the island) is prone to collide with turbine towers. By employing a BACI-approach, we tested if painting the lower parts of turbine towers black would reduce the collision risk.Overall, there was a 48% reduction in the number of recorded ptarmigan carcasses per search at painted turbines relative to neighboring control (unpainted) ones, with significant variation both within and between years.Using contrast painting to the turbine towers resulted in significantly reduced number of ptarmigan carcasses found, emphasizing the effectiveness of such a relatively simple mitigation measure.

High-Resolution Modeling of Uplift Landscapes can Inform Micrositing of Wind Turbines for Soaring Raptors.

Collision risk of soaring birds is partly associated with updrafts to which they are attracted. To identify the risk-enhancing landscape features, a micrositing tool was developed to model orographic and thermal updraft velocities from high-resolution remote sensing data. The tool was applied to the island of Hitra, and validated using GPS-tracked white-tailed eagles (Haliaeetus albicilla). Resource selection functions predicted that eagles preferred ridges with high orographic uplift, especially at flight altitudes within the rotor-swept zone (40-110 m). Flight activity was negatively associated with the widely distributed areas with high thermal uplift at lower flight altitudes (<110 m). Both the existing wind-power plant and planned extension are placed at locations rendering maximum orographic updraft velocities around the minimum sink rate for white-tailed eagles (0.75 m/s) but slightly higher thermal updraft velocities. The tool can contribute to improve micrositing of wind turbines to reduce the environmental impacts, especially for soaring raptors.

Resource utilization by the Kori bustard in the Serengeti ecosystem.

This study aimed to understand the movement behaviour and utilization distributions of Kori bustards in space and time in the Serengeti ecosystem. A total of 14 individuals were tracked with the aid of GPS (Geographical positioning system) satellite transmitters, and their sexes were identified using DNA analysis. A species utilization distribution was estimated using the Brownian bridge movement model (hereafter dBBMM) in which the probability of being in an area is conditioned by starting and ending (GPS) relocations. Resource selections were analysed by comparing the GPS relocations with locations randomly placed within each individual's region of utilization in a spatio-temporal approach. Vegetation information was derived from a Serengeti GIS vegetation map and Data Centre and was reclassified as Open grassland, Dense grassland, Shrubbed grassland, Treed grassland, Shrubland, and Woodland. The Shannon diversity index for vegetation was calculated based on the original vegetation classification. Used versus non-used habitats were contrasted using a generalized linear mixed-effects model with a binomial distribution. The results indicated that males were 21.5% more mobile than females, and movements were 6.3% more diffuse during the non-breeding period compared to the breeding period (7.59 versus 7.14, respectively). Contrasting models indicated that males preferred more open grasslands during the non-breeding period and also preferred closed and shrubbed grassland during the breeding period. Females preferred more woody vegetation during the non-breeding season compared to the breeding season. The most parsimonious model indicated that females preferred to stay closer to rivers and diverse areas during the non-breeding period whereas males preferred areas that were farther from rivers and homogenous. Homogeneous areas were preferred during the breeding period, and heterogeneous areas were preferred during the non-breeding period. We conclude that the movement behaviours of Kori bustards changes with the season and habitat. Further research is needed to understand the factors driving the seasonal movement of Kori bustards in the Serengeti ecosystem.

No evidence of handling-induced mortality in Serengeti's African wild dog population.

The disappearance of an endangered African wild dog population from Serengeti National Park (SNP) led to international debate centered around one question: were researchers to blame? The "Burrows' hypothesis" postulated that stress induced by research-related immobilization and handling reactivated a latent rabies virus, eliminating the population. Insufficient data inhibited hypothesis testing, but since wild dogs persisted alongside SNP and have been studied since 2005, the hypothesis can be tested 25 years after its proposition. To be supported, wild dog immobilization interventions should have resulted in high mortality rates. However, 87.6% of 121 handled wild dogs (2006-2016) survived >12 months post-handling. Some argued that viral reactivation would necessitate long-term stress. Following immobilization, 67 animals were captured, transported, and held in a translocation enclosure. Despite the longer-term stress, 95.5% survived >12 months. Furthermore, the stable number of wild dog packs in the ecosystem over the past decade, and lack of recolonization of SNP, strongly oppose Burrows' hypothesis. Instead, factors such as heightened levels of interspecific competition are likely to have contributed to the wild dog disappearance and subsequent avoidance of the Serengeti plains. Handling and radio telemetry are invaluable when studying elusive endangered species, yielding information pertinent to their conservation and management, and had no effect on Serengeti wild dog survival.

Erratum: Being stressed outside the park-conservation of African elephants (Loxodonta africana) in Namibia.

[This corrects the article DOI: 10.1093/conphys/cox067.][This corrects the article DOI: 10.1093/conphys/cox067.].

Modeling Net Land Occupation of Hydropower Reservoirs in Norway for Use in Life Cycle Assessment.

Increasing hydropower electricity production constitutes a unique opportunity to mitigate climate change impacts. However, hydropower electricity production also impacts aquatic and terrestrial biodiversity through freshwater habitat alteration, water quality degradation, and land use and land use change (LULUC). Today, no operational model exists that covers any of these cause-effect pathways within life cycle assessment (LCA). This paper contributes to the assessment of LULUC impacts of hydropower electricity production in Norway in LCA. We quantified the inundated land area associated with 107 hydropower reservoirs with remote sensing data and related it to yearly electricity production. Therewith, we calculated an average net land occupation of 0.027 m2·yr/kWh of Norwegian storage hydropower plants for the life cycle inventory. Further, we calculated an adjusted average land occupation of 0.007 m2·yr/kWh, accounting for an underestimation of water area in the performed maximum likelihood classification. The calculated land occupation values are the basis to support the development of methods for assessing the land occupation impacts of hydropower on biodiversity in LCA at a damage level.

Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black-legged kittiwake.

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long-term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick-rearing black-legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony.

Being stressed outside the park-conservation of African elephants (Loxodonta africana) in Namibia.

The conservation of the African savanna elephant (Loxodonta africana) is of prime importance for many African countries. Interactions between elephants and humans are known to induce stress and thereby have the potential to affect elephants' fitness. In Namibia, anthropogenic disturbances are increasing due to increasing human population size and development, particularly near protected areas, such as national parks. In this study, we investigated elephant stress levels in relation to their land use, specifically their protection status, comparing elephants within Etosha National Park in Namibia with elephants residing outside the park. We noninvasively collected dung samples of 91 elephants and determined the concentration of faecal glucocorticoid metabolites (fGCM), an indicator of physiological stress. Elephants outside the park (N = 35) had significantly higher concentrations of fGCM than those inside ENP (N = 56), suggesting that, despite including community-based conservancies, unprotected areas are more stressful for elephants than protected areas, most likely due to increased interactions with humans. We also found that males had lower fGCM concentrations than females, but no significant effect of age, body size or group size was detected. Additionally, herd sizes were significantly smaller and calf recruitment was potentially lower in unprotected areas. These findings underpin the importance of protected areas such as ENP, while encouraging decision-makers to continue reducing and mitigating potential human-induced disturbances.

Performance test and verification of an off-the-shelf automated avian radar tracking system.

Microwave radar is an important tool for observation of birds in flight and represents a tremendous increase in observation capability in terms of amount of surveillance space that can be covered at relatively low cost. Based on off-the-shelf radar hardware, automated radar tracking systems have been developed for monitoring avian movements. However, radar used as an observation instrument in biological research has its limitations that are important to be aware of when analyzing recorded radar data. This article describes a method for exploring the detection capabilities of a dedicated short-range avian radar system used inside the operational Smøla wind-power plant. The purpose of the testing described was to find the maximum detection range for various sized birds, while controlling for the effects of flight tortuosity, flight orientation relative to the radar and ground clutter. The method was to use a dedicated test target in form of a remotely controlled unmanned aerial vehicle (UAV) with calibrated radar cross section (RCS), which enabled the design of virtually any test flight pattern within the area of interest. The UAV had a detection probability of 0.5 within a range of 2,340 m from the radar. The detection performance obtained by the RCS-calibrated test target (-11 dBm2, 0.08 m2 RCS) was then extrapolated to find the corresponding performance of differently sized birds. Detection range depends on system sensitivity, the environment within which the radar is placed and the spatial distribution of birds. The avian radar under study enables continuous monitoring of bird activity within a maximum range up to 2 km dependent on the size of the birds in question. While small bird species may be detected up to 0.5-1 km, larger species may be detected up to 1.5-2 km distance from the radar.

Match between soaring modes of black kites and the fine-scale distribution of updrafts.

Understanding how soaring birds use updrafts at small spatial scales is important to identify ecological constraints of movement, and may help to prevent conflicts between wind-energy development and the conservation of wildlife. We combined high-frequency GPS animal tracking and fine-spatial-scale uplift modelling to establish a link between flight behaviour of soaring birds and the distribution of updrafts. We caught 21 black kites (Milvus migrans) and GPS-tracked them while flying over the Tarifa region, on the Spanish side of the Strait of Gibraltar. This region has a diverse topography and land cover, favouring a heterogeneous updraft spatial distribution. Bird tracks were segmented and classified into flight modes from motion parameters. Thermal and orographic uplift velocities were modelled from publically available remote-sensing and meteorological data. We found that birds perform circular soaring in areas of higher predicted thermal uplift and linear soaring in areas of higher predicted orographic uplift velocity. We show that updraft maps produced from publically available data can be used to predict where soaring birds will concentrate their flight paths and how they will behave in flight. We recommend the use of this methodological approach to improve environmental impact assessments of new wind-energy installations.

How many routes lead to migration? Comparison of methods to assess and characterize migratory movements.

Decreasing rate of migration in several species as a consequence of climate change and anthropic pressure, together with increasing evidence of space-use strategies intermediate between residency and complete migration, are very strong motivations to evaluate migration occurrence and features in animal populations. The main goal of this paper was to perform a relative comparison between methods for identifying and characterizing migration at the individual and population level on the basis of animal location data. We classified 104 yearly individual trajectories from five populations of three deer species as migratory or non-migratory, by means of three methods: seasonal home range overlap, spatio-temporal separation of seasonal clusters and the Net Squared Displacement (NSD) method. For migratory cases, we also measured timing and distance of migration and residence time on the summer range. Finally, we compared the classification in migration cases across methods and populations. All methods consistently identified migration at the population level, that is, they coherently distinguished between complete or almost complete migratory populations and partially migratory populations. However, in the latter case, methods coherently classified only about 50% of the single cases, that is they classified differently at the individual-animal level. We therefore infer that the comparison of methods may help point to 'less-stereotyped' cases in the residency-to-migration continuum. For cases consistently classified by all methods, no significant differences were found in migration distance, or residence time on summer ranges. Timing of migration estimated by NSD was earlier than by the other two methods, both for spring and autumn migrations. We suggest three steps to identify improper inferences from migration data and to enhance understanding of intermediate space-use strategies. We recommend (i) classifying migration behaviours using more than one method, (ii) performing sensitivity analysis on method parameters to identify the extent of the differences and (iii) investigating inconsistently classified cases as these may often be ecologically interesting (i.e. less-stereotyped migratory behaviours).

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves.

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.

Habitat differentiation within the large-carnivore community of Norway's multiple-use landscapes.

The re-establishment of large carnivores in Norway has led to increased conflicts and the adoption of regional zoning for these predators. When planning the future distribution of large carnivores, it is important to consider details of their potential habitat tolerances and strength of inter-specific differentiation. We studied differentiation in habitat and kill sites within the large-carnivore community of south-eastern Norway.We compared habitat selection of the brown bear Ursus arctos L., Eurasian lynx Lynx lynx L., wolf Canis lupus L. and wolverine Gulo gulo L., based on radio-tracking data. Differences in kill site locations were explored using locations of documented predator-killed sheep Ovis aries L. We modelled each species' selection for, and differentiation in, habitat and kill sites on a landscape scale using resource selection functions and multinomial logistic regression. Based on projected probability of occurrence maps, we estimated continuous patches of habitat within the study area.Although bears, lynx, wolves and wolverines had overlapping distributions, we found a clear differentiation for all four species in both habitat and kill sites. The presence of bears, wolves and lynx was generally associated with rugged, forested areas at lower elevations, whereas wolverines selected rugged terrain at higher elevations. Some degree of sympatry was possible in over 40% of the study area, although only 1.5% could hold all four large carnivores together.Synthesis and applications. A geographically differentiated management policy has been adopted in Norway, aimed at conserving viable populations of large carnivores while minimizing the potential for conflicts. Sympatry of all four carnivores will be most successful if regional zones are established of adequate size spanning an elevational gradient. High prey densities, low carnivore densities, low dietary overlap and scavenging opportunities have most probably led to reduced competitive exclusion. Although regional sympatry enhances the conservation of an intact guild of large carnivores, it may well increase conflict levels and resistance to carnivore conservation locally.