High latitude northern bats (Eptesicus nilssonii) reveal adaptations to both high and low ambient temperatures.

Insectivorous bats at northern latitudes need to cope with long periods of no food for large parts of the year. Hence, bats which are resident at northern latitudes throughout the year will need to undergo a long hibernation season and a short reproductive season where foraging time is limited by extended daylight periods. Eptesicus nilssonii is the northernmost occurring bat species worldwide and hibernates locally when ambient temperatures (Ta) limit prey availability. Therefore, we investigated the energy spent maintaining normothermy at different Ta, as well as how much bats limit energy expenditure while in torpor. We found that, despite being exposed to Ta as low as 1.1°C, bats did not increase torpid metabolic rate, thus indicating that E. nilssonii can survive and hibernate at low ambient temperatures. Furthermore, we found a lower critical temperature (Tlc) of 27.8°C, which is lower than in most other vespertilionid bats, and we found no indication of any metabolic response to Ta up to 37.1°C. Interestingly, carbon dioxide production increased with increasing Ta above the Tlc, presumably caused by a release of retained CO2 in bats that remained in torpor for longer and aroused at Ta above the Tlc. Our results indicate that E. nilssonii can thermoconform at near-freezing Ta, and hence maintain longer torpor bouts with limited energy expenditure, yet also cope with high Ta when sun exposed in roosts during long summer days. These physiological traits are likely to enable the species to cope with ongoing and predicted climate change.

A species-level trait dataset of bats in Europe and beyond.

Knowledge of species' functional traits is essential for understanding biodiversity patterns, predicting the impacts of global environmental changes, and assessing the efficiency of conservation measures. Bats are major components of mammalian diversity and occupy a variety of ecological niches and geographic distributions. However, an extensive compilation of their functional traits and ecological attributes is still missing. Here we present EuroBaTrait 1.0, the most comprehensive and up-to-date trait dataset covering 47 European bat species. The dataset includes data on 118 traits including genetic composition, physiology, morphology, acoustic signature, climatic associations, foraging habitat, roost type, diet, spatial behaviour, life history, pathogens, phenology, and distribution. We compiled the bat trait data obtained from three main sources: (i) a systematic literature and dataset search, (ii) unpublished data from European bat experts, and (iii) observations from large-scale monitoring programs. EuroBaTrait is designed to provide an important data source for comparative and trait-based analyses at the species or community level. The dataset also exposes knowledge gaps in species, geographic and trait coverage, highlighting priorities for future data collection.

State dependence of arousal from torpor in brown long-eared bats (Plecotus auritus).

To cope with periods of low food availability and unsuitable environmental conditions (e.g., short photoperiod or challenging weather), many heterothermic mammals can readily go into torpor to save energy. However, torpor also entails several potential costs, and quantitative energetics can, therefore, be influenced by the individual state, such as available energy reserves. We studied the thermal energetics of brown long-eared bats (Plecotus auritus) in the northern part of its distributional range, including torpor entry, thermoregulatory ability during torpor and how they responded metabolically to an increasing ambient temperature (Ta) during arousal from torpor. Torpor entry occurred later in bats with higher body mass (Mb). During torpor, only 10 out of 21 bats increased oxygen consumption (V̇O2) to a greater extent above the mean torpor metabolic rates (TMR) when exposed to low Ta. The slope of the torpid thermoregulatory curve was shallower than that of resting metabolic rate (RMR) during normothermic conditions, indicating a higher thermal insulation during torpor. During exposure to an increasing Ta, all bats increased metabolic rate exponentially, but the bats with higher Mb aroused at a lower Ta than those with lower Mb. In bats with low Mb, arousal was postponed to an Ta above the lower critical temperature of the thermoneutral zone. Our results demonstrate that physiological traits, which are often considered fixed, can be more flexible than previously assumed and vary with individual state. Thus, future studies of thermal physiology should to a greater extent take individual state-dependent effects into account.

Drivers and consequences of partial migration in an alpine bird species.

Partial migration, where a portion of the population migrates between winter and summer (breeding) areas and the rest remain year-round resident, is a common phenomenon across several taxonomic groups. Several hypotheses have been put forward to explain why some individuals migrate while others stay resident, as well as the fitness consequences of the different strategies. Yet, the drivers and consequences of the decision to migrate or not are poorly understood.We used data from radio-tagged female (n = 73) willow ptarmigan Lagopus lagopus in an alpine study area in Central Norway to test if (i) the decision to migrate was dependent on individual state variables (age and body weight), (ii) individuals repeated migratory decisions between seasons, and (iii) the choice of migratory strategy was related to reproductive success.Partially supporting our prediction that migratory strategy depends on individual state, we found that juvenile birds with small body sizes were more likely to migrate, whereas large juveniles remained resident. For adult females, we found no relationship between the decision to migrate or stay resident and body weight. We found evidence for high individual repeatability of migratory decision between seasons. Migratory strategy did not explain variation in clutch size or nest fate among individuals, suggesting no direct influence of the chosen strategy on reproductive success.Our results indicate that partial migration in willow ptarmigan is related to juvenile body weight, and that migratory behavior becomes a part of the individual life history as a fixed strategy. Nesting success was not affected by migratory strategy in our study population, but future studies should assess other traits to further test potential fitness consequences.

Ontogenetic niche shifts in a locally endangered tree species (Olea europaea subsp. cuspidata) in a disturbed forest in Northern Ethiopia: Implications for conservation.

Understanding the responses of different ontogenetic stages to environmental and human disturbance factors is essential for developing efficient conservation strategies for endangered plant species. We examined how three ontogenetic stages of a locally endangered tree species, Olea europaea subsp. cuspidata, responded to environmental factors and human disturbance in Hugumburda dry Afromontane forest in Ethiopia. We counted individual seedlings, saplings and adults of O. europaea in 70 20 × 20 m quadrats over ca. 2.8 ha, and measured biotic (woody species richness, canopy cover, aboveground tree biomass, herbaceous cover), abiotic (soil and topographic variables), and human disturbance factors (logging and tracks). To detect ontogenetic niche shifts, we compared observed vs. simulated locations of trees in the three life stages and how they related to the environmental and human disturbance factors. We found that the population structure of O. europaea showed generally low recruitment, with few seedlings per hectare compared with the abundance of saplings and adults. The probability of finding O. europaea individuals was influenced by biotic (woody species richness) and abiotic (soil depth, slope) environmental conditions and human disturbance (logging intensity), but the direction, strength and shape of the relationships differed between seedling, sapling and adult life stages, indicating ontogenetic niche shifts. All life stages showed a positive relationship with elevation. The observed environmental niches of the different lifestages of O. europaea, and their association with human disturbance levels, should be considered when conservation strategies are developed for this species. Human disturbance in terms of logging decreases the abundance of saplings, but may facilitate emerging seedlings through creation of gaps with improved light conditions. Recruitment is, however, very low in the study area, and seedlings should be protected from browsing to enhance survival. Woody species richness in general should be conserved to optimize conditions also for O. europaea saplings.

Enhancing pollination is more effective than increased conventional agriculture inputs for improving watermelon yields.

Agricultural practices to improve yields in small-scale farms in Africa usually focus on improving growing conditions for the crops by applying fertilizers, irrigation, and/or pesticides. This may, however, have limited effect on yield if the availability of effective pollinators is too low. In this study, we established an experiment to test whether soil fertility, soil moisture, and/or pollination was limiting watermelon (Citrullus lanatus) yields in Northern Tanzania. We subjected the experimental field to common farming practices while we treated selected plants with extrafertilizer applications, increased irrigation and/or extra pollination in a three-way factorial experiment. One week before harvest, we assessed yield from each plant, quantified as the number of mature fruits and their weights. We also assessed fruit shape since this may affect the market price. For the first fruit ripening on each plant, we also assessed sugar content (brix) and flesh color as measures of fruit quality for human consumption. Extra pollination significantly increased the probability of a plant producing a second fruit of a size the farmer could sell at the market, and also the fruit sugar content, whereas additional fertilizer applications or increased irrigation did not improve yields. In addition, we did not find significant effects of increased fertilizer or watering on fruit sugar, weight, or color. We concluded that, insufficient pollination is limiting watermelon yields in our experiment and we suggest that this may be a common situation in sub-Saharan Africa. It is therefore critically important that small-scale farmers understand the role of pollinators and understand their importance for agricultural production. Agricultural policies to improve yields in developing countries should therefore also include measures to improve pollination services by giving education and advisory services to farmers on how to develop pollinator-friendly habitats in agricultural landscapes.

High frequency GPS bursts and path-level analysis reveal linear feature tracking by red foxes.

There is a need to quantify and better understand how wildlife interact with linear features, as these are integral elements of most landscapes. One potentially important aspect is linear feature tracking (LFT), yet studies rarely succeed in directly revealing or quantifying this behavior. In a proof-of-concept study, we employed short-term intensive GPS monitoring of red foxes (Vulpes vulpes) in a multiple-use landscape in southern Norway. Using periodic bursts of high frequency GPS position fixes, we performed modified path selection analyses to estimate the propensity of foxes to track natural and man-made linear features (roads, forest edges, and streams) once they are encountered. Foxes in our study tracked primarily forest edges and roads. Forty-three percent of bursts that encountered any linear feature resulted in LFT. LFT, although prominent, was manifested as a short-lived behavior, with overall median times to linear feature abandonment around two minutes. Movement speeds were highest along roads, perhaps due to greater ease of travel or higher perceived risk. In the highly heterogeneous habitats that characterize human-dominated landscapes, LFT may be manifested at such a fine spatio-temporal scale that it would remain hidden during telemetry studies employing conventional position fix frequencies. The approach described here may aid others studying spatial behaviors that are manifested over very short durations, yet are biologically significant.

Univariate Community Assembly Analysis (UniCAA): Combining hierarchical models with null models to test the influence of spatially restricted dispersal, environmental filtering, and stochasticity on community assembly.

Identifying the influence of stochastic processes and of deterministic processes, such as dispersal of individuals of different species and trait-based environmental filtering, has long been a challenge in studies of community assembly. Here, we present the Univariate Community Assembly Analysis (UniCAA) and test its ability to address three hypotheses: species occurrences within communities are (a) limited by spatially restricted dispersal; (b) environmentally filtered; or (c) the outcome of stochasticity-so that as community size decreases-species that are common outside a local community have a disproportionately higher probability of occurrence than rare species. The comparison with a null model allows assessing if the influence of each of the three processes differs from what one would expect under a purely stochastic distribution of species. We tested the framework by simulating "empirical" metacommunities under 15 scenarios that differed with respect to the strengths of spatially restricted dispersal (restricted vs. not restricted); habitat isolation (low, intermediate, and high immigration rates); and environmental filtering (strong, intermediate, and no filtering). Through these tests, we found that UniCAA rarely produced false positives for the influence of the three processes, yielding a type-I error rate ≤5%. The type-II error rate, that is, production of false negatives, was also acceptable and within the typical cutoff (20%). We demonstrate that the UniCAA provides a flexible framework for retrieving the processes behind community assembly and propose avenues for future developments of the framework.

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion.

Vast areas of the African savanna landscapes are characterized by tree-covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two-month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.

The effects of habitat management on the species, phylogenetic and functional diversity of bees are modified by the environmental context.

Anthropogenic landscape elements, such as roadsides, hedgerows, field edges, and power line clearings, can be managed to provide important habitats for wild bees. However, the effects of habitat improvement schemes in power line clearings on components of diversity are poorly studied. We conducted a large-scale experiment to test the effects of different management practices on the species, phylogenetic, and functional diversity of wild bees in power line clearings (n = 19 sites across southeastern Norway) and explored whether any treatment effects were modified by the environmental context. At each site, we conducted the following treatments: (1) Cut: all trees cut and left to decay in the clearing; (2) Cut + Remove: all trees cut and removed from the plot; and (3) Uncut: uncleared. The site-specific environmental context (i.e., elevation and floral diversity) influenced the species, phylogenetic, and functional diversity within bee species assemblages. The largest number of species was found in the Cut + Remove treatment in plots with a high forb species richness, indicating that the outcome of management practices depends on the environmental context. Clearing of treatment plots with many forb species also appeared to alter the phylogenetic composition of bee species assemblages, that is, more closely related species were found in the Cut and the Cut + Remove plots than in the Uncut plots. Synthesis and applications: Our experimental simulation of management practices in power line clearings influenced the species, phylogenetic, and functional diversity of bee species assemblages. Frequent clearing and removal of the woody debris at low elevations with a high forb species richness can increase the value of power line clearings for solitary bees. It is therefore important for managers to consider the environmental context when designing habitat improvement schemes for solitary bees.

Inter-assemblage facilitation: the functional diversity of cavity-producing beetles drives the size diversity of cavity-nesting bees.

Inter-specific interactions are important drivers and maintainers of biodiversity. Compared to trophic and competitive interactions, the role of non-trophic facilitation among species has received less attention. Cavity-nesting bees nest in old beetle borings in dead wood, with restricted diameters corresponding to the body size of the bee species. The aim of this study was to test the hypothesis that the functional diversity of cavity-producing wood boring beetles - in terms of cavity diameters - drives the size diversity of cavity-nesting bees. The invertebrate communities were sampled in 30 sites, located in forested landscapes along an elevational gradient. We regressed the species richness and abundance of cavity nesting bees against the species richness and abundance of wood boring beetles, non-wood boring beetles and elevation. The proportion of cavity nesting bees in bee species assemblage was regressed against the species richness and abundance of wood boring beetles. We also tested the relationships between the size diversity of cavity nesting bees and wood boring beetles. The species richness and abundance of cavity nesting bees increased with the species richness and abundance of wood boring beetles. No such relationship was found for non-wood boring beetles. The abundance of wood boring beetles was also related to an increased proportion of cavity nesting bee individuals. Moreover, the size diversity of cavity-nesting bees increased with the functional diversity of wood boring beetles. Specifically, the mean and dispersion of bee body sizes increased with the functional dispersion of large wood boring beetles. The positive relationships between cavity producing bees and cavity nesting bees suggest that non-trophic facilitative interactions between species assemblages play important roles in organizing bee species assemblages. Considering a community-wide approach may therefore be required if we are to successfully understand and conserve wild bee species assemblages in forested landscapes.

Female offspring desertion and male-only care increase with natural and experimental increase in food abundance.

In species with biparental care, one parent may escape the costs of parental care by deserting and leaving the partner to care for the offspring alone. A number of theoretical papers have suggested a link between uniparental offspring desertion and ecological factors, but empirical evidence is scarce. We investigated the relationship between uniparental desertion and food abundance in a natural population of Tengmalm's owl Aegolius funereus, both by means of a 5-year observational study and a 1-year experimental study. Parents and offspring were fitted with radio-transmitters in order to reveal the parental care strategy (i.e. care or desert) of individual parents, and to keep track of the broods post-fledging. We found that 70 per cent of the females from non-experimental nests deserted, while their partner continued to care for their joint offspring alone. Desertion rate was positively related to natural prey population densities and body reserves of the male partner. In response to food supplementation, a larger proportion of the females deserted, and females deserted the offspring at an earlier age. Offspring survival during the post-fledging period tended to be lower in deserted than in non-deserted broods. We argue that the most important benefit of deserting may be remating (sequential polyandry).