Showing papers in "Journal of Animal Ecology in 2017"

Low migratory connectivity is common in long-distance migrant birds.

Abstract: Summary Estimating how much long-distance migrant populations spread out and mix during the non-breeding season (migratory connectivity) is essential for understanding and predicting population dynamics in the face of global change. We quantify variation in population spread and inter-population mixing in long-distance, terrestrial migrant land-bird populations (712 individuals from 98 populations of 45 species, from tagging studies in the Neotropic and Afro-Palearctic flyways). We evaluate the Mantel test as a metric of migratory connectivity, and explore the extent to which variance in population spread can be explained simply by geography. The mean distance between two individuals from the same population during the non-breeding season was 743 km, covering 10–20% of the maximum width of Africa/South America. Individuals from different breeding populations tended to mix during the non-breeding season, although spatial segregation was maintained in species with relatively large non-breeding ranges (and, to a lesser extent, those with low population-level spread). A substantial amount of between-population variation in population spread was predicted simply by geography, with populations using non-breeding zones with limited land availability (e.g. Central America compared to South America) showing lower population spread. The high levels of population spread suggest that deterministic migration tactics are not generally adaptive; this makes sense in the context of the recent evolution of the systems, and the spatial and temporal unpredictability of non-breeding habitat. The conservation implications of generally low connectivity are that the loss (or protection) of any non-breeding site will have a diffuse but widespread effect on many breeding populations. Although low connectivity should engender population resilience to shifts in habitat (e.g. due to climate change), we suggest it may increase susceptibility to habitat loss. We hypothesize that, because a migrant species cannot adapt to both simultaneously, migrants generally may be more susceptible to population declines in the face of concurrent anthropogenic habitat and climate change.

The many faces of fear: a synthesis of the methodological variation in characterizing predation risk

Abstract: The NSF Graduate Research Fellowship Program (RJM), the Michigan State University MasterCard Foundation Scholars Program (ABM), CNPq-Brasil (LA), and the University of Montana Boone and Crockett Program (JJM).

Repeatability, heritability, and age-dependence of seasonal plasticity in aggressiveness in a wild passerine bird.

Abstract: 1.Labile characters allow individuals to flexibly adjust their phenotype to changes in environmental conditions. There is growing evidence that individuals can differ both in average expression of and level of plasticity in this type of character. Both of these aspects are studied in conjunction within a reaction norm framework. 2.Theoreticians have investigated the factors promoting variation in reaction norm intercepts (average phenotype) and slopes (level of plasticity) of a key labile character: behaviour. A general prediction from their work is that selection will favour the evolution of repeatable individual variation in level of plasticity only under certain ecological conditions. While factors promoting individual repeatability of plasticity have thus been identified, empirical estimates of this phenomenon are largely lacking for wild populations. 3.We assayed aggressiveness of individual male great tits (Parus major) twice during their egg-laying stage and twice during their egg-incubation stage to quantify each male's level of seasonal plasticity. This procedure was applied during six consecutive years; all males breeding in our plots during those years were assayed, resulting in repeated measures of individual reaction norms for any individual breeding in multiple years. We quantified among- and within-individual variation in reaction norm components, allowing us to estimate repeatability of seasonal plasticity. Using social pedigree information, we further partitioned reaction norm components into their additive genetic and permanent environmental counterparts. 4.Cross-year individual repeatability for the intercepts (average aggressiveness) and slopes (level of seasonal plasticity) of the aggressiveness reaction norms were 0.574 and 0.516 respectively. The posterior modes of the estimates suggested modest heritabilities (h2=0.260 for intercepts; h2=0.266 for slopes) that were relatively uncertain. Males behaved more aggressively in areas with higher breeding densities, and became less aggressive and less plastic with increasing age; plasticity thus varied within individuals and was multidimensional in nature. 5.This empirical study quantified cross-year individual repeatability, heritability, and age-related reversible plasticity in behaviour. Acknowledging such patterns of multi-level variation is important not only for testing behavioural ecology theory concerning the evolution of repeatable differences in behavioural plasticity but also for predicting how reversible plasticity may evolve in natural populations. This article is protected by copyright. All rights reserved.

Forbidden fruit: human settlement and abundant fruit create an ecological trap for an apex omnivore

Abstract: Summary Habitat choice is an evolutionary product of animals experiencing increased fitness when preferentially occupying high-quality habitat. However, an ecological trap (ET) can occur when an animal is presented with novel conditions and the animal's assessment of habitat quality is poorly matched to its resulting fitness. We tested for an ET for grizzly (brown) bears using demographic and movement data collected in an area with rich food resources and concentrated human settlement. We derived measures of habitat attractiveness from occurrence models of bear food resources and estimated demographic parameters using DNA mark–recapture information collected over 8 years (2006–2013). We then paired this information with grizzly bear mortality records to investigate kill and movement rates. Our results demonstrate that a valley high in both berry resources and human density was more attractive than surrounding areas, and bears occupying this region faced 17% lower apparent survival. Despite lower fitness, we detected a net flow of bears into the ET, which contributed to a study-wide population decline. This work highlights the presence and pervasiveness of an ET for an apex omnivore that lacks the evolutionary cues, under human-induced rapid ecological change, to assess trade-offs between food resources and human-caused mortality, which results in maladaptive habitat selection.

Temporal shifts and temperature sensitivity of avian spring migratory phenology: a phylogenetic meta-analysis

Abstract: There are wide reports of advances in the timing of spring migration of birds over time and in relation to rising temperatures, though phenological responses vary substantially within and among species. An understanding of the ecological, life-history and geographic variables that predict this intra- and interspecific variation can guide our projections of how populations and species are likely to respond to future climate change. Here, we conduct phylogenetic meta-analyses addressing slope estimates of the timing of avian spring migration regressed on (i) year and (ii) temperature, representing a total of 413 species across five continents. We take into account slope estimation error and examine phylogenetic, ecological and geographic predictors of intra- and interspecific variation. We confirm earlier findings that on average birds have significantly advanced their spring migration time by 2·1 days per decade and 1·2 days °C-1 . We find that over time and in response to warmer spring conditions, short-distance migrants have advanced spring migratory phenology by more than long-distance migrants. We also find that larger bodied species show greater advance over time compared to smaller bodied species. Our results did not reveal any evidence that interspecific variation in migration response is predictable on the basis of species' habitat or diet. We detected a substantial phylogenetic signal in migration time in response to both year and temperature, suggesting that some of the shifts in migratory phenological response to climate are predictable on the basis of phylogeny. However, we estimate high levels of species and spatial variance relative to phylogenetic variance, which is consistent with plasticity in response to climate evolving fairly rapidly and being more influenced by adaptation to current local climate than by common descent. On average, avian spring migration times have advanced over time and as spring has become warmer. While we are able to identify predictors that explain some of the true among-species variation in response, substantial intra- and interspecific variation in migratory response remains to be explained.

The effects of ant nests on soil fertility and plant performance: a meta-analysis

Abstract: Ants are recognized as one of the major sources of soil disturbance world-wide. However, this view is largely based on isolated studies and qualitative reviews. Here, for the first time, we quantitatively determined whether ant nests affect soil fertility and plant performance, and identified the possible sources of variation of these effects. Using Bayesian mixed-models meta-analysis, we tested the hypotheses that ant effects on soil fertility and plant performance depend on the substrate sampled, ant feeding type, latitude, habitat and the plant response variable measured. Ant nests showed higher nutrient and cation content than adjacent non-nest soil samples, but similar pH. Nutrient content was higher in ant refuse materials than in nest soils. The fertilizer effect of ant nests was also higher in dry habitats than in grasslands or savannas. Cation content was higher in nests of plant-feeding ants than in nests of omnivorous species, and lower in nests from agro-ecosystems than in nests from any other habitat. Plants showed higher green/root biomass and fitness on ant nests soils than in adjacent, non-nest sites; but plant density and diversity were unaffected by the presence of ant nests. Root growth was particularly higher in refuse materials than in ant nest soils, in leaf-cutting ant nests and in deserts habitats. Our results confirm the major role of ant nests in influencing soil fertility and vegetation patterns and provide information about the factors that mediate these effects. First, ant nests improve soil fertility mainly through the accumulation of refuse materials. Thus, different refuse dump locations (external or in underground nest chambers) could benefit different vegetation life-forms. Second, ant nests could increase plant diversity at larger spatial scales only if the identity of favoured plants changes along environmental gradients (i.e. enhancing β-diversity). Third, ant species that feed on plants play a relevant role fertilizing soils, which may balance their known influence as primary consumers. Fourth, the effects of ant nests as fertility islands are larger in arid lands, possibly because fertility is intrinsically lower in these habitats. Overall, this study provide novel and quantitative evidence confirming that ant nests are key soil modifiers, emphasizing their role as ecological engineers.

An Arctic predator–prey system in flux: climate change impacts on coastal space use by polar bears and ringed seals

Abstract: Norwegian Polar Institute The World Wild Fund for Nature (WWF) Statoil The Norwegian Academy of Science and Letters

African departure rather than migration speed determines variation in spring arrival in pied flycatchers

Abstract: Properly timed spring migration enhances reproduction and survival. Climate change requires organisms to respond to changes such as advanced spring phenology. Pied flycatchers Ficedula hypoleuca have become a model species to study such phenological adaptations of long-distance migratory songbirds to climate change, but data on individuals' time schedules outside the breeding season are still lacking. Using light-level geolocators, we studied variation in migration schedules across the year in a pied flycatcher population in the Netherlands, which sheds light on the ability for individual adjustments in spring arrival timing to track environmental changes at their breeding grounds. We show that variation in arrival dates to breeding sites in 2014 was caused by variation in departure date from sub-Saharan Africa and not by environmental conditions encountered en route. Spring migration duration was short for all individuals, on average 2 weeks. Males migrated ahead of females in spring, while migration schedules in autumn were flexibly adjusted according to breeding duties. Individuals were therefore not consistently early or late throughout the year. In fast migrants like our Dutch pied flycatchers, advancement of arrival to climate change likely requires changes in spring departure dates. Adaptation for earlier arrival may be slowed down by harsh circumstances in winter, or years with high costs associated with early migration.

Sexual selection can both increase and decrease extinction probability: reconciling demographic and evolutionary factors.

Abstract: Previous theoretical models of the effect of sexual selection on average individual fitness in a population have mostly predicted that sexually selected populations should adapt faster and clear deleterious mutations more quickly than populations where sexual selection is not operating. While some laboratory studies have supported these predictions, others have not and studies of field systems have tended to find negative effects of sexual selection, or no effect. The negative effects of sexual selection found in field and other studies are usually ascribed to the costs associated with strong sexual selection acting on the population. Here, using an individual-based model that allows feedback between demographic and evolutionary processes, we find that sexual selection can lead to both increases and decreases in population-level fitness measures such as extinction probability and adaptation rate. Whether fitness increases or decreases depends on a variety of environmental and demographic factors including the nature of environmental change, the carrying capacity of the environment, the average fecundity of the population in question and the strength of condition dependence. In many cases, our model predicts that sexual selection leads to higher extinction probability in small populations because of an increased risk of demographic stochasticity, but lower extinction probability in larger populations because of faster adaptation rates. This is consistent with field studies that have mostly focussed on very small populations such as recently introduced birds, and tend to find negative effects, and also with laboratory studies that tend to use larger populations and have tended to find positive effects. These findings go at least some way towards an understanding of the apparent contradictions between theoretical predictions, laboratory studies and field data.

Sensitivity of UK butterflies to local climatic extremes: which life stages are most at risk?

Abstract: Summary There is growing recognition as to the importance of extreme climatic events (ECEs) in determining changes in species populations In fact, it is often the extent of climate variability that determines a population's ability to persist at a given site This study examined the impact of ECEs on the resident UK butterfly species (n = 41) over a 37-year period The study investigated the sensitivity of butterflies to four extremes (drought, extreme precipitation, extreme heat and extreme cold), identified at the site level, across each species' life stages Variations in the vulnerability of butterflies at the site level were also compared based on three life-history traits (voltinism, habitat requirement and range) This is the first study to examine the effects of ECEs at the site level across all life stages of a butterfly, identifying sensitive life stages and unravelling the role life-history traits play in species sensitivity to ECEs Butterfly population changes were found to be primarily driven by temperature extremes Extreme heat was detrimental during overwintering periods and beneficial during adult periods and extreme cold had opposite impacts on both of these life stages Previously undocumented detrimental effects were identified for extreme precipitation during the pupal life stage for univoltine species Generalists were found to have significantly more negative associations with ECEs than specialists With future projections of warmer, wetter winters and more severe weather events, UK butterflies could come under severe pressure given the findings of this study

Habitat connectivity and local conditions shape taxonomic and functional diversity of arthropods on green roofs.

Abstract: Summary Increasing development of urban environments creates high pressure on green spaces with potential negative impacts on biodiversity and ecosystem services. There is growing evidence that green roofs – rooftops covered with vegetation – can contribute mitigate the loss of urban green spaces by providing new habitats for numerous arthropod species. Whether green roofs can contribute to enhance taxonomic and functional diversity and increase connectivity across urbanized areas remains, however, largely unknown. Furthermore, only limited information is available on how environmental conditions shape green roof arthropod communities. We investigated the community composition of arthropods (Apidae, Curculionidae, Araneae and Carabidae) on 40 green roofs and 40 green sites at ground level in the city of Zurich, Switzerland. We assessed how the site's environmental variables (such as area, height, vegetation, substrate and connectivity among sites) affect species richness and functional diversity using generalized linear models. We used an extension of co-inertia analysis (RLQ) and fourth-corner analysis to highlight the mechanism underlying community assemblages across taxonomic groups on green roof and ground communities. Species richness was higher at ground-level sites, while no difference in functional diversity was found between green roofs and ground sites. Green roof arthropod diversity increased with higher connectivity and plant species richness, irrespective of substrate depth, height and area of green roofs. The species trait analysis reviewed the mechanisms related to the environmental predictors that shape the species assemblages of the different taxa at ground and roof sites. Our study shows the important contribution of green roofs in maintaining high functional diversity of arthropod communities across different taxonomic groups, despite their lower species richness compared with ground sites. Species communities on green roofs revealed to be characterized by specific trait assemblages. The study also provides details on the environmental conditions that influence arthropod diversity and gives new perspectives on how the design of green roofs can be improved to increase their ecological value. Furthermore, the study highlights the importance of integrating green roofs in planning policies which aim to enhance urban habitat connectivity.

Why are some plant-pollinator networks more nested than others?

Abstract: Summary Empirical studies have found that the mutualistic interactions forming the structure of plant-pollinator networks are typically more nested than expected by chance alone. Additionally, theoretical studies have shown a positive association between the nested structure of mutualistic networks and community persistence. Yet, it has been shown that some plant-pollinator networks may be more nested than others, raising the interesting question of which factors are responsible for such enhanced nested structure. It has been argued that ordered network structures may increase the persistence of ecological communities under less predictable environments. This suggests that nested structures of plant-pollinator networks could be more advantageous under highly seasonal environments. While several studies have investigated the link between nestedness and various environmental variables, unfortunately, there has been no unified answer to validate these predictions. Here, we move from the problem of describing network structures to the problem of comparing network structures. We develop comparative statistics, and apply them to investigate the association between the nested structure of 59 plant-pollinator networks and the temperature seasonality present in their locations. We demonstrate that higher levels of nestedness are associated with a higher temperature seasonality. We show that the previous lack of agreement came from an extended practice of using standardized measures of nestedness that cannot be compared across different networks. Importantly, our observations complement theory showing that more nested network structures can increase the range of environmental conditions compatible with species coexistence in mutualistic systems, also known as structural stability. This increase in nestedness should be more advantageous and occur more often in locations subject to random environmental perturbations, which could be driven by highly changing or seasonal environments. This synthesis of theory and observations could prove relevant for a better understanding of the ecological processes driving the assembly and persistence of ecological communities. This article is protected by copyright. All rights reserved.

Functional and phylogenetic structure of island bird communities

Abstract: Biodiversity change in anthropogenically transformed habitats is often nonrandom, yet the nature and importance of the different mechanisms shaping community structure are unclear. Here, we extend the classic Theory of Island Biogeography (TIB) to account for nonrandom processes by incorporating species traits and phylogenetic relationships into a study of faunal relaxation following habitat loss and fragmentation. Two possible mechanisms can create nonrandom community patterns on fragment islands. First, small and isolated islands might consist of similar or closely related species because they are environmentally homogeneous or select for certain shared traits, such as dispersal ability. Alternatively, communities on small islands might contain more dissimilar or distantly related species than on large islands because limited space and resource availability result in greater competitive exclusion among species with high niche overlap. Breeding birds were surveyed on 36 islands and two mainland sites annually from 2010 to 2014 in the Thousand Island Lake region, China. We assessed community structure of breeding birds on these subtropical land-bridge islands by integrating species' trait and evolutionary distances. We additionally analysed habitat heterogeneity and variance in size ratios to distinguish biotic and abiotic processes of community assembly. Results showed that functional-phylogenetic diversity increased with island area, and decreased with isolation. Bird communities on the mainland were more diverse and generally less clustered than island bird communities and not different than randomly assembled communities. Bird communities on islands tend to be functionally similar and phylogenetically clustered, especially on small and isolated islands. The nonrandom decline in species diversity and change in bird community structure with island area and isolation, along with the relatively homogeneous habitats on small islands, support the environmental filtering hypothesis. Our study demonstrates the importance of integrating multiple forms of diversity for understanding the effects of habitat loss and fragmentation, and further reveals that TIB could be extended to community measures by moving beyond assumptions of species equivalency in colonisation rates and extinction susceptibilities.

Land-use type and intensity differentially filter traits in above- and below-ground arthropod communities

Abstract: Along with the global decline of species richness goes a loss of ecological traits. Associated biotic homogenization of animal communities and narrowing of trait diversity threaten ecosystem functioning and human well-being. High management intensity is regarded as an important ecological filter, eliminating species that lack suitable adaptations. Below-ground arthropods are assumed to be less sensitive to such effects than above-ground arthropods. Here, we compared the impact of management intensity between (grassland vs. forest) and within land-use types (local management intensity) on the trait diversity and composition in below- and above-ground arthropod communities. We used data on 722 arthropod species living above-ground (Auchenorrhyncha and Heteroptera), primarily in soil (Chilopoda and Oribatida) or at the interface (Araneae and Carabidae). Our results show that trait diversity of arthropod communities is not primarily reduced by intense local land use, but is rather affected by differences between land-use types. Communities of Auchenorrhyncha and Chilopoda had significantly lower trait diversity in grassland habitats as compared to forests. Carabidae showed the opposite pattern with higher trait diversity in grasslands. Grasslands had a lower proportion of large Auchenorrhyncha and Carabidae individuals, whereas Chilopoda and Heteroptera individuals were larger in grasslands. Body size decreased with land-use intensity across taxa, but only in grasslands. The proportion of individuals with low mobility declined with land-use intensity in Araneae and Auchenorrhyncha, but increased in Chilopoda and grassland Heteroptera. The proportion of carnivorous individuals increased with land-use intensity in Heteroptera in forests and in Oribatida and Carabidae in grasslands. Our results suggest that gradients in management intensity across land-use types will not generally reduce trait diversity in multiple taxa, but will exert strong trait filtering within individual taxa. The observed patterns for trait filtering in individual taxa are not related to major classifications into above- and below-ground species. Instead, ecologically different taxa resembled each other in their trait diversity and compositional responses to land-use differences. These previously undescribed patterns offer an opportunity to develop management strategies for the conservation of trait diversity across taxonomic groups in permanent grassland and forest habitats.

Artificial light at night alters trophic interactions of intertidal invertebrates.

Abstract: Summary Despite being globally widespread in coastal regions, the impacts of light pollution on intertidal ecosystems has received little attention. Intertidal species exhibit many night-time-dependent ecological strategies, including feeding, reproduction, orientation and predator avoidance, which are likely negatively affected by shifting light regimes, as has been observed in terrestrial and aquatic taxa. Coastal lighting may shape intertidal communities through its influence on the nocturnal foraging activity of dogwhelks (Nucella lapillus), a widespread predatory mollusc that structures biodiversity in temperate rocky shores. In the laboratory, we investigated whether the basal and foraging activity of this predator was affected by exposure to night-time lighting both in the presence and absence of olfactory predator cues (Carcinus maenas, common shore crab). Assessments of dogwhelks’ behavioural responses to night-time white LED lighting were performed on individuals that had been acclimated to night-time white LED lighting conditions for 16 days and individuals that had not previously been exposed to artificial light at night. Dogwhelks acclimated to night-time lighting exhibited natural refuge-seeking behaviour less often compared to control animals, but were more likely to respond to and handle prey irrespective of whether olfactory predator cues were present. These responses suggest night-time lighting likely increased the energetic demand of dogwhelks through stress, encouraging foraging whenever food was available, regardless of potential danger. Contrastingly, whelks not acclimated under night-time lighting were more likely to respond to the presence of prey under artificial light at night when olfactory predator cues were present, indicating an opportunistic shift towards the use of visual instead of olfactory cues in risk evaluation. These results demonstrate that artificial night-time lighting influences the behaviour of intertidal fauna such that the balance of interspecific interactions involved in community structuring may be affected.

Seasonal detours by soaring migrants shaped by wind regimes along the East Atlantic Flyway.

Abstract: 1. Avian migrants often make substantial detours between their seasonal destinations. It is likely some species do this to make the most of predictable wind regimes along their respective flyways. We test this hypothesis by studying orientation behaviour of a long-distance soaring migrant in relation to prevailing winds along the East Atlantic Flyway. 2. We tracked 62 migratory journeys of 12 adult European Honey Buzzards Pernis apivorus with GPS loggers. Hourly fixes were annotated with local wind vectors from a global atmospheric model to determine orientation behaviours with respect to the buzzards’ seasonal goal destinations. This enabled us to determine hot spots where buzzards overdrifted and overcompensated for side winds. We then determined whether winds along the buzzards’ detours differed from winds prevailing elsewhere in the flyway. 3. Honey Buzzards cross western Africa using different routes in autumn and spring. In autumn, they overcompensated for westward winds to circumvent the Atlas Mountains on the eastern side and then overdrifted with south-westward winds while crossing the Sahara. In spring, however, they frequently overcompensated for eastward winds to initiate a westward detour at the start of their journey. They later overdrifted with side winds north-westward over the Sahel and north-eastward over the Sahara, avoiding adverse winds over the central Sahara. 4. We conclude that Honey Buzzards make seasonal detours to utilize more supportive winds further en route and thereby expend less energy while crossing the desert. Lifelong tracking studies will be helpful to elucidate how honey buzzards and other migrants learn complex routes to exploit atmospheric circulation patterns from local to synoptic scales.

Hot dogs: High ambient temperatures impact reproductive success in a tropical carnivore

Abstract: Climate change imposes an urgent need to recognise and conserve the species likely to be worst affected. However, while ecologists have mostly explored indirect effects of rising ambient temperatures on temperate and polar species, physiologists have predicted direct impacts on tropical species. The African wild dog (Lycaon pictus), a tropical species, exhibits few of the traits typically used to predict climate change vulnerability. Nevertheless, we predicted that wild dog populations might be sensitive to weather conditions, because the species shows strongly seasonal reproduction across most of its geographical range. We explored associations between weather conditions, reproductive costs, and reproductive success, drawing on long-term wild dog monitoring data from sites in Botswana (20°S, 24 years), Kenya (0°N, 12 years), and Zimbabwe (20°S, 6 years). High ambient temperatures were associated with reduced foraging time, especially during the energetically costly pup-rearing period. Across all three sites, packs which reared pups at high ambient temperatures produced fewer recruits than did those rearing pups in cooler weather; at the non-seasonal Kenya site such packs also had longer inter-birth intervals. Over time, rising ambient temperatures at the (longest-monitored) Botswana site coincided with falling wild dog recruitment. Our findings suggest a direct impact of high ambient temperatures on African wild dog demography, indicating that this species, which is already globally endangered, may be highly vulnerable to climate change. This vulnerability would have been missed by simplistic trait-based assessments, highlighting the limitations of such assessments. Seasonal reproduction, which is less common at low latitudes than at higher latitudes, might be a useful indicator of climate change vulnerability among tropical species.

Niche conservatism and the invasive potential of the wild boar

Abstract: Niche conservatism, i.e. the retention of a species' fundamental niche through evolutionary time, is cornerstone for biological invasion assessments. The fact that species tend to maintain their original climate niche allows predictive maps of invasion risk to anticipate potential invadable areas. Unravelling the mechanisms driving niche shifts can shed light on the management of invasive species. Here, we assessed niche shifts in one of the world's worst invasive species: the wild boar Sus scrofa. We also predicted potential invadable areas based on an ensemble of three ecological niche modelling methods, and evaluated the performance of models calibrated with native vs. pooled (native plus invaded) species records. By disentangling the drivers of change on the exotic wild boar population's niches, we found strong evidence for niche conservatism during biological invasion. Ecological niche models calibrated with both native and pooled range records predicted convergent areas. Also, observed niche shifts are mostly explained by niche unfilling, i.e. there are unoccupied areas in the exotic range where climate is analogous to the native range. Niche unfilling is expected as result of recent colonization and ongoing dispersal, and was potentially stronger for the Neotropics, where a recent wave of introductions for pig-farming and game-hunting has led to high wild boar population growth rates. The invasive potential of wild boar in the Neotropics is probably higher than in other regions, which has profound management implications if we are to prevent their invasion into species-rich areas, such as Amazonia, coupled with expansion of African swine fever and possibly great economic losses. Although the originally Eurasian-wide distribution suggests a pre-adaptation to a wide array of climates, the wild boar world-wide invasion does not exhibit evidence of niche evolution. The invasive potential of the wild boar therefore probably lies on the reproductive, dietary and morphological characteristics of this species, coupled with behavioural thermoregulation.

Genetic mixture of multiple source populations accelerates invasive range expansion.

Abstract: A wealth of population genetic studies have documented that many successful biological invasions stem from multiple introductions from genetically distinct source populations. Yet, mechanistic understanding of whether and how genetic mixture promotes invasiveness has lagged behind documentation that such mixture commonly occurs. We conducted a laboratory experiment to test the influence of genetic mixture on the velocity of invasive range expansion. The mechanistic basis for effects of genetic mixture could include evolutionary responses (mixed invasions may harbour greater genetic diversity and thus elevated evolutionary potential) and/or fitness advantages of between-population mating (heterosis). If driven by evolution, positive effects of source population mixture should increase through time, as selection sculpts genetic variation. If driven by heterosis, effects of mixture should peak following first reproductive contact and then dissipate. Using a laboratory model system (beetles spreading through artificial landscapes), we quantified the velocity of range expansion for invasions initiated with one, two, four or six genetic sources over six generations. Our experiment was designed to test predictions corresponding to the evolutionary and heterosis mechanisms, asking whether any effects of genetic mixture occurred in early or later generations of range expansion. We also quantified demography and dispersal for each experimental treatment, since any effects of mixture should be manifest in one or both of these traits. Over six generations, invasions with any amount of genetic mixture (two, four and six sources) spread farther than single-source invasions. Our data suggest that heterosis provided a 'catapult effect', leaving a lasting signature on range expansion even though the benefits of outcrossing were transient. Individual-level trait data indicated that genetic mixture had positive effects on local demography (reduced extinction risk and enhanced population growth) during the initial stages of invasion but no consistent effects on dispersal ability. Our work is the first to demonstrate that genetic mixture can alter the course of spatial expansion, the stage of invasion typically associated with the greatest ecological and economic impacts. We suggest that similar effects of genetic mixture may be a common feature of biological invasions in nature, but that these effects can easily go undetected.

Thermal physiology: A new dimension of the pace-of-life syndrome.

Abstract: 1.Current syndrome research focuses primarily on behavior with few incorporating components of physiology. One such syndrome is the Pace-of-Life Syndrome (POLS) which describes covariation between behaviour, metabolism immunity, hormonal response, and life history traits. Despite the strong effect temperature has on behavior, thermal physiology has yet to be considered within this syndrome framework. 2.We proposed the POLS to be extended to include a new dimension, the cold-hot axis. Under this premise, it is predicted that thermal physiology and behavior would covary whereby individual positioning along the thermal continuum would coincide with that of the behavioral continuum. 3.This hypothesis was tested by measuring thermal traits of delicate skinks (Lampropholis delicata) and linking it to their behavior. Principal components analysis and structural equation modelling were used to determine if traits were structured within the Pace-of-Life Syndrome (POLS) and to characterize the direction of their interactions. 4.Model results supported the inclusion of the cold-hot axis into the POLS and indicated that thermal physiology was the driver of this relationship, in that thermal traits either constrained or promoted activity, exploration, boldness, and social behavior. 5.This study highlights the need to integrate thermal physiology within a syndrome framework. This article is protected by copyright. All rights reserved.

Caching reduces kleptoparasitism in a solitary, large felid

Abstract: Food caching is a common strategy used by a diversity of animals, including carnivores, to store and/or secure food. Despite its prevalence, the drivers of caching behaviour, and its impacts on individuals, remain poorly understood, particularly for short-term food cachers. Leopards Panthera pardus exhibit a unique form of short-term food caching, regularly hoisting, storing and consuming prey in trees. We explored the factors motivating such behaviour among leopards in the Sabi Sand Game Reserve, South Africa, associated with four not mutually exclusive hypotheses: food-perishability, consumption-time, resource-pulse and kleptoparasitism-avoidance. Using data from 2032 prey items killed by 104 leopards from 2013 to 2015, we built generalized linear mixed models to examine how hoisting behaviour, feeding time and the likelihood of a kill being kleptoparasitized varied with leopard sex and age, prey size and vulnerability, vegetation, elevation, climate, and the immediate and long-term risk posed by dominant competitors. Leopards hoisted 51% of kills. They were more likely to hoist kills of an intermediate size, outside of a resource pulse and in response to the presence of some competitors. Hoisted kills were also fed on for longer than non-hoisted kills. At least 21% of kills were kleptoparasitized, mainly by spotted hyaenas Crocuta crocuta. Kills were more likely to be kleptoparasitized at lower temperatures and if prey were larger, not hoisted, and in areas where the risk of encountering hyaenas was greatest. Female leopards that suffered higher rates of kleptoparasitism exhibited lower annual reproductive success than females that lost fewer kills. Our results strongly support the kleptoparasitism-avoidance hypothesis and suggest hoisting is a key adaptation that enables leopards to coexist sympatrically with high densities of competitors. We further argue that leopards may select smaller-sized prey than predicted by optimal foraging theory, to balance trade-offs between kleptoparasitic losses and the energetic gains derived from killing larger prey. Although caching may provide the added benefits of delaying food perishability and enabling consumption over an extended period, the behaviour primarily appears to be a strategy for leopards, and possibly other short-term cachers, to reduce the risks of kleptoparasitism.

Intraspecific competition, not predation, drives lizard tail loss on islands

Abstract: Summary Tail autotomy is mainly considered an antipredator mechanism. Theory suggests that predation pressure relaxes on islands, subsequently reducing autotomy rates. Intraspecific aggression, which may also cause tail loss, probably intensifies on islands due to the higher abundance. We studied whether tail autotomy is mostly affected by predation pressure or by intraspecific competition. We further studied whether predator abundance or predator richness is more important in this context. To test our predictions, we examined multiple populations of two gecko species: Kotschy's gecko (Mediodactylus kotschyi; mainland and 41 islands) and the Mediterranean house gecko (Hemidactylus turcicus; mainland and 17 islands), and estimated their abundance together with five indices of predation. In both species, autotomy rates are higher on islands and decline with most predation indices, in contrast with common wisdom, and increase with gecko abundance. In M. kotschyi, tail-loss rates are higher on predator and viper-free islands, but increase with viper abundance. We suggest that autotomy is not simply, or maybe even mainly, an antipredatory mechanism. Rather, such defence mechanisms are a response to complex direct and indirect biotic interactions and perhaps, in the case of tail autotomy in insular populations, chiefly to intraspecific aggression.

Sex-dependent carry-over effects on timing of reproduction and fecundity of a migratory bird.

Abstract: 1.Life of many organisms flows as a sequence of annual cycles. Timing of cyclical events is shaped by natural selection also via the domino effects that any life-history stage has on the stages that follow. Such ‘carry-over effects’ have major consequences for evolutionary, ecological and demographic processes, but the causes that generate their individual-level variation, including the effect of sex, are poorly understood. 2.We used light-level geolocators to study carry-over effects on the year-round life-cycle of the long-distance migratory barn swallow (Hirundo rustica) and sex-dependent variation in their strength. 3.Correlation analyses showed that timing of breeding influenced departure time for autumn migration in females but not in males. In addition, strong, time-mediated carry-over effects of timing of departure from the wintering areas in sub-Saharan Africa for spring migration on timing of arrival to the breeding grounds in Italy and Switzerland operated in both sexes. However, carry-over effects of spring migration phenology on breeding date and seasonal fecundity were observed among females but not among males. 4.We used partial least squares path modelling to unveil the complex carry-over effects of phenology during the non-breeding season in combination with the ecological conditions experienced by individual swallows in the wintering area, as gauged by Normalized Difference Vegetation Index values (NDVI), on breeding performance. Phenology during the non-breeding season combined with NDVI during wintering accounted for as much as 65-70% of variation in subsequent seasonal fecundity in females, while such carry-over effects on breeding success of males were weaker. 5.Intense, sex-specific carry-over effects can have impacted on evolutionary processes, including sexual selection, and affected phenological response to climate change, causing the large population decline observed in this species. This article is protected by copyright. All rights reserved.

The benefits of coinfection: trematodes alter disease outcomes associated with virus infection.

Abstract: Summary Coinfections are increasingly recognized as important drivers of disease dynamics. Consequently, greater emphasis has been placed on integrating principles from community ecology with disease ecology to understand within-host interactions among parasites. Using larval amphibians and two amphibian parasites (ranaviruses and the trematode Echinoparyphium sp.), we examined the influence of coinfection on disease outcomes. Our first objective was to examine how priority effects (the timing and sequence of parasite exposure) influence infection and disease outcomes in the laboratory. We found that interactions between the parasites were asymmetric; prior infection with Echinoparyphium reduced ranaviral loads by 9% but there was no reciprocal effect of prior ranavirus infection on Echinoparyphium load. Additionally, survival rates of hosts (larval gray treefrogs; Hyla versicolor) infected with Echinoparyphium 10 days prior to virus exposure were 25% greater compared to hosts only exposed to virus. Our second objective was to determine whether these patterns were generalizable to multiple amphibian species under more natural conditions. We conducted a semi-natural mesocosm experiment consisting of four larval amphibian hosts [gray treefrogs, American toads (Anaxyrus americanus), leopard frogs (Lithobates pipiens) and spring peepers (Pseudacris crucifer)] to examine how prior Echinoparyphium infection influenced ranavirus transmission within the community, using ranavirus-infected larval wood frogs (Lithobates sylvaticus) as source of ranavirus. Consistent with the laboratory experiment, we found that prior Echinoparyphium infection reduced ranaviral loads by 19 to 28% in three of the four species. Collectively, these results suggest that macroparasite infection can reduce microparasite replication rates across multiple amphibian species, possibly through cross-reactive immunity. Although the immunological mechanisms driving this outcome are in need of further study, trematode infections appear to benefit hosts that are exposed to ranaviruses. Additionally, these results suggest that consideration of priority effects and timing of exposure are vital for understanding parasite interactions within hosts and disease outcomes.

Carnivore carcasses are avoided by carnivores

Abstract: Ecologists have traditionally focused on herbivore carcasses as study models in scavenging research. However, some observations of scavengers avoiding feeding on carnivore carrion suggest that different types of carrion may lead to differential pressures. Untested assumptions about carrion produced at different trophic levels could therefore lead ecologists to overlook important evolutionary processes and their ecological consequences. Our general goal was to investigate the use of mammalian carnivore carrion by vertebrate scavengers. In particular, we aimed to test the hypothesis that carnivore carcasses are avoided by other carnivores, especially at the intraspecific level, most likely to reduce exposure to parasitism. We take a three-pronged approach to study this principle by: (i) providing data from field experiments, (ii) carrying out evolutionary simulations of carnivore scavenging strategies under risks of parasitic infection, and (iii) conducting a literature-review to test two predictions regarding parasite life-history strategies. First, our field experiments showed that the mean number of species observed feeding at carcasses and the percentage of consumed carrion biomass were substantially higher at herbivore carcasses than at carnivore carcasses. This occurred even though the number of scavenger species visiting carcasses and the time needed by scavengers to detect carcasses were similar between both types of carcasses. In addition, we did not observe cannibalism. Second, our evolutionary simulations demonstrated that a risk of parasite transmission leads to the evolution of scavengers with generally low cannibalistic tendencies, and that the emergence of cannibalism-avoidance behaviour depends strongly on assumptions about parasite-based mortality rates. Third, our literature review indicated that parasite species potentially able to follow a carnivore-carnivore indirect cycle, as well as those transmitted via meat consumption, are rare in our study system. Our findings support the existence of a novel coevolutionary relation between carnivores and their parasites, and suggest that carnivore and herbivore carcasses play very different roles in food webs and ecosystems.

Hunting promotes sexual conflict in brown bears.

Abstract: Summary The removal of individuals through hunting can destabilize social structure, potentially affecting population dynamics. Although previous studies have shown that hunting can indirectly reduce juvenile survival through increased sexually selected infanticide (SSI), very little is known about the spatiotemporal effects of male hunting on juvenile survival. Using detailed individual monitoring of a hunted population of brown bears (Ursus arctos) in Sweden (1991–2011), we assessed the spatiotemporal effect of male removal on cub survival. We modelled cub survival before, during and after the mating season. We used three proxies to evaluate spatial and temporal variation in male turnover; distance and timing of the closest male killed and number of males that died around a female's home range centre. Male removal decreased cub survival only during the mating season, as expected in seasonal breeders with SSI. Cub survival increased with distance to the closest male killed within the previous 1·5 years, and it was lower when the closest male killed was removed 1·5 instead of 0·5 year earlier. We did not detect an effect of the number of males killed. Our results support the hypothesis that social restructuring due to hunting can reduce recruitment and suggest that the distribution of the male deaths might be more important than the overall number of males that die. As the removal of individuals through hunting is typically not homogenously distributed across the landscape, spatial heterogeneity in hunting pressure may cause source–sink dynamics, with lower recruitment in areas of high human-induced mortality.

Seal mothers expend more on offspring under favourable conditions and less when resources are limited.

Abstract: Summary 1.In mammals, maternal expenditure on offspring is a complex mix of several factors including the species’ mating system, offspring sex and the condition and age of the mother. While theory suggests that in polygynous species mothers should wean larger male offspring than females when resources and maternal conditions allow, the evidence for this remains equivocal. 2.Southern elephant seals are highly dimorphic, polygynous capital breeders existing in an environment with highly variable resources and should therefore provide clear evidence to support the theoretical expectations of differential maternal expenditure in male and female pups. 3.We quantified maternal size (mass and length) and pup size at birth and weaning for 342 elephant seal mothers at Macquarie Island. The study was conducted over 11 years of contrasting sea-ice and Southern Annular Mode values, both indices of maternal prey resources. 4.Overall, large females weaned male pups that weighed 17 kg (15.5%) more than female pups. Maternal condition varied by as much as 59 kg among years, and was positively related to SAM, and negatively to maximum sea-ice extent. Smaller mothers weaned relatively larger male pups under favourable conditions, this effect was less apparent for larger mothers. 5.We developed a simple model linking environmental variation to maternal masses post-partum, followed by maternal masses post-partum to weaning masses and then weaning masses to pup survival and demonstrated that environmental conditions affected predicted survival so that the pups of small mothers had an estimated 7% increase in first year survival in “good” vs “bad” years compared to 1% for female pups of large mothers. 6.Co-occurrence of environmental quality and conservative reproductive tactics suggests that mothers retain substantial plasticity in maternal care, enhancing their lifetime reproductive success by adjusting reproductive expenditure relative to both prevailing environmental conditions and their own capabilities. This article is protected by copyright. All rights reserved.

A framework for modelling range shifts and migrations: asking when, whither, whether and will it return

Abstract: Summary Many animals undertake movements that are longer scaled and more directed than their typical home ranging behaviour. These movements include seasonal migrations (e.g. between breeding and feeding grounds), natal dispersal, nomadic range shifts and responses to local environmental disruptions. While various heuristic tools exist for identifying range shifts and migrations, none explicitly model the movement of the animals within a statistical framework that facilitates quantitative comparisons. We present the mechanistic range shift analysis (MRSA), a method to estimate a suite of range shift parameters: times of initiation, duration of transitions, centroids and areas of respective ranges. The method can take the autocorrelation and irregular sampling that is characteristic of much movement data into account. The mechanistic parameters suggest an intuitive measure, the range shift index, for the extent of a range shift. The likelihood based estimation further allows for statistical tests of several relevant hypotheses, including a range shift test, a stopover test and a site fidelity test. The analysis tools are provided in an R package (marcher). We applied the MRSA to a population of GPS tracked roe deer (Capreolus capreolus) in the Italian Alps between 2005 and 2008. With respect to seasonal migration, this population is extremely variable and difficult to classify. Using the MRSA, we were able to quantify the behaviours across the population and among individuals across years, identifying extents, durations and locations of seasonal range shifts, including cases that would have been ambiguous to detect using existing tools. The strongest patterns were differences across years: many animals simply did not perform a seasonal migration to wintering grounds during the mild winter of 2006–2007, even though some of these same animals did move extensively in other, harsher winters. For seasonal migrants, however, site fidelity across years was extremely high, even after skipping an entire seasonal migration. These results suggest that for roe deer behavioural plasticity and tactical responses to immediate environmental cues are reflected in the decision of whether rather than where to migrate. The MRSA also revealed a trade-off between the probability of migrating and the size of a home range.

Boldness predicts an individual's position along an exploration-exploitation foraging trade-off

Abstract: Individuals do not have complete information about the environment and therefore they face a trade-off between gathering information (exploration) and gathering resources (exploitation). Studies have shown individual differences in components of this trade-off but how stable these strategies are in a population and the intrinsic drivers of these differences is not well understood. Top marine predators are expected to experience a particularly strong trade-off as many species have large foraging ranges and their prey often have a patchy distribution. This environment leads these species to exhibit pronounced exploration and exploitation phases but differences between individuals are poorly resolved. Personality differences are known to be important in foraging behaviour but also in the trade-off between exploration and exploitation. Here we test whether personality predicts an individual exploration-exploitation strategy using wide ranging wandering albatrosses (Diomedea exulans) as a model system. Using GPS tracking data from 276 wandering albatrosses, we extract foraging parameters indicative of exploration (searching) and exploitation (foraging) and show that foraging effort, time in patch and size of patch are strongly correlated, demonstrating these are indicative of an exploration-exploitation strategy. Furthermore, we show these are consistent within individuals and appear stable in the population, with no reproductive advantage. The searching and foraging behaviour of bolder birds placed them towards the exploration end of the trade-off, whereas shy birds showed greater exploitation. This result provides a mechanism through which individual foraging strategies may emerge. Age and sex affected components of the trade-off, but not the trade-off itself, suggesting these factors may drive behavioural compensation to maintain resource acquisition and this was supported by the evidence that there were no fitness consequence of any EE trait nor the trade-off itself. These results demonstrate a clear trade-off between information gathering and exploitation of prey patches, and reveals for the first time that boldness may drive these differences. This provides a mechanism through which widely reported links between personality and foraging may emerge. This article is protected by copyright. All rights reserved.

Singing from North to South: Latitudinal variation in timing of dawn singing under natural and artificial light conditions.

Abstract: Summary 1. Animals breeding at northern latitudes experience drastic changes in daily light conditions during the breeding season with decreasing periods of darkness, whereas those living at lower latitudes are exposed to naturally dark nights throughout the year. Nowadays, many animals are also exposed to artificial night lighting (often referred to as light pollution). 2. Animals strongly rely on variation in light levels to time their daily and seasonal behaviour. Previous work on passerine birds showed that artificial night lighting leads to earlier onset of dawn song. However, these studies were carried out at intermediate latitudes with more limited seasonal changes in daylength, and we still lack an understanding of the impact of artificial night lighting in relation to variation in natural light conditions. 3. We investigated the influence of natural and artificial light conditions on the timing of dawn singing in five common songbird species in each of three regions in Europe that differed in natural variation in daylength (northern Finland, 65°N; southern Germany, 48°N; southern Spain, 37°N). In each region, we selected five peri-urban forest sites with and five without street lighting, and recorded dawn singing at the beginning of the local breeding season. 4. Our results show that the earliest natural singers, i.e., European robins (Erithacus rubecula) and common blackbirds (Turdus merula), started dawn singing earlier along with the natural increase in night brightness in Finland, with no additional effects of artificial night lighting. In contrast, the later singers, i.e., great tits (Parus major), blue tits (Cyanistes caeruleus), and chaffinches (Fringilla coelebs), showed similar onsets of dawn song relative to sunrise across the season and similar effects of artificial night lighting at all latitudes. 5. Artificial night lighting affected great tits, blue tits and chaffinches even in northern Finland where nights became very bright. Proximate factors such as differential light sensitivities may explain why early singers showed more plastic behavioural responses to naturally and artificially bright nights. The maintenance of rhythmicity in the late singers during bright northern nights and under artificial night lighting may also be an adaptive response to predation risk or costs of sleep loss. This article is protected by copyright. All rights reserved.