An economic model of the limits to foraging range in central place foragers with numerical solutions for bumblebees

1. A model is described that evaluates the maximum economic foraging range in central place foragers by using optimality criteria to discriminate between foraging sites at different distances from the forager's central place. 2. The basic model can be varied to suit foragers that optimise either their rate of net energy uptake or their foraging efficiency. 3. The model requires specification of the time and energy budgets of travel and foraging, and of the rewards obtainable at potential foraging sites. 4. The specific case of bumblebees, whose foraging ranges are poorly known, is considered. 5. Numerical solutions of the model for parameter values that represent bumblebees and their forage predict economic foraging ranges exceeding several kilometres. The model demonstrates that economics alone can explain extensive flight ranges in bees.     

Climate change is the primary driver of white‐tailed deer (Odocoileus virginianus) range expansion at the northern extent of its range; land use is secondary

Quantifying the relative influence of multiple mechanisms driving recent range expansion of non‐native species is essential for predicting future changes and for informing adaptation and management plans to protect native species. White‐tailed deer (Odocoileus virginianus) have been expanding their range into the North American boreal forest over the last half of the 20th century. This has already altered predator–prey dynamics in Alberta, Canada, where the distribution likely reaches the northern extent of its continuous range. Although current white‐tailed deer distribution is explained by both climate and human land use, the influence each factor had on the observed range expansion would depend on the spatial and temporal pattern of these changes. Our objective was to quantify the relative importance of land use and climate change as drivers of white‐tailed deer range expansion and to predict decadal changes in white‐tailed deer distribution in northern Alberta for the first half of the 21st century. An existing species distribution model was used to predict past decadal distributions of white‐tailed deer which were validated using independent data. The effects of climate and land use change were isolated by comparing predictions under theoretical “no‐change between decades” scenarios, for each factor, to predictions under observed climate and land use change. Climate changes led to more than 88%, by area, of the increases in probability of white‐tailed deer presence across all decades. The distribution is predicted to extend 100 km further north across the northeastern Alberta boreal forest as climate continues to change over the first half of the 21st century.     

The influences of snow cover,vegetation and topography on the upper range limit of common wombats Vombatus ursinus in the subalpine zone,Australia

Aim In subalpine and alpine environments, range shifts of species to higher altitudes are predicted to occur in response to reductions in the snow cover from climate change. However, the distribution of key resources may constrain the range of animal populations and prevent any upward migration. This study examined the local resource constraints on the upper range limit of a large, native herbivore in Australia, the common wombat Vombatus ursinus. Location The subalpine zone of the Snowy Mountains, Australia. Methods Logistic regression analyses of snow and habitat predictors were conducted on the presence/absence of wombat signs recorded along an altitudinal gradient during winter and summer, using parametric and nonparametric methods. Results Wombats responded strongly to the altitudinal gradient, but snow cover alone did not fully explain their upper range limit. Wombat occurrence in the subalpine zone was influenced by local habitat features in combination with maximum snow depth. More rugged, high‐relief terrain was important to wombats in winter, allowing individuals access to a wider range of altitudes, snow depths and shelter sites. During summer, high soil bulk density was an important predictor of occurrence, and in both seasons, occurrence declined in response to a higher cover of burnt grass. Main conclusions These models demonstrate that local habitat factors play a role even where there are strong regulating environmental factors. For wombats, this may limit future range expansion into the alpine zone despite the potential for an increase in abundance at their present range limit. These findings show the need for local ecological studies to be conducted in parallel with broad scale climate modelling if we are to understand shifts in species distributions as the climate rapidly changes.     

Temperature thresholds for germination in 20 short‐range endemic plant species from a Greenstone Belt in southern Western Australia

• The study of climate‐driven effects on seed traits such as germination has gained momentum over the past decade as the impact of global warming becomes more apparent on the health and survival of plant diversity.
• Seed response to warming was evaluated in a suite of short‐range endemic species from the biodiverse Greenstone Belt of southern Western Australia. The temperature dimensions for germination in 20 woody perennials were identified using small unreplicated samples over 6 weeks on a temperature gradient plate (constant and fluctuating temperatures between 5 and 40 °C). These data were subsequently modelled against current and forecast (2070) mean monthly minimum and maximum temperatures to illustrate seasonal changes to germination timing and final percentage germination.
• All but one species attained full germination in at least one cell on the gradient plate. Modelling of the data suggested only minimal changes to percentage germination despite a forecast rise in diurnal temperatures over the next 50 years. Nine species were predicted to experience declines of between <1% and 7%, whilst 11 species were predicted to increase their germination by <1% to 3%. Overall, the speed of germination is predicted to increase but the timing of germination for most species shifts seasonally (both advances and delays) as a result of changing diurnal temperatures.
• The capacity of this suite of species to cope with warmer temperatures during a critical early life stage shows a degree of adaptation to heterogeneous environments. Predicting the effects of global change on terrestrial plant communities is crucial to managing and conserving plant diversity.

     

A test for a shift in the boundary of the geographical range of a species

One predicted impact of climate change is a poleward shift in the boundaries of species ranges. Existing methods for identifying such a boundary shift based on changes in the observed pattern of occupancy within a grid of cells are sensitive to changes in the overall rate of sightings and their latitudinal distribution that are unconnected to a boundary shift. A formal test for a boundary shift is described that allows for such changes. The test is applied to detect northward shifts in the northern boundary of the Essex skipper (Thymelicus lineola) butterfly and the European goldfinch (Carduelis carduelis) in Great Britain. A shift is detected in the latter case but not in the former. Results from a simulation study are presented showing that the test performs well.     

Expressed mutational load increases toward the edge of a species’ geographic range

There is no general explanation for why species have restricted geographic distributions. One hypothesis posits that range expansion or increasing scarcity of suitable habitat results in accumulation of mutational load due to enhanced genetic drift, which constrains population performance toward range limits and further expansion. We tested this hypothesis in the North American plant, Arabidopsis lyrata. We experimentally assessed mutational load by crossing plants of 20 populations from across the entire species range and by raising the offspring of within- and between-population crosses at five common garden sites within and beyond the range. Offspring performance was tracked over three growing seasons. The heterosis effect, depicting expressed mutational load, was increased in populations with heightened genomic estimates of load, longer expansion distance or long-term isolation, and a selfing mating system. The decline in performance of within-population crosses amounted to 80%. Mutation accumulation due to past range expansion and long-term isolation of populations in the area of range margins is therefore a strong determinant of population-mean performance, and the magnitude of effect may be sufficient to cause range limits.     

Interrogating recent range changes in South African birds: confounding signals from land use and climate change present a challenge for attribution

Aim Apparent anthropogenic warming has been underway in South Africa for several decades, a period over which significant range shifts have been observed in some indigenous bird species. We asked whether these range shifts by birds are clearly consistent with either climate change or land use change being the primary driver. Location South Africa. Methods We categorized recent range changes among 408 South African terrestrial bird species and, using generalized linear mixed models, analysed ecological attributes of those species that have and have not changed their ranges. Results Fifty‐six of the 408 taxa studied have undergone significant range shifts. Most extended their ranges towards the south (towards cooler latitudes, consistent with climate‐change drivers) or west (towards drier and warmer habitats, inconsistent with climate drivers but consistent with land use drivers); very few moved east or north. Both southward and westward movers were habitat generalists. Furthermore, southward movers were mobile taxa (migrants and nomads), whereas westward movers were associated with human‐modified elements in the landscape, such as croplands, plantations or buildings. Main conclusions The results suggest that both land use changes and climate change may simultaneously be influencing dynamic range shifts by South African birds, but separating the relative strengths of these two drivers is challenging, not least because both are operating concurrently and may influence some species simultaneously. Those species that respond to land use change by contracting their ranges are likely to be among the species that will be most impacted by climate change if land use practices with negative impacts are occurring in areas anticipated to become climatic refugia for these species. This highlights a pressing need to develop dynamic models of species’ potential range shifts and changing abundances that incorporate population and dispersal processes, as well as ecological processes that influence habitat suitability.     

BIODIVERSITY RESEARCH: Current distribution and predicted geographic expansion of the Rufous‐backed Robin in Mexico: a fading endemism?

Aim The Rufous‐backed Robin Turdus rufopalliatus is a bird endemic to the Pacific slope of Mexico. The species recently established populations in several localities in the Mexican Central Highlands. Based on available data, we modelled the range expansion of the Rufous‐backed Robin in Mexico to understand the pattern, mechanisms and ecological and biogeographic implications of its expansion. Location Mexico. Methods We assessed the species’ presence and habitat requirements at two spatial scales. At the site level, we evaluated the relationship between land use and species presence in an urban environment. At the country level, we generated a niche model. We then produced a dispersion model through the interpolation of points generated from information derived from the niche model, the location of records within and outside its native distribution range, the species’ natural history, habitat requirements and its estimated dispersion rate (4.2 km year^?1). Results The dispersion model predicted that the species will significantly increase its distribution range in Mexico in the coming decades. Its expansion would occur by a stepping‐stone colonization of suitable habitat in areas of native vegetation and human settlements. The model predicted that the species should arrive on the Gulf slope of Mexico before 2025. Main conclusions Mechanisms that could explain the species’ success in establishing viable populations outside its native distribution include its dispersion ability, competitive release, the urban heat island phenomenon and the trade of wild birds. The geographic range expansion of the Rufous‐backed Robin will probably create new interactions with other species, particularly with close taxonomic and ecological relatives. The increase in the distribution range of the Rufous‐backed Robin has resulted from direct and indirect human‐induced dispersion; therefore, it cannot be considered a fading endemism. In part of its expanded range (to date the Mexican Central Highlands), it should be considered an invasive alien species.     

Reconstructing range dynamics and range fragmentation of European bison for the last 8000 years

Aim Understanding what constituted species’ ranges prior to large‐scale human influence, and how past climate and land use change have affected range dynamics, provides conservation planners with important insights into how species may respond to future environmental change. Our aim here was to reconstruct the Holocene range of European bison (Bison bonasus) by combining a time‐calibrated species distribution models (SDM) with a dynamic vegetation model. Location Europe. Method We used European bison occurrences from the Holocene in a maximum entropy model to assess bison range dynamics during the last 8000 years. As predictors, we used bioclimatic variables and vegetation reconstructions from the generalized dynamic vegetation model LPJ‐GUESS. We compared our range maps with maps of farmland and human population expansion to identify the main species range constraints. Results The Holocene distribution of European bison was mainly determined by vegetation patterns, with bison thriving in both broadleaved and coniferous forests, as well as by mean winter temperature. The heartland of European bison was in Central and Eastern Europe, whereas suitable habitat in Western Europe was scarce. While environmentally suitable regions were overall stable, the expansion of settlements and farming severely diminished available habitat. Main conclusions European bison habitat preferences may be wider than previously assumed, and our results suggest that the species had a more eastern and northern distribution than previously reported. Vegetation and climate transformation during the Holocene did not affect the bison’s range substantially. Conversely, human population growth and the spread of farming resulted in drastic bison habitat loss and fragmentation, likely reaching a tipping point during the last 1000 years. Combining SDM and dynamic vegetation models can improve range reconstructions and projections, and thus help to identify resilient conservation strategies for endangered species.     

Modelling the effect of habitat fragmentation on range expansion in a butterfly

There is an increasing need for conservation programmes to make quantitative predictions of biodiversity responses to changed environments. Such predictions will be particularly important to promote species recovery in fragmented landscapes, and to understand and facilitate distribution responses to climate change. Here, we model expansion rates of a test species (a rare butterfly, Hesperia comma) in five landscapes over 18 years (generations), using a metapopulation model (the incidence function model). Expansion rates increased with the area, quality and proximity of habitat patches available for colonization, with predicted expansion rates closely matching observed rates in test landscapes. Habitat fragmentation constrained expansion, but in a predictable way, suggesting that it will prove feasible both to understand variation in expansion rates and to develop conservation programmes to increase rates of range expansion in such species.     

Stream temperature and the potential growth and survival of juvenile Oncorhynchus mykiss in a southern California creek

1. We asked whether an increase in food supply in the field would increase the ability of fish populations to withstand climate warming, as predicted by certain bioenergetic models and aquarium experiments. 2. We subsidised the in situ food supply of wild juvenile steelhead (Oncorhynchus mykiss) in a small stream near the species’ southern limit. High‐quality food (10% of fish biomass per day) was added to the drift in eight in‐stream enclosures along a naturally‐occurring thermal gradient. 3. The temperatures during the experiment were well below the upper thermal limit for the species (means of enclosures ranged from 15.1 to 16.5 °C). Food supplements had no discernible effect on survival, but raised mean (± SD) specific growth rate substantially, from 0.038 ± 0.135 in controls to 2.28 ± 0.51 in feeding treatments. Food supplements doubled the variation in growth among fish. 4. The mean and variance of water temperature were correlated across the enclosures, and were therefore transformed into principal component scores T₁ (which expressed the stream‐wide correlation pattern) and T₂ (which expressed local departures from the pattern). Even though T₁ accounted for 96% of the variation in temperature mean and variance, it was not a significant predictor of fish growth. T₂ was a significant predictor of growth. The predicted time to double body mass in an enclosure with a large T₂ score (cool‐variable) was half that in an enclosure with a low T₂ score (warm‐stable). 5. Contrary to expectation, temperature effects were neutral, at least with respect to the main axis of variation among enclosures (cool‐stable versus warm‐variable). Along the orthogonal axis (cool‐variable versus warm‐stable), the effect was opposite from expectations, probably because of temperature variation. Subtle patterns of temperature heterogeneity in streams can be important to potential growth of O. mykiss.     

Expanding northward: influence of climate change,forest connectivity,and population processes on a threatened species' range shift

Species' ranges are dynamic, shifting in response to a large number of interrelated ecological and anthropogenic processes. Climate change is thought to be one of the most influential drivers of range shifts, but the effects of other confounded ecological processes are often ignored even though these processes may modify expected range responses to climate change. To determine the relative effects of climate, forest availability, connectivity, and biotic processes such as immigration and establishment, we examine range changes occurring in a species of bird, the Hooded Warbler (Wilsonia citrina). We focus predominantly on the periphery of the species' northern range in Canada but we also examine data from the entire species' range. Nesting records in southern Ontario were obtained from two breeding bird Atlases of Ontario separated by a period of 20 years (1981–1985 and 2001–2005), and the rate of range expansion was estimated by comparing the number of occupied areas in each Atlas. Twelve hypotheses of the relationship between the rate of range expansion and factors known to influence range change were examined using model‐selection techniques and a mixed modeling approach (zero‐inflated Poisson's regression). Cooler temperatures were positively related to a lack of range expansion indicating that climate constrained the species' distribution. Establishment probability (based on the number of occupied, neighboring Atlas squares) and immigration from populations to the south (estimated using independent data from the North American Breeding Bird Survey) were also important predictors of range expansion. These biotic process variables can mask the effects of forest availability and connectivity on range expansion. Expansion due to climate change may be slower in fragmented systems, but the rate of expansion will be influenced largely by biotic processes such as proximity to neighboring populations.     

Population genetic analysis of a recent range expansion: mechanisms regulating the poleward range limit in the volcano barnacle Tetraclita rubescens

As range shifts coincident with climate change have become increasingly well documented, efforts to describe the causes of range boundaries have increased. Three mechanisms—genetic impoverishment, migration load, or a physical barrier to dispersal—are well described theoretically, but the data needed to distinguish among them have rarely been collected. We describe the distribution, abundance, genetic variation, and environment of Tetraclita rubescens, an intertidal barnacle that expanded its northern range limit by several hundreds of kilometres from San Francisco, CA, USA, since the 1970s. We compare geographic variation in abundance with abiotic and biotic patterns, including sea surface temperatures and the distributions of 387 co‐occurring species, and describe genetic variation in cytochrome c oxidase subunit I, mitochondrial noncoding region, and nine microsatellite loci from 27 locations between Bahia Magdalena (California Baja Sur, Mexico) and Cape Mendocino (CA, USA). We find very high gene flow, high genetic diversity, and a gradient in physical environmental variation coincident with the range limit. We infer that the primary cause of the northern range boundary in T. rubescens is migration load arising from flow of maladapted alleles into peripheral locations and that environmental change, which could have reduced selection against genotypes immigrating into the newly colonized portion of the range, is the most likely cause of the observed range expansion. Because environmental change could similarly affect all taxa in a region whose distributional limits are established by migration load, these mechanisms may be common causes of range boundaries and largely synchronous multi‐species range expansions.     

Timing of diapause induction and its life‐history consequences in Nezara viridula: is it costly to expand the distribution range?

Abstract. 1. From the early 1960s to 2000 Nezara viridula (Heteroptera: Pentatomidae) expanded its range northwards in Japan and reached Osaka following climate warming recorded in the region. 2. The timing of diapause induction and its effect on life‐history traits were studied under quasi‐natural conditions in Osaka. Egg masses were placed outdoors in six series in July–November 1999. Developmental events were monitored until September 2000. 3. Adult diapause was induced in September–October in agreement with the photoperiodic response obtained under laboratory conditions. Induction of diapause in early October ensured the highest winter survival. Nymphs that hatched after late September died by December–April showing that the species cannot survive winter in the nymphal stage. 4. Life‐history traits varied between the early (non‐diapause reproduction) and late (post‐diapause reproduction) series. Thus, non‐diapause females produced significantly fewer egg masses than did females that reproduced only after diapause. The timing of diapause induction strongly affected overwintering success and post‐diapause performance: females that became adults and entered diapause in October lived longer, had a longer period of oviposition, and produced more eggs in larger egg masses than females that attained adulthood and entered diapause in September. 5. Females from the early series reproduced until late November, although progeny from the late September eggs were destined to die during the winter. Pre‐winter reproduction of adults that emerged in mid‐September or later was a result of the imperfect timing of diapause induction. It is an ineffective allocation of resources and may be considered the ecological cost of range expansion. 6. To establish in the region, N. viridula will probably evolve a lengthening of the critical photoperiod of the diapause induction response. This will allow the species to enter diapause earlier and, thus, avoid maladaptive late‐autumn reproduction but, perhaps, increase the cost of diapause because of a possible adverse impact of pre‐winter high temperature conditions on overwintering.     

Evidence for evolutionary change associated with the recent range expansion of the British butterfly, Aricia agestis, in response to climate change

Poleward range expansions are widespread responses to recent climate change and are crucial for the future persistence of many species. However, evolutionary change in traits such as colonization history and habitat preference may also be necessary to track environmental change across a fragmented landscape. Understanding the likelihood and speed of such adaptive change is important in determining the rate of species extinction with ongoing climate change. We conducted an amplified fragment length polymorphism (AFLP)‐based genome scan across the recently expanded UK range of the Brown Argus butterfly, Aricia agestis, and used outlier‐based (DFDIST and BayeScan) and association‐based (Isolation‐By‐Adaptation) statistical approaches to identify signatures of evolutionary change associated with range expansion and habitat use. We present evidence for (i) limited effects of range expansion on population genetic structure and (ii) strong signatures of selection at approximately 5% AFLP loci associated with both the poleward range expansion of A. agestis and differences in habitat use across long‐established and recently colonized sites. Patterns of allele frequency variation at these candidate loci suggest that adaptation to new habitats at the range margin has involved selection on genetic variation in habitat use found across the long‐established part of the range. Our results suggest that evolutionary change is likely to affect species’ responses to climate change and that genetic variation in ecological traits across species’ distributions should be maximized to facilitate range shifts across a fragmented landscape, particularly in species that show strong associations with particular habitats.