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.     

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.     

Marginal range expansion in a host-limited butterfly species Gonepteryx rhamni

1. The British distribution of the butterfly Gonepteryx rhamni (L.) follows closely the range of its natural host plants, Rhamnus catharticus L. and Frangula alnus Miller, suggesting that it is one of the few British butterflies that has a host‐limited distribution. In North Wales, this species has its range margin, and it was recorded only occasionally in a 35‐km² area prior to the 1980s. Frangula alnus bushes were planted in the area in about 1986, allowing the hypothesis that G. rhamni would expand its range following increased host plant availability to be tested. 2. From 1996 to 1998, the distribution of the butterfly and its host plants, R. catharticus (native), Rhamnus alaternus L. (introduced), and F. alnus (introduced to the area but native to Britain), was mapped in the study area. It was found that the butterfly was more widespread than any of its host plants. Frangula alnus was the most widespread of the host plants, and received most eggs, suggesting that the carrying capacity of the habitat would have increased substantially following the planting of this species. Gonepteryx rhamni was able to complete its lifecycle on both introduced species in the study area. 3. A mark–release–recapture study showed that adult G. rhamni moved an average of 512 m, and 50% of movements were further than 400 m; these values are underestimates. The relatively high mobility of this species suggests that it probably perceives host plants and nectar sources as resource patches (patchy population) in this fragmented landscape, and this population now represents a satellite population of the butterfly's main distribution in Britain. 4. The results presented here confirm empirically the host‐limited distribution of G. rhamni, which expanded following the planting of extra host plants.     

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.     

The genetic signature of rapid range expansion by flying squirrels in response to contemporary climate warming

Climate is an important factor limiting species distributions. Historic climate‐change related range movements have modified the genetic diversity of species by the merging and splitting of gene pools and by the effects associated with recurrent founder events. These effects are often inferred, either from retrospective analyses of current genetic patterns or from simulations. Rarely has it been possible for the population genetic effects of range expansion to be examined with contemporaneous demographic data. We characterized the genetic signature of rapid range expansion by southern flying squirrels (Glaucomys volans) and compared these results to a stationary population of the closely related northern flying squirrel (Glaucomys sabrinus) in Ontario, Canada. Samples were taken during an approximately 200 km range expansion by G. volans (1994–2003) and genotyped at 6 (G. sabrinus) and 8 (G. volans) microsatellite loci. For G. volans, but not G. sabrinus, we found evidence of a latitudinal gradient in allele frequencies and a decrease in allelic richness along the axis of expansion. We found no evidence of isolation‐by‐distance in either species or of genetic bottlenecks in the area of G. volans expansion. These results suggest that serial founder events can cause an immediate reduction in genetic diversity following rapid range expansion with high levels of gene flow giving rise to heterogeneity within what would classically be termed panmixia. Given the pace of anthropogenic climate change, and the increasing incidence of range movements in response, this may be an important, immediate consequence of climate change.     

The distributions of a wide range of taxonomic groups are expanding polewards

Evidence is accumulating of shifts in species' distributions during recent climate warming. However, most of this information comes predominantly from studies of a relatively small selection of taxa (i.e., plants, birds and butterflies), which may not be representative of biodiversity as a whole. Using data from less well‐studied groups, we show that a wide variety of vertebrate and invertebrate species have moved northwards and uphill in Britain over approximately 25 years, mirroring, and in some cases exceeding, the responses of better‐known groups.     

Rapid range expansion and community reorganization in response to warming

Species ranges are expected to expand along their cooler boundaries in response to rising temperatures associated with current global climate change. However, this ‘fingerprint’ of climate change is yet to be assessed for an entire flora. Here, we examine patterns of altitudinal range change in the complete native vascular flora of sub‐Antarctic Marion Island. We demonstrate a rapid mean upslope expansion in the flora since 1966, in response to 1.2 °C warming on the island. The 3.4±0.8 m yr^?1 (mean±SE) upslope expansion rate documented is amongst the highest estimates from partial floras. However, less than half of the species in the flora were responsible for the expansion trend, demonstrating that the global fingerprint of warming may be driven by a highly responsive subset of the species pool. Individual range expansion rates varied greatly, with species‐specific niche requirements explaining some of this variation. As a result of the idiosyncratic expansion rates, altitudinal patterns of species richness and community composition changed considerably, with the formation of no‐analog communities at high and intermediate altitudes. Therefore, both species‐ and community‐level changes have occurred in the flora of Marion Island over a relatively short period of rapid warming, demonstrating the sensitivity of high latitude communities to climate change. Patterns of change within this flora illustrate the range of variation in species responses to climate change and the consequences thereof for species distributions and community reorganization.     

The demography of range boundaries versus range cores in eastern US tree species

Regional species–climate correlations are well documented, but little is known about the ecological processes responsible for generating these patterns. Using the data from over 690 000 individual trees I estimated five demographic rates—canopy growth, understorey growth, canopy lifespan, understorey lifespan and per capita reproduction—for 19 common eastern US tree species, within the core and the northern and southern boundaries, of the species range. Most species showed statistically significant boundary versus core differences in most rates at both boundary types. Differences in canopy and understorey growth were relatively small in magnitude but consistent among species, being lower at the northern (average −17%) and higher at the southern (average +12%) boundaries. Differences in lifespan were larger in magnitude but highly variable among species, except for a marked trend for reduced canopy lifespan at the northern boundary (average −49%). Differences in per capita reproduction were large and statistically significant for some species, but highly variable among species. The rate estimates were combined to calculate two performance indices: R₀ (a measure of lifetime fitness in the absence of competition) was consistently lower at the northern boundary (average −86%) whereas Z^* (a measure of competitive ability in closed forest) showed no sign of a consistent boundary–core difference at either boundary.     

Historical changes in the phenology of British Odonata are related to climate

Responses of biota to climate change take a number of forms including distributional shifts, behavioural changes and life history changes. This study examined an extensive set of biological records to investigate changes in the timing of life history transitions (specifically emergence) in British Odonata between 1960 and 2004. The results show that there has been a significant, consistent advance in phenology in the taxon as a whole over the period of warming that is mediated by life history traits. British odonates significantly advanced the leading edge (first quartile date) of the flight period by a mean of 1.51 ±0.060 (SEM, n=17) days per decade or 3.08±1.16 (SEM, n=17) days per degree rise in temperature when phylogeny is controlled for. This study represents the first review of changes in odonate phenology in relation to climate change. The results suggest that the damped temperature oscillations experienced by aquatic organisms compared with terrestrial organisms are sufficient to evoke phenological responses similar to those of purely terrestrial taxa.     

The structure and function of the genitalia in the Libellulidae (Odonata)

An outline of the morphology of the secondary genitalia of libellulid dragonflies is given together with a more detailed treatment of the components of the fourth (distal) segment of the penis. The variations of penis structure in 70 species, representing ten of the 11 subfamilies, are surveyed and catalogued. There is remarkable diversity in the presence and form of penis structures, particularly those associated with the medial process, namely the flagelium, cornua and inner lobes. A comparable survey of females shows that they can be provisionally divided into ten types according to the number and size of their sperm-storage organs. Comparisons of penis structures with the sperm-storage organs of females suggests that in species whose females store large amounts of sperm, usually doing so in one or a pair of organs, males possess highly extensible, lobate and bristly penes, commonly equipped with cornua. By contrast, in those species in which the females store small volumes of sperm, usually doing so in three organs, the penes bear either flagella equipped with proximally directed spines or with large terminal barbs, or they have narrow, spiny inner-lobes which resemble flagella. Some functional interpretations are proposed in the light of sperm competition. The variability of genital structure can be interpreted adaptively and does not closely reflect phylogenetic relationships. Functionally convergent structures have evolved a number of times from different origins.     

Range shift promotes the formation of stable range edges

Aim I investigate the counter‐intuitive possibility that range shift promotes the formation of stable range edges. This might be expected because: (1) range‐shifting populations typically evolve increased dispersal on the expanding range edge; (2) increased dispersal steepens the relative slope of environmental gradients (gradients appear steeper to a more dispersive population); and (3) environmental gradients that are steep relative to dispersal encourage the formation of stable range edges (when gradients appear steep, adaptation on the range edge is swamped by maladapted genes). Methods I test the idea that populations take longer to evolve across an environmental gradient when those populations have already undergone a period of spread. I do this using an individual‐based coupled map lattice simulation, in which individuals carry heritable traits for dispersal probability and environment‐specific fitness. Results Numerous simulations across parameter space confirm that a period of range shift almost always results in a longer time to evolve through an environmental gradient. This occurs because of both the mechanism described above and the erosion of adaptive variation resulting from the serial foundering that occurs during range advance. Main conclusions This result suggests that species may often shift their range due to intrinsic changes in the population rather than extrinsic changes in the environment. The result also suggests a new mechanism regulating the speed of invasion, and sounds a cautionary note for climate change impacts: the longer a species tracks climate change, the less able it may be to track that change into the future.     

Flowering range changes across an elevation gradient in response to warming summer temperatures

Many studies have demonstrated plant response to warming temperatures, both as advancement in the timing of phenological events and in range shifts. Mountain gradients are ideal laboratories for studying species range changes. In this study of 363 plant species in bloom collected in five segments across a 1200 m (4158 ft) elevation gradient, we look for changes in species flowering ranges over a 20-year period. Ninety-three species (25.6%) exhibited a significant change in the elevation at which they flowered from the first half to the second half of the record, with many of these changes occurring at higher elevations. Most of the species exhibiting the changes were perennial plants. Interestingly, though many changes in flowering range were specific to higher elevations, range changes occurred all across the gradient. The changes reported in this study are concurrent with significant increases in summer temperatures across the region and are consistent with observed changes around the globe.     

Do species’ traits predict recent shifts at expanding range edges?

Although some organisms have moved to higher elevations and latitudes in response to recent climate change, there is little consensus regarding the capacity of different species to track rapid climate change via range shifts. Understanding species' abilities to shift ranges has important implications for assessing extinction risk and predicting future community structure. At an expanding front, colonization rates are determined jointly by rates of reproduction and dispersal. In addition, establishment of viable populations requires that individuals find suitable resources in novel habitats. Thus, species with greater dispersal ability, reproductive rate and ecological generalization should be more likely to expand into new regions under climate change. Here, we assess current evidence for the relationship between leading-edge range shifts and species' traits. We found expected relationships for several datasets, including diet breadth in North American Passeriformes and egg-laying habitat in British Odonata. However, models generally had low explanatory power. Thus, even statistically and biologically meaningful relationships are unlikely to be of predictive utility for conservation and management. Trait-based range shift forecasts face several challenges, including quantifying relevant natural history variation across large numbers of species and coupling these data with extrinsic factors such as habitat fragmentation and availability.     

Dynamics of range margins for metapopulations under climate change

We link spatially explicit climate change predictions to a dynamic metapopulation model. Predictions of species'' responses to climate change, incorporating metapopulation dynamics and elements of dispersal, allow us to explore the range margin dynamics for two lagomorphs of conservation concern. Although the lagomorphs have very different distribution patterns, shifts at the edge of the range were more pronounced than shifts in the overall metapopulation. For Romerolagus diazi (volcano rabbit), the lower elevation range limit shifted upslope by approximately 700 m. This reduced the area occupied by the metapopulation, as the mountain peak currently lacks suitable vegetation. For Lepus timidus (European mountain hare), we modelled the British metapopulation. Increasing the dispersive estimate caused the metapopulation to shift faster on the northern range margin (leading edge). By contrast, it caused the metapopulation to respond to climate change slower, rather than faster, on the southern range margin (trailing edge). The differential responses of the leading and trailing range margins and the relative sensitivity of range limits to climate change compared with that of the metapopulation centroid have important implications for where conservation monitoring should be targeted. Our study demonstrates the importance and possibility of moving from simple bioclimatic envelope models to second-generation models that incorporate both dynamic climate change and metapopulation dynamics.     

Combined effects of climate and biotic interactions on the elevational range of a phytophagous insect

1. The ranges of many species have expanded in cool regions but contracted at warm margins in response to recent climate warming, but the mechanisms behind such changes remain unclear. Particular debate concerns the roles of direct climatic limitation vs. the effects of interacting species in explaining the location of low latitude or low elevation range margins. 2. The mountains of the Sierra de Guadarrama (central Spain) include both cool and warm range margins for the black-veined white butterfly, Aporia crataegi, which has disappeared from low elevations since the 1970s without colonizing the highest elevations. 3. We found that the current upper elevation limit to A. crataegi's distribution coincided closely with that of its host plants, but that the species was absent from elevations below 900 m, even where host plants were present. The density of A. crataegi per host plant increased with elevation, but overall abundance of the species declined at high elevations where host plants were rare. 4. The flight period of A. crataegi was later at higher elevations, meaning that butterflies in higher populations flew at hotter times of year; nevertheless, daytime temperatures for the month of peak flight decreased by 6.2 degrees C per 1 km increase in elevation. 5. At higher elevations A. crataegi eggs were laid on the south side of host plants (expected to correspond to hotter microclimates), whereas at lower sites the (cooler) north side of plants was selected. Field transplant experiments showed that egg survival increased with elevation. 6. Climatic limitation is the most likely explanation for the low elevation range margin of A. crataegi, whereas the absence of host plants from high elevations sets the upper limit. This contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins. 7. Studies that have modelled distribution changes in response to climate change may have underestimated declines for many specialist species, because range contractions will be exacerbated by mismatch between the future distribution of suitable climate space and the availability of resources such as host plants.     

Rapid range expansion of a wing-dimorphic bush-cricket after the 2003 climatic anomaly

During recent decades, many species have responded to global warming by poleward range expansions. We require a better mechanistic understanding of the nature and extent of such processes to assess how climate change might affect biodiversity. Wing-dimorphic bush-crickets are excellent objects to study dispersal and colonization processes at the range margin because the long-winged morphs (macropters) represent dispersal units of otherwise flightless species. Moreover, these insects produce noisy songs and can easily be mapped. The present study comprised a detailed investigation of the population dynamics and genetics at the edge of the range of Roesel's bush-cricket, Metrioptera roeselii . We mapped the distribution of this insect in a previously unoccupied area of 185 km² and examined the genetic structure at the range margin using four polymorphic microsatellite loci. The results obtained demonstrate that the European heat wave in 2003 induced a strong immigration of macropters in the area stemming from multiple sources, whereas only few immigrants were recorded in the two subsequent years. Macropters were genotyped in a distance of up to 19.1 km from their origin, considerably exceeding the known dispersal distances for this species. Moreover, the data show that strong local founder effects are equalized on a large scale by the high number of immigrants from multiple sources. The present study demonstrates that macropters are of high significance for the range expansion of wing-dimorphic insects because a single-year climatic anomaly can induce strong dispersal processes.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 118–127.