Predicting range expansion of the map butterfly in Northern Europe using bioclimatic models

The two main goals of this study are: (i) to examine the range shifts of a currently northwards expanding species, the map butterfly (Araschnia levana), in relation to annual variation in weather, and (ii) to test the capability of a bioclimatic envelope model, based on broad-scale European distribution data, to predict recent distributional changes (2000–2004) of the species in Finland. A significant relationship between annual maximum dispersal distance of the species and late summer temperature was detected. This suggests that the map butterfly has dispersed more actively in warmer rather than cooler summers, the most notable dispersal events being promoted by periods of exceptionally warm weather and southerly winds. The accuracy of the broad-scale bioclimatic model built for the species with European data using Generalized Additive Models (GAM) was good based on split-sample evaluation for a single period. However, the model’s performance was poor when applied to predict range shifts in Finland. Among the many potential explanations for the poor success of the transferred bioclimatic model, is the fact that bioclimatic envelope models do not generally account for species dispersal. This and other uncertainties support the view that bioclimatic models should be applied with caution when they are used to project future range shifts of species.     

Role of larval host plants in the climate-driven range expansion of the butterfly Polygonia c-album

1. Some species have expanded their ranges during recent climate warming and the availability of breeding habitat and species' dispersal ability are two important factors determining expansions. The exploitation of a wide range of larval host plants should increase an herbivorous insect species' ability to track climate by increasing habitat availability. Therefore we investigated whether the performance of a species on different host plants changed towards its range boundary, and under warmer temperatures. 2. We studied the polyphagous butterfly Polygonia c-album, which is currently expanding its range in Britain and apparently has altered its host plant preference from Humulus lupulus to include other hosts (particularly Ulmus glabra and Urtica dioica). We investigated insect performance (development time, larval growth rate, adult size, survival) and adult flight morphology on these host plants under four rearing temperatures (18-28.5 degrees C) in populations from core and range margin sites. 3. In general, differences between core and margin populations were small compared with effects of rearing temperature and host plant. In terms of insect performance, host plants were generally ranked U. glabra > or = U. dioica > H. lupulus at all temperatures. Adult P. c-album can either enter diapause or develop directly and higher temperatures resulted in more directly developing adults, but lower survival rates (particularly on the original host H. lupulus) and smaller adult size. 4. Adult flight morphology of wild-caught individuals from range margin populations appeared to be related to increased dispersal potential relative to core populations. However, there was no difference in laboratory reared individuals, and conflicting results were obtained for different measures of flight morphology in relation to larval host plant and temperature effects, making conclusions about dispersal potential difficult. 5. Current range expansion of P. c-album is associated with the exploitation of more widespread host plants on which performance is improved. This study demonstrates how polyphagy may enhance the ability of species to track climate change. Our findings suggest that observed differences in climate-driven range shifts of generalist vs. specialist species may increase in the future and are likely to lead to greatly altered community composition.     

Impacts of landscape structure on butterfly range expansion

Since the 1940s, the distributions of several butterfly species have been expanding in northern Europe, probably in response to climate warming. We focus on the speckled wood butterfly Pararge aegeria in order to determine impacts of habitat availability on expansion rates. We analyse observed expansion rates since 1940 and also use a spatially explicit mechanistic model (MIGRATE) to simulate range expansion in two areas of the UK which differ in their distribution of breeding habitat (woodland). Observed and simulated expansion rates were in very close agreement but were 42%–45% slower in an area that had 24% less woodland. Unlike P. aegeria, the majority of butterfly species are not currently expanding, almost certainly because of lack of suitable habitat. Incorporating the spatial distribution of habitat into investigations of range changes is likely to be important in determining those species that can and cannot expand, and for predicting potential future range changes.     

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.     

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.     

The response of two butterfly species to climatic variation at the edge of their range and the implications for poleward range shifts

To predict changes in species' distributions due to climate change we must understand populations at the poleward edge of species' ranges. Ecologists generally expect range shifts under climate change caused by the expansion of edge populations as peripheral conditions increasingly resemble the range core. We tested whether peripheral populations of two contrasting butterflies, a small-bodied specialist (Erynnis propertius) and a large-bodied generalist (Papilio zelicaon), respond favorably to warmer conditions. Performance of populations related to climate was evaluated in seven peripheral populations spanning 1.2 degrees latitude (160 km) using: (1) population density surveys, an indirect measure of site suitability; and (2) organismal fitness in translocation experiments. There was evidence that population density increased with temperature for P. zelicaon whose population density declined with latitude in 1 of 3 sample years. On the other hand, E. propertius showed a positive relationship of population density with latitude, apparently unrelated to climate or measured habitat variables. Translocation experiments showed increased larval production at increased temperatures for both species, and in P. zelicaon, larval production also increased under drier conditions. These findings suggest that both species may increase at their range edge with warming but the preference for core-like conditions may be stronger in P. zelicaon. Further, populations of E. propertius at the range boundary may be large enough to act as sources of colonists for range expansions, but range expansion in this species may be prevented by a lack of available host plants further north. In total, the species appear to respond differently to climate and other factors that vary latitudinally, factors that will likely affect poleward expansion.     

Temperature constraints on foraging behaviour of male Alpine ibex (Capra ibex) in summer

In arctic and alpine environments, warm summer temperatures may force a reduction in foraging time of large herbivores, whose tolerance for heat is lower than for species adapted to warmer weather. We constructed time budgets for marked ibex (Capra ibex) males over two summers to test whether warm temperatures constrained foraging behaviour and forced altitudinal migrations. As daily temperature and solar radiation increased, feeding activity was reduced at midday and evening, but increased in the early morning, probably to anticipate for an expected reduction in foraging later in the day. With increasing temperature and solar radiation, ibex moved to higher elevations where they spent very little time feeding. Changes in forage quality and availability could not explain altitudinal migration. Temperatures above 15–20°C apparently result in heat discomfort in male Alpine ibex. As temperature and solar radiation increased, older and larger ibex spent less time feeding during daylight and showed a steeper decrease in feeding time than younger and smaller ibex. Larger males may be more sensitive to temperature and solar radiation, or may have more flexibility in allocating time to different activities, given their lower relative energetic requirements. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Evolution of flight morphology in a butterfly that has recently expanded its geographic range

Individuals colonizing unoccupied habitats typically possess characters associated with increased dispersal and, in insects, colonization success has been related to flight morphology. The speckled wood butterfly, Pararge aegeria, has undergone recent major expansions in its distribution: in the north of its range, P. aegeria has colonized many areas in north and east England, and in the south, it was first recorded on Madeira in 1976. We examined morphological traits associated with flight and reproduction in the northern subspecies tircis, and in the southern subspecies aegeria, from sites colonized about 20 years ago in northern England and on Madeira, respectively. Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were from individuals reared in a common environment and there were significant family effects in most of the variables measured. Compared with individuals from sites continuously occupied in recent history, colonizing individuals were larger (adult live mass). In the subspecies tircis, colonizing individuals also had relatively larger thoraxes and lower wing aspect ratios indicating that evolutionary changes in flight morphology may be related to colonization. However, sex by site interactions in analyses of thorax mass and abdomen mass suggest different selection pressures on flight morphology between the sexes in relation to colonization. Overall, the subspecies aegeria was smaller (adult live mass) and had a relatively larger thorax and wings, and smaller abdomen than subspecies tircis. Evolutionary changes in flight morphology and dispersal rate may be important determinants of range expansion, and may affect responses to future climate change. Received: 1 March 1999 / Accepted: 30 June 1999     

Modelling species' range shifts in a changing climate: the impacts of biotic interactions, dispersal distance and the rate of climate change

There is an urgent need for accurate prediction of climate change impacts on species ranges. Current reliance on bioclimatic envelope approaches ignores important biological processes such as interactions and dispersal. Although much debated, it is unclear how such processes might influence range shifting. Using individual-based modelling we show that interspecific interactions and dispersal ability interact with the rate of climate change to determine range-shifting dynamics in a simulated community with two growth forms--mutualists and competitors. Interactions determine spatial arrangements of species prior to the onset of rapid climate change. These lead to space-occupancy effects that limit the rate of expansion of the fast-growing competitors but which can be overcome by increased long-distance dispersal. As the rate of climate change increases, lower levels of long-distance dispersal can drive the mutualists to extinction, demonstrating the potential for subtle process balances, non-linear dynamics and abrupt changes from species coexistence to species loss during climate change.     

Predicting impacts of climate-induced range expansion: an experimental framework and a test involving key grazers on temperate rocky shores

Climate change has strong potential to modify the structure and functioning of ecosystems, but experimental field studies into its effects are rare. On rocky shores, grazing limpets strongly affect ecosystem structure and their distribution in NW Europe is changing in response to climate change. Three limpet species co-occur in SW Britain ( Patella vulgata, Patella ulyssiponensis and Patella depressa ) on open rock and in pools. Shores in Ireland are similar, but currently lack P. depressa . It is anticipated that P. depressa will expand its range into Ireland as the climate warms, but we currently lack an empirical basis to predict the consequences of this change. Recent studies show that increasing abundance of P. depressa on British shores has been accompanied by a decline of P. vulgata suggesting interspecific competition. In this study, a new experimental framework was used to examine the potential for P. depressa to affect P. vulgata on Irish shores. P. vulgata was experimentally transplanted into enclosures on open rock and in pools in both Ireland and Britain. In pools, treatments also included transplanted P. ulyssiponensis to mimic natural assemblages. Growth and mortality of P. vulgata were measured over 6 months with no differences between Ireland and Britain. In Britain, P. vulgata caged in pools with transplanted P. depressa and P. ulyssiponensis showed reduced growth, compared with when caged in pools with P. ulyssiponensis alone. There was no effect of P. depressa on the growth rate of P. vulgata on open rock. Results indicate that if the range of P. depressa extends into Ireland, it would reduce the growth of P. vulgata where it co-occurs with P. ulyssiponensis in pools. The framework used here provides a field-based approach that could be used to examine the impacts of climate-induced range expansions on the structure and functioning of other ecosystems.     

Delayed ovarian maturation in the butterfly Hipparchia semele as a possible response to summer drought

ABSTRACT.

• 1 Several references indicate that the period of flight of the European satyrine butterfly Hipparchia semele (L.) (Nymphalidae, Satyrinae) starts earlier in southern latitudes, where summers are longer and drier than in the north. However, summer drought has an adverse effect on the growth of grasses on which larval feeding depends. Growth of the grasses is delayed as long as the drought lasts.
• 2 From laboratory and field observations in a mid altitude area near the centre of the Iberian Peninsula, a mechanism that can be interpreted as an adjustment of this insect's life cycle to the host plant's phenology has been observed, i.e. delayed gonadal maturation of adult females. This delay is not associated with female diapause. Although the mean delay in oviposition after copulation was 43 days some captive females were able to oviposit much earlier, and this suggests variability in oviposition dates which might have an environmental or a genetic basis.
• 3 A mechanism of delayed ovarian maturation similar to that of H.semele is also known to occur in the satyrine Maniola jurtina (L.); it is suggested that this adaptation enables these species to occupy wider geographical ranges than other univoltine satyrines in Europe.

     

Poleward range expansion without a southern contraction in the ground beetle Agonum viridicupreum (Coleoptera, Carabidae)

We investigated the extent of poleward shifts in the distribution range of Agonum viridicupreum due to climate change in the western Palaearctic. Species' records were obtained from extensive literature sources as well as from collections, and consistent amateur entomologists' recordings. Within the general geographic range of the species, we analyzed in detail two parts of both, the northern and southern distribution range boundaries: (1 and 2) north-western Germany (leading or high-latitude edge), (3) Israel and (4) southern Italy (rear or low-latitude edge). Temporal changes in the occurrence data of the species indicated a northward shift of the leading edge of a minimum of 100 km within the last 50 to 100 years. In contrast, according to the data gathered, the rear edge has not changed during the last decades. Further studies are needed in order to fully understand the underlying mechanisms of the different behaviour of leading and rear range edges of Agonum viridicupreum in the current context of global change. Despite our incomplete understanding, chronosequences of the occurrence of the given species have the potential to optimize climate niche modelling to predict trends in the distribution range in the future.     

Foraging on buds and bark of mulberry trees by Japanese monkeys and their range utilization

The Japanese monkey (Macaca fuscata) winter range utilization and the effects of foraging on mulberry trees (Morus bombycis) were studied in the Shimokita Peninsula during four winter seasons. The monkeys ate mainly winter dormancy buds when they visited the mulberry tree clumps for the first time within the winter, but they ate mainly bark when they visited for the second or third times. In the areas utilized by the monkeys over the recent three years, the mulberry trees compensated for the decrease in their number of shoots by producing longer shoots with more buds against the monkey foraging. In the areas used every year for more than four years, however, the mulberry trees were unable to compensate for the foraging pressure. Thus, although the monkeys had apparently operated prudent herbivory within three years, they did not do so on a longer time-scale. They shifted their utilization ranges after having over-exploited the mulberry trees.     

Climate change and the potential for range expansion of the Lyme disease vector Ixodes scapularis in Canada

We used an Ixodes scapularis population model to investigate potential northward spread of the tick associated with climate change. Annual degree-days >0 degrees C limits for I. scapularis establishment, obtained from tick population model simulations, were mapped using temperatures projected for the 2020s, 2050s and 2080s by two Global Climate Models (the Canadian CGCM2 and the UK HadCM3) for two greenhouse gas emission scenario enforcings 'A2'and 'B2' of the Intergovernmental Panel on Climate Change. Under scenario 'A2' using either climate model, the theoretical range for I. scapularis establishment moved northwards by approximately 200 km by the 2020s and 1000 km by the 2080s. Reductions in emissions (scenario 'B2') had little effect on projected range expansion up to the 2050s, but the range expansion projected to occur between the 2050s and 2080s was less than that under scenario 'A2'. When the tick population model was driven by projected annual temperature cycles (obtained using CGCM2 under scenario 'A2'), tick abundance almost doubled by the 2020s at the current northern limit of I. scapularis, suggesting that the threshold numbers of immigrating ticks needed to establish new populations will fall during the coming decades. The projected degrees of theoretical range expansion and increased tick survival by the 2020s, suggest that actual range expansion of I. scapularis may be detectable within the next two decades. Seasonal tick activity under climate change scenarios was consistent with maintenance of endemic cycles of the Lyme disease agent in newly established tick populations. The geographic range of I. scapularis-borne zoonoses may, therefore, expand significantly northwards as a consequence of climate change this century.     

Plant–soil interactions in the expansion and native range of a poleward shifting plant species

Climate warming causes range shifts of many species toward higher latitudes and altitudes. However, range shifts of host species do not necessarily proceed at the same rates as those of their enemies and symbionts. Here, we examined how a range shifting plant species performs in soil from its original range in comparison with soil from the expansion range. Tragopogon dubius is currently expanding from southern into north-western Europe and we examined how this plant species responds to soil communities from its original and expansion ranges. We compared the performance of T. dubius with that of the closely related Tragopogon pratensis , which has a natural occurrence along the entire latitudinal gradient. Inoculation with the rhizosphere soil from T. dubius populations of the original range had a more negative effect on plant biomass production than inoculation with rhizosphere soil from the expansion range. Interestingly, the nonrange expander T. pratensis experienced a net negative soil effect throughout this entire range. The effects observed in this species pair may be due to release from soil born enemies or accumulation of beneficial soil born organisms. If this phenomenon applies broadly to other species, then range expansion may enable plants species to show enhanced performance.     

Changes in the high-mountain vegetation of the Central Iberian Peninsula as a probable sign of global warming

Aerial images of the high summits of the Spanish Central Range reveal significant changes in vegetation over the period 1957 to 1991. These changes include the replacement of high-mountain grassland communities dominated by Festuca aragonensis, typical of the Cryoro-Mediterranean belt, by shrub patches of Juniperus communis ssp. alpina and Cytisus oromediterraneus from lower altitudes (Oro-Mediterranean belt). Climatic data indicate a shift towards warmer conditions in this mountainous region since the 1940s, with the shift being particularly marked from 1960. Changes include significantly higher minimum and maximum temperatures, fewer days with snow cover and a redistribution of monthly rainfall. Total yearly precipitation showed no significant variation. There were no marked changes in land use during the time frame considered, although there were minor changes in grazing species in the 19th century. It is hypothesized that the advance of woody species into higher altitudes is probably related to climate change, which could have acted in conjunction with discrete variations in landscape management. The pronounced changes observed in the plant communities of the area reflect the susceptibility of high-mountain Mediterranean species to environmental change.     

Effects of elevated temperature and carbon dioxide on the nutritional quality of leaves of oak (Quercus robur L.) as food for the Winter Moth (Operophtera brumata L.)

1. Pedunculate Oak trees were grown in ambient and elevated temperatures and CO₂. Leaves were fed to Winter Moth caterpillars reared either in constant conditions or with the trees (caged or on-tree).
2. Caterpillars in constant conditions ate the same mass and produced the same mass of faeces whether fed elevated or ambient temperature leaves. However, less was assimilated from elevated leaves, resulting in lighter pupae and fewer, lighter eggs.
3. Caterpillars in constant conditions ate more and produced more faeces when fed elevated CO₂ leaves than when fed ambient CO₂ leaves, but the mass assimilated and pupal mass were unchanged.
4. Caged caterpillars reared with the trees from which they were fed had constant pupal mass in all treatments, but pupated earlier at elevated temperature. Pupal mass was also unaffected when caterpillars fed on the trees.
5. Nitrogen was reduced in both elevated temperature and elevated CO₂ leaves. Increased fibre in the former prevented increased consumption and resulted in reduced pupal mass and fecundity. Reduced fibre in the latter allowed increased consumption, resulting in pupae of normal mass.
6. Despite the clear effect of nutrient quality, experiments rearing caterpillars and trees together suggest that anticipated climatic change will have no nutritional effect on Winter Moth development.     

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.