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.     

Field transplants reveal summer constraints on a butterfly range expansion

The geographic ranges of most species are expected to shift to higher elevations and latitudes in response to global warming. But species react to specific environmental changes in individualistic ways, and we are far from a detailed understanding of range-shifts. Summer temperature often limits the ranges of insects and plants, so many range-shifts are expected to track summer warming. I explore this potential range-limiting factor in a case study of a northwardly expanding American butterfly, Atalopedes campestris (Lepidoptera, Hesperiidae). This species has recently colonized the Pacific Northwest, USA, where the mean annual temperature has risen 0.8–1.8°C over the past 100 years. Using field transplant experiments across the current range edge, I measured development time, survivorship, fecundity and predation rates along a naturally occurring thermal gradient of 3°C. Development time was significantly slower outside the current range in eastern Washington (WA), as expected because of cooler temperatures there. Slower development would reduce the number of generations possible per year outside the current range, dramatically lowering the probability that a population could survive there. Differences in survivorship, fecundity and predation rate across the range edge were not significant. The interaction between summer and winter temperature appears to be crucial in defining the current range limit. The estimated difference in temperature required to affect the number of generations is greater than the extent of summer warming observed over the past century, however, and thus historically winter temperature alone probably limited the range in southeastern WA. Nonetheless, extraordinarily warm summers may have improved colonization success, increasing the probability of a range expansion. These results suggest that extreme climatic events may influence rates of response to long-term climate change. They also demonstrate that range-limiting factors can change over time, and that the asymmetry in seasonal warming trends will have biological consequences.     

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.     

How interactions between animal movement and landscape processes modify local range dynamics and extinction risk

Forecasts of range dynamics now incorporate many of the mechanisms and interactions that drive species distributions. However, connectivity continues to be simulated using overly simple distance-based dispersal models with little consideration of how the individual behaviour of dispersing organisms interacts with landscape structure (functional connectivity). Here, we link an individual-based model to a niche-population model to test the implications of this omission. We apply this novel approach to a turtle species inhabiting wetlands which are patchily distributed across a tropical savannah, and whose persistence is threatened by two important synergistic drivers of global change: predation by invasive species and overexploitation. We show that projections of local range dynamics in this study system change substantially when functional connectivity is modelled explicitly. Accounting for functional connectivity in model simulations causes the estimate of extinction risk to increase, and predictions of range contraction to slow. We conclude that models of range dynamics that simulate functional connectivity can reduce an important source of bias in predictions of shifts in species distributions and abundances, especially for organisms whose dispersal behaviours are strongly affected by landscape structure.     

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.     

Modelling single nucleotide effects in phosphoglucose isomerase on dispersal in the Glanville fritillary butterfly: coupling of ecological and evolutionary dynamics

Dispersal comprises a complex life-history syndrome that influences the demographic dynamics of especially those species that live in fragmented landscapes, the structure of which may in turn be expected to impose selection on dispersal. We have constructed an individual-based evolutionary sexual model of dispersal for species occurring as metapopulations in habitat patch networks. The model assumes correlated random walk dispersal with edge-mediated behaviour (habitat selection) and spatially correlated stochastic local dynamics. The model is parametrized with extensive data for the Glanville fritillary butterfly. Based on empirical results for a single nucleotide polymorphism (SNP) in the phosphoglucose isomerase (Pgi) gene, we assume that dispersal rate in the landscape matrix, fecundity and survival are affected by a locus with two alleles, A and C, individuals with the C allele being more mobile. The model was successfully tested with two independent empirical datasets on spatial variation in Pgi allele frequency. First, at the level of local populations, the frequency of the C allele is the highest in newly established isolated populations and the lowest in old isolated populations. Second, at the level of sub-networks with dissimilar numbers and connectivities of patches, the frequency of C increases with decreasing network size and hence with decreasing average metapopulation size. The frequency of C is the highest in landscapes where local extinction risk is high and where there are abundant opportunities to establish new populations. Our results indicate that the strength of the coupling of the ecological and evolutionary dynamics depends on the spatial scale and is asymmetric, demographic dynamics having a greater immediate impact on genetic dynamics than vice versa.     

Population structure of spotted salamanders (Ambystoma maculatum) in a fragmented landscape

Understanding the impacts of landscape-level processes on the population biology of amphibians is critical, especially for species inhabiting anthropogenically modified landscapes. Many pond-breeding amphibians are presumed to exist as metapopulations, but few studies demonstrate the extent and consequences of this metapopulation structure. Gene flow measures may facilitate the construction of more realistic models of population structure than direct measures of migration. This is especially true for species that are cryptic, such as many amphibians. We used eight polymorphic microsatellite loci to determine the genetic population structure of spotted salamanders ( Ambystoma maculatum ) breeding at 17 ponds in northeastern Ohio, a landscape fragmented by roads, agriculture, urban areas and the Cuyahoga River. Using a variety of analyses (Bayesian clustering, F -statistics, AMOVA) we generated a model of salamander population genetic structure. Our data revealed patterns of genetic connectivity that were not predicted by geographical distances between ponds (no isolation by distance). We also tested for a relationship between population structure and several indices of landscape resistance, but found no effect of potential barriers to dispersal on genetic connectivity. Strong overall connectivity among ponds, despite the hostile habitat matrix, may be facilitated by a network of riparian corridors associated with the Cuyahoga River; however, high gene flow in this system may indicate a general ability to disperse and colonize beyond particular corridors.     

Population genetic structure and dispersal across a fragmented landscape in cerulean warblers (<Emphasis Type="Italic">Dendroica cerulea</Emphasis>)

Cerulean warblers (Dendroica cerulea) have experienced significant declines across their breeding range and presently exist in disjunct populations, largely because of extensive loss and fragmentation of their breeding and wintering habitat. Despite this overall decline, a recent north-eastern expansion of the breeding range has been proposed, and some researchers have suggested that the eastern Ontario population may be acting as a source population maintaining sink populations elsewhere. However, little is known about either the geographic distribution of genetic variation or dispersal in these birds. We assayed variation in five microsatellite loci and a 366 base-pair fragment of the mitochondrial control region among 154 cerulean warblers from five populations throughout the breeding range. No evidence of population genetic structure was found. Assignment tests suggested that six individuals were either inter-population migrants or descendants of recent migrants. The lack of population genetic structure is probably due to a combination of historical association and contemporary dispersal. Population decline does not appear to have reduced genetic variation yet. Overall results suggest that cerulean warblers from Ontario, Illinois, Arkansas and Tennessee should be considered a single genetic management unit for conservation.     

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.     

Population structure of a large blue butterfly and its specialist parasitoid in a fragmented landscape

Habitat fragmentation may interrupt trophic interactions if herbivores and their specific parasitoids respond differently to decreasing connectivity of populations. Theoretical models predict that species at higher trophic levels are more negatively affected by isolation than lower trophic level species. By combining ecological data with genetic information from microsatellite markers we tested this hypothesis on the butterfly Maculinea nausithous and its specialist hymenopteran parasitoid Neotypus melanocephalus. We assessed the susceptibility of both species to habitat fragmentation by measuring population density, rate of parasitism, overall genetic differentiation (theta(ST)) and allelic richness in a large metapopulation. We also simulated the dynamics of genetic differentiation among local populations to asses the relative effects of migration rate, population size, and haplodiploid (parasitoid) and diploid (host) inheritance on metapopulation persistence. We show that parasitism by N. melanocephalus is less frequent at larger distances to the nearest neighbouring population of M. nausithous hosts, but that host density itself is not affected by isolation. Allelic richness was independent of isolation, but the mean genetic differentiation among local parasitoid populations increased with the distance between these populations. Overall, genetic differentiation in the parasitoid wasp was much greater than in the butterfly host and our simulations indicate that this difference is due to a combination of haplodiploidy and small local population sizes. Our results thus support the hypothesis that Neotypus parasitoid wasps are more sensitive to habitat fragmentation than their Maculinea butterfly hosts.     

Intraspecific variation in habitat availability among ectothermic animals near their climatic limits and their centres of range

1. In a modelling exercise, the quantity and distribution of habitat patches within a heathland biotope for four ectothermic heathland animals (silver-studded blue butterfly [ Plebejus argus ] , a red ant [ Myrmica sabuleti ] , heath grasshopper [ Chorthippus vagans ] and sand lizard [ Lacerta agilis ]) were compared in space and time assuming two climates: that experienced at the northern edge of the species' ranges and that 300–400 km further south, where mean summer temperatures are 2–3 °C warmer.
2. Habitats both at the northern edge of their ranges and 300–400 km further south for the four species were defined qualitatively from existing sources and then expressed quantitatively in terms of the attributes recorded in the Dorset Heathland Survey. The Survey was then used as a GIS to map the occurrence of the habitats of the four species under two climates and a decade apart.
3. The model predicts that an increase of 2–3 °C can result in a large increase in the area of habitat available to these north temperate species, that the length of time that individual patches of successional habitat may be occupied increases and that the distance between habitat patches within the biotope decreases.
4. The warmer conditions should result in a more stable metapopulation structure for P. argus , with fewer metapopulations existing in the landscape but each, on average, containing a greater number of larger and more stable constituent populations.
5. These predictions are of significance to ectothermic species which currently live at the northern limits of their ranges in the British Isles. The reverse effect is likely for species at the southern limits of their ranges. Conservationists who wish to maintain the status quo may be able to reduce some effects of these changes by appropriate habitat management.