Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

Habitat shifts of endangered species under altered climate conditions: importance of biotic interactions

Predicting changes in potential habitat for endangered species as a result of global warming requires considering more than future climate conditions; it is also necessary to evaluate biotic associations. Most distribution models predicting species responses to climate change include climate variables and occasionally topographic and edaphic parameters, rarely are biotic interactions included. Here, we incorporate biotic interactions into niche models to predict suitable habitat for species under altered climates. We constructed and evaluated niche models for an endangered butterfly and a threatened bird species, both are habitat specialists restricted to semiarid shrublands of southern California. To incorporate their dependency on shrubs, we first developed climate‐based niche models for shrubland vegetation and individual shrub species. We also developed models for the butterfly's larval host plants. Outputs from these models were included in the environmental variable dataset used to create butterfly and bird niche models. For both animal species, abiotic–biotic models outperformed the climate‐only model, with climate‐only models over‐predicting suitable habitat under current climate conditions. We used the climate‐only and abiotic–biotic models to calculate amounts of suitable habitat under altered climates and to evaluate species' sensitivities to climate change. We varied temperature (+0.6, +1.7, and +2.8 °C) and precipitation (50%, 90%, 100%, 110%, and 150%) relative to current climate averages and within ranges predicted by global climate change models. Suitable habitat for each species was reduced at all levels of temperature increase. Both species were sensitive to precipitation changes, particularly increases. Under altered climates, including biotic variables reduced habitat by 68–100% relative to the climate‐only model. To design reserve systems conserving sensitive species under global warming, it is important to consider biotic interactions, particularly for habitat specialists and species with strong dependencies on other species.     

Human impacts on environment–diversity relationships: evidence for biotic homogenization from butterfly species richness patterns

Aim Broad‐scale spatial variation in species richness relates to climate and physical heterogeneity but human activities may be changing these patterns. We test whether climate and heterogeneity predict butterfly species richness regionally and across Canada and whether these relationships change in areas of human activity. Location Canada. Methods We modelled the ranges of 102 butterfly species using genetic algorithms for rule‐set production (GARP). We then measured butterfly species richness and potentially important aspects of human activity and the natural environment. These were included in a series of statistical models to determine which factors are likely to affect butterfly species richness in Canada. We considered patterns across Canada, within predominantly natural areas, human‐dominated areas and particular ecozones. We examined independent observations of butterfly species currently listed under Canada's endangered species legislation to test whether these were consistent with findings from statistical models. Results Growing season temperature is the main determinant of butterfly species richness across Canada, with substantial contributions from habitat heterogeneity (measured using elevation). Only in the driest areas does precipitation emerge as a leading predictor of richness. The slope of relationships between all of these variables and butterfly species richness becomes shallower in human‐dominated areas, but butterfly richness is still highest there. Insecticide applications, habitat loss and road networks reduce butterfly richness in human‐dominated areas, but these effects are relatively small. All of Canada's at‐risk butterfly species are located in these human‐dominated areas. Main conclusions Temperature affects butterfly species richness to a greater extent than habitat heterogeneity at fine spatial scales and is generally far more important than precipitation, supporting both the species richness–energy and habitat heterogeneity hypotheses. Human activities, especially in southern Canada, appear to cause surprisingly consistent trends in biotic homogenization across this region, perhaps through range expansion of common species and loss of range‐restricted species.     

An elevational shift of cryophilous bryophytes in the last century – an effect of climate warming?

Aim To investigate altitudinal range shifts of bryophytes in Switzerland by comparing recent altitudinal distributions with historical distributions derived from herbarium specimens. Location Switzerland, covering 41,285 km² in Central Europe. Methods We used a dataset of 8520 herbarium specimens of 61 bryophyte species and compared altitudinal data between the two periods 1880–1920 and 1980–2005. The records we used were not specifically sampled for climatological analyses, but originate from non‐systematic fieldwork by various collectors. Historical and recent records were distributed all over Switzerland with occurrences in all major biogeographical areas. To account for different sampling efforts in the two time periods, different subsampling procedures were applied. Results Overall, we found a significant mean increase in altitude of 89 ± 29 m which was mainly driven by the cryophilous species (+222 ± 50 m). The mean increase in altitude of cryophilous species corresponds to a decadal upward shift of 24 m. The upper range limit of cryophilous species also increased by 189 ± 55 m, but there was no effect on the lower range limit. For intermediate and thermophilous species neither mean, nor upper or lower range limits changed. However, the proportion of records of thermophilous to cryophilous species increased considerably at lower altitudes, but levelled off above approximately 1800 m. Main conclusions We conclude that cryophilous bryophytes are expanding their range to higher elevations in Switzerland and that at lower elevations, a slow extinction process is going on, probably as a result of climate warming trends. The observed decadal upward shifts of cryophilous species closely match those reported from vascular plants in Europe and those expected, given recent estimates of climate warming trends. We emphasize that herbaria provide valuable data that can be used to detect ongoing changes in the distribution of species.     

Recent trends in butterfly populations from north-east Spain and Andorra in the light of habitat and climate change

Although butterfly declines have been reported across Europe, no assessment based on detailed quantitative data has ever been made for any extensive area in the Mediterranean Basin. In 1994, a Butterfly Monitoring Scheme was launched in Catalonia (NE Spain), and in 2005 a similar, albeit much smaller, scheme started in the neighbouring Pyrenean country of Andorra. Here we provide a first thorough assessment of butterfly trends in both areas for the last 15 years. Several patterns emerged, above all a worrying decline of a substantial part of the fauna. It was also evident that habitat specialists are experiencing greater declines than habitat generalists, thereby butterfly communities becoming progressively dominated by common species. However, habitat indicators based on characteristic species also revealed that trends are actually associated with habitat types, grassland and scrub specialists declining strongly but woodland specialists showing a marginal increase. These differences are certainly related to profound landscape changes, mainly a dramatic reduction of semi-natural grasslands and open Mediterranean scrub, and a major increase in woodlands. The general effect of climatic warming on butterfly populations was investigated by using the temperature community index (CTI) approach. The thermal structure of butterfly communities remained very stable over time, except in one case where, contrary to the expectations, a significant negative trend in the CTI was noted. However, this surprising result can be explained by taking into account the above-reported pattern of butterfly communities becoming dominated by common species, characterized by low thermal indices in comparison with declining Mediterranean specialists.     

Significant decrease in local butterfly community during the last 15 years in a calcareous hill of the middle Korea

Recently the significant decreases of species richness and abundance among terrestrial animals including butterflies are reported due to habitat change, overexploitation, and global warming. We compared the butterfly species composition and abundance from 1999 and 2014–2015 in a calcareous hill site of the middle part of Korea using a line transect method. There was a significant decrease in the number of individuals (abundance) and the number of species (richness) from 1999 to 2014–2015. This decrease was more prevalent among northern species than southern species, and the local extinct species were more prevalent among northern species, showing the influence of global warming on butterfly assemblages. However, no impact of habitat change was observed because of maintenance of the grasslands, which is caused by the dry soils of the calcareous region.     

Habitat associations of thermophilous butterflies are reduced despite climatic warming

Climate warming threatens the survival of species at their warm, trailing‐edge range boundaries but also provides opportunities for the ecological release of populations at the cool, leading edges of their distributions. Thus, as the climate warms, leading‐edge populations are expected to utilize an increased range of habitat types, leading to larger population sizes and range expansion. Here, we test the hypothesis that the habitat associations of British butterflies have expanded over three decades of climate warming. We characterize the habitat breadth of 27 southerly distributed species from 77 monitoring transects between 1977 and 2007 by considering changes in densities of butterflies across 11 habitat types. Contrary to expectation, we find that 20 of 27 (74%) butterfly species showed long‐term contractions in their habitat associations, despite some short‐term expansions in habitat breadth in warmer‐than‐usual years. Thus, we conclude that climatic warming has ameliorated habitat contractions caused by other environmental drivers to some extent, but that habitat degradation continues to be a major driver of reductions in habitat breadth and population density of butterflies.     

Long‐term monitoring of an amphibian community after a climate change‐ and infectious disease‐driven species extirpation

Infectious disease and climate change are considered major threats to biodiversity and act as drivers behind the global amphibian decline. This is, to a large extent, based on short‐term studies that are designed to detect the immediate and strongest biodiversity responses to a threatening process. What few long‐term studies are available, although typically focused on single species, report outcomes that often diverge significantly from the short‐term species responses. Here, we report the results of an 18‐year survey of an amphibian community exposed to both climate warming and the emergence of lethal chytridiomycosis. Our study shows that the impacts of infectious disease are ongoing but restricted to two out of nine species that form the community, despite the fact all species can become infected with the fungus. Climate warming appears to be affecting four out of the nine species, but the response of three of these is an increase in abundance. Our study supports a decreasing role of infectious disease on the community, and an increasing and currently positive effect of climate warming. We caution that if the warming trends continue, the net positive effect will turn negative as amphibian breeding habitat becomes unavailable as water bodies dry, a pattern that already may be underway.     

Climate and land-use interactively shape butterfly diversity in tropical rainforest and savanna ecosystems of southwestern China

Human-induced habitat conversion and degradation, along with accelerating climatic change, have resulted in considerable global biodiversity loss. Nevertheless, how local ecological assemblages respond to the interplay between climate and land-use change remains poorly understood. Here, we examined the effects of climate and land-use interactions on butterfly diversity in different ecosystems of southwestern China. Specifically, we investigated variation in the alpha and beta diversities of butterflies in different landscapes along human-modified and climate gradients. We found that increasing land-use intensity not only caused a dramatic decrease in butterfly alpha diversity but also significantly simplified butterfly species composition in tropical rainforest and savanna ecosystems. These findings suggest that habitat modification by agricultural activities increases the importance of deterministic processes and leads to biotic homogenization. The land-use intensity model best explained species richness variation in the tropical rainforest, whereas the climate and land-use intensity interaction model best explained species richness variation in the savanna. These results indicate that climate modulates the effects of land-use intensity on butterfly alpha diversity in the savanna ecosystem. We also found that the response of species composition to climate varied between sites: specifically, species composition was strongly correlated with climatic distance in the tropical rainforest but not in the savanna. Taken together, our long-term butterfly monitoring data reveal that interactions between human-modified habitat change and climate change have shaped butterfly diversity in tropical rainforest and savanna. These findings also have important implications for biodiversity conservation under the current era of rapid human-induced habitat loss and climate change.     

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.     

Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.     

Environmental controls on butterfly occurrence and species richness in Israel: The importance of temperature over rainfall

Butterflies are considered important indicators representing the state of biodiversity and key ecosystem functions, but their use as bioindicators requires a better understanding of how their observed response is linked to environmental factors. Moreover, better understanding how butterfly faunas vary with climate and land cover may be useful to estimate the potential impacts of various drivers, including climate change, botanical succession, grazing, and afforestation. It is particularly important to establish which species of butterflies are sensitive to each environmental driver.The study took place in Israel, including the West Bank and Golan Heights.To develop a robust and systematic approach for identifying how butterfly faunas vary with the environment, we analyzed the occurrence of 73 species and the abundance of 24 species from Israeli Butterfly Monitoring Scheme (BMS‐IL) data. We used regional generalized additive models to quantify butterfly abundance, and generalized linear latent variable models and generalized linear models to quantify the impact of temperature, rainfall, soil type, and habitat on individual species and on the species community.Species richness was higher for cooler transects, and also for hilly and mountainous transects in the Mediterranean region (rendzina and Terra rossa soils) compared with the coastal plain (Hamra soil) and semiarid northern Jordan Vale (loessial sierozem soil). Species occurrence was better explained by temperature (negative correlation) than precipitation, while for abundance the opposite pattern was found. Soil type and habitat were insignificant drivers of occurrence and abundance.Butterfly faunas responded very strongly to temperature, even when accounting for other environmental factors. We expect that some butterfly species will disappear from marginal sites with global warming, and a large proportion will become rarer as the region becomes increasingly arid.     

Influence of thermal conditions on habitat use by a rare spring‐emerging butterfly Euphydryas editha taylori

We may expect butterflies as ectotherms to have particularly active life‐history stages that occur in the warmest and lightest times of the year; however, there are temperate species that are active when climatic conditions seem unfavourable and photoperiod short, such as the Taylor's checkerspot (Euphydryas editha taylori). For such species, studies suggest that even subtle changes to microclimate can potentially impact populations. Thus, understanding how in situ variations in microclimate influence the Taylor's checkerspot butterfly could provide much needed insights into more effective management. We conducted a series of surveys that explored (i) adult habitat use, (ii) final instar larval distribution and (iii) adult movement up to and across site boundaries at two sites in Oregon, USA, in 2010 and 2011. We found that in situ habitat use by the Taylor's checkerspot butterfly was strongly influenced by microclimate. Both adult activities and final instar larvae distribution were clustered within the warmest areas of the sites. Moreover, adults did not use up to 59% and larva up to 90% of their sites, despite vegetation structure and composition being uniform. More specifically, butterfly habitat use increased with increasing ground temperatures, and we found that areas with the highest ground temperatures were more exposed to direct sunlight. Similarly, we found that butterflies tended to only move through sunlit site boundaries. We conclude that the Taylor's checkerspot is sensitive to changes in its thermal environment at fine spatial scales. Our results highlight the importance of microclimate as an indicator of habitat quality, and establishing the thermal criteria in which species of concern exists may provide valuable insights into the implications of climate change.     

Extracting useful data from imperfect monitoring schemes: endangered butterflies at San Bruno Mountain,San Mateo County,California (1982–2000) and implications for habitat management

Managers surveyed for sensitive butterfly species in the San Bruno Mountain Habitat Conservation Plan area between 1982 and 2000 using an opportunistic “wandering transect” method. To extract as much valuable information as possible from the data collected by this method we analyzed patterns of surveys and butterfly presence and absence within 250 m square cells gridded across the area within a Geographic Information System. While estimates of butterfly abundance were not possible, the data could be tested for trends in butterfly occupancy. For those cells surveyed during at least 10 years, no trends in the total number of occupied cells was evident for either Callippe silverspot or mission blue butterfly. There were cells, however, that showed positive or negative trends (P < 0.20) in occupancy for each species (Callippe silverspot: 14 positive, 15 negative, 6 cells occupied all years; mission blue butterfly: 40 positive, 40 negative, 2 cells occupied all years). We conclude that for the period 1982–2000 the population of each species was stable in overall total distribution, but indicate geographic areas of concern for each, specifically the edges of the northeast ridge for Callippe silverspot butterfly and the northwest of the study area for mission blue butterfly. Vegetation composition analysis using orthophotography with field corroboration indicates that those areas with declines in occupancy for these species experienced native coastal scrub succession and a corresponding loss in grassland butterfly habitat, while positive trending and stable cells had stable grassland proportions. Habitat managers at San Bruno Mountain should therefore incorporate programs for protecting grassland butterfly habitat not only from invasive weeds but also from succession to native coastal scrub. This approach illustrates the feasibility of using occupancy as an indicator to track butterfly status in a protected area even when suboptimal data collection methods are used, but the difficulties of using these data also reinforces the need for managers to devise monitoring schemes appropriate for their objectives before implementing them.     

Shift in Plant Species Composition Reveals Environmental Changes During the Last Decades: A Long-Term Study in Beech (Fagus sylvatica) Forests in Bavaria, Germany

Changes in vegetation composition due to the increasing temperatures in the past few decades have already been reported from several parts of Europe. It has been shown that single species move either northwards or to higher elevations. We expected that the species composition of forest stands should also have changed, i.e., an increase of thermophilous species. Another site factor changing for decades is nitrogen availability; we therefore also expected an increase of nitrophilous species, which was one main result in former long-term studies. We studied the species composition of beech (Fagus sylvatica) forests in southern Germany (Bavaria), comparing old (from 1949 to 1985) and young (2010) phytosociological relevés. Ellenberg indicator values representing plant species specific environmental factors combined with climatic data were used in a partial canonical correspondence analysis (pCCA) for vegetation comparisons. Changes in plant species composition were analyzed considering species frequency, distribution of Ellenberg indicator values, shares of i) non-native plants and ii) tree, shrub and herbaceous species. Contrary to our expectations, global warming in Bavaria during the past decades resulted only in an explained dispersion of 5 % in the species composition. On the species level, an overall increase of thermophilous walnut tree saplings (Juglans regia) was conspicuous in some study areas. Nitrophilous species, however, generally increased in frequency throughout the study areas. Throughout Bavaria the most significant change was a striking increase of juvenile tree species and a decrease of herbaceous species. Up to now the increased nitrogen input into forests had a stronger provable influence on species composition shift than the global warming of the last decades. Additionally general changes in forest management also had effects on forest species. Therefore community reorganization mirroring temperature factors in beech forests in central Europe seems to be only at its very beginning.     

Terrestrial mesofauna in above- and below-ground habitats: Taylor Valley,Antarctica

In the McMurdo Dry Valleys region of Antarctica, above-ground production is often limited to mosses and algae that occur near seasonally available liquid water such as ephemeral streams and ice-covered lakes. Compared to surrounding dry soils these critical transition zones are highly productive and harbor a more diverse assemblage of soil animals, including rotifers, tardigrades, nematodes and microarthropods. Current cooling trends punctuated by warming events, and predicted future climate warming are expected to affect the hydrology of this region and thereby biodiversity and ecosystem functioning. Above-ground communities are exposed to more variable temperature, relative humidity and greater UV radiation, and may be more vulnerable to climate change than sediments beneath, which are buffered from short-term changes. In this study, we compared above- and below-ground communities associated with either moss or cyanobacterial mats along glacial-fed streams and lakes differing in biological complexity (diversity, productivity and habitat suitability). All groups of soil fauna were more abundant in the above-ground material compared to the sediment beneath. Common indicators of habitat suitability (chlorophyll a, soil pH, soil salinity, and soil nitrogen) did not differ between vegetation types but were significantly different among sites. Variables most correlated with invertebrate abundances were sediment salinity, chlorophyll a content and nitrogen concentration. The McMurdo Dry Valleys are expected to become warmer and wetter as a result of climate change. This will likely increase the area of suitable habitat for most soil animals as areas of liquid water potentially increase and become available for longer periods of time.     

Extinction vulnerability of tropical montane endemism from warming and upslope displacement: a preliminary appraisal for the highest massif in Madagascar

One of the predicted biological responses to climate warming is the upslope displacement of species distributions. In the tropics, because montane assemblages frequently include local endemics that are distributed close to summits, these species may be especially vulnerable to experiencing complete habitat loss from warming. However, there is currently a dearth of information available for tropical regions. Here, we present a preliminary appraisal of this extinction threat using the herpetological assemblage of the Tsaratanana Massif in northern Madagascar (the island's highest massif), which is rich with montane endemism. We present meteorological evidence (individual and combined regional weather station data and reanalysis forecast data) for recent warming in Madagascar, and show that this trend is consistent with recent climate model simulations. Using standard moist adiabatic lapse rates, these observed meteorological warming trends in northern Madagascar predict upslope species displacement of 17–74 m per decade between 1993 and 2003. Over this same period, we also report preliminary data supporting a trend for upslope distribution movements, based on two surveys we completed at Tsaratanana. For 30 species, representing five families of reptiles and amphibians, we found overall mean shifts in elevational midpoint of 19–51 m upslope (mean lower elevation limit 29–114 m; mean upper elevation limit ?8 to 53 m). We also found upslope trends in mean and median elevational observations in seven and six of nine species analysed. Phenological differences between these surveys do not appear to be substantial, but these upslope shifts are consistent with the predictions based on meteorological warming. An elevational range displacement analysis projects complete habitat loss for three species below the 2 °C ‘dangerous’ warming threshold. One of these species is not contracting its distribution, but the other two were not resampled in 2003. A preliminary review of the other massifs in Madagascar indicates potential similar vulnerability to habitat loss and upslope extinction. Consequently, we urgently recommend additional elevational surveys for these and other tropical montane assemblages, which should also include, when possible, the monitoring of local meteorological conditions and habitat change.     

Beyond climate envelopes: effects of weather on regional population trends in butterflies

Although the effects of climate change on biodiversity are increasingly evident by the shifts in species ranges across taxonomical groups, the underlying mechanisms affecting individual species are still poorly understood. The power of climate envelopes to predict future ranges has been seriously questioned in recent studies. Amongst others, an improved understanding of the effects of current weather on population trends is required. We analysed the relation between butterfly abundance and the weather experienced during the life cycle for successive years using data collected within the framework of the Dutch Butterfly Monitoring Scheme for 40 species over a 15-year period and corresponding climate data. Both average and extreme temperature and precipitation events were identified, and multiple regression was applied to explain annual changes in population indices. Significant weather effects were obtained for 39 species, with the most frequent effects associated with temperature. However, positive density-dependence suggested climatic independent trends in at least 12 species. Validation of the short-term predictions revealed a good potential for climate-based predictions of population trends in 20 species. Nevertheless, data from the warm and dry year of 2003 indicate that negative effects of climatic extremes are generally underestimated for habitat specialists in drought-susceptible habitats, whereas generalists remain unaffected. Further climatic warming is expected to influence the trends of 13 species, leading to an improvement for nine species, but a continued decline in the majority of species. Expectations from climate envelope models overestimate the positive effects of climate change in northwestern Europe. Our results underline the challenge to include population trends in predicting range shifts in response to climate change.     

Effect of grazing intensity on the interaction between Shijimiaeoides divinus asonis (Matsumura, 1929) and its attendant ants

This study was carried out in the natural herbaceous grassland of Mt. Aso, which had been almost completely grazed, and which is subjected to routine grassland burning every spring (February or March) to conserve Shijimiaeoides divinus asonis (Matsumura, 1929). We clarify that ants protect the larvae of this butterfly and evaluate the effects of grazing intensity on the attendant ant population. The results obtained are summarized as follows: (i) Five species of attendant ant were identified, with the dominant species being Formica japonica (Motschulsky, 1866) and Camponotus japonicus (Mayr, 1866). (ii) The number of attendant ants decreased at night time; however, only one or two ants attended the larva until the following morning. (iii) Ten species of insect excluding ants and three species of spiders that approached the larva were recorded on the larval host‐plant. Formica japonica and C. japonicus fought off most newcomers of other insects including the natural enemies of these butterfly larvae. (iv) The number of ants in this butterfly habitat under regular grazing intensity was significantly higher than during low grazing intensity and non‐grazing periods. (v) A positive correlation was found between the number of attendant ants and the number of butterfly larvae on the host‐plant. We concluded that the interaction between this butterfly and attendant ants is one of facultative mutualism because the attendant ants protect the butterfly larva. Therefore, the numbers of this butterfly species may decrease if the number of attendant ants decreases due to the cessation of pasturage.     

Population resilience to an extreme drought is influenced by habitat area and fragmentation in the local landscape

Most studies on the biological impact of climate change have focussed on incremental climate warming, rather than extreme events. Yet responses of species’ populations to climatic extremes may be one of the primary drivers of ecological change. We assess the resilience of individual populations in terms of their sensitivity to‐ and ability to recover from‐ environmental perturbation. We demonstrate the method using a model species, the ringlet butterfly Aphantopus hyperantus, and analyse the effects of an extreme drought event using data from 79 British sites over 10 yr. We find that populations crashed most severely in drier regions but, additionally, the landscape structure around sites influenced population responses. Larger and more connected patches of woodland habitat reduced population sensitivity to the drought event and also facilitated faster recovery. Having enough, sufficiently connected habitat appears essential for species’ populations to be resilient to the increased climatic variability predicted under future scenarios.