Habitat amount and distribution modify community dynamics under climate change.

Habitat fragmentation may present a major impediment to species range shifts caused by climate change, but how it affects local community dynamics in a changing climate has so far not been adequately investigated empirically. Using long-term monitoring data of butterfly assemblages, we tested the effects of the amount and distribution of semi-natural habitat (SNH), moderated by species traits, on climate-driven species turnover. We found that spatially dispersed SNH favoured the colonisation of warm-adapted and mobile species. In contrast, extinction risk of cold-adapted species increased in dispersed (as opposed to aggregated) habitats and when the amount of SNH was low. Strengthening habitat networks by maintaining or creating stepping-stone patches could thus allow warm-adapted species to expand their range, while increasing the area of natural habitat and its spatial cohesion may be important to aid the local persistence of species threatened by a warming climate.

The decline of butterflies in Europe: Problems, significance, and possible solutions.

We review changes in the status of butterflies in Europe, focusing on long-running population data available for the United Kingdom, the Netherlands, and Belgium, based on standardized monitoring transects. In the United Kingdom, 8% of resident species have become extinct, and since 1976 overall numbers declined by around 50%. In the Netherlands, 20% of species have become extinct, and since 1990 overall numbers in the country declined by 50%. Distribution trends showed that butterfly distributions began decreasing long ago, and between 1890 and 1940, distributions declined by 80%. In Flanders (Belgium), 20 butterflies have become extinct (29%), and between 1992 and 2007 overall numbers declined by around 30%. A European Grassland Butterfly Indicator from 16 European countries shows there has been a 39% decline of grassland butterflies since 1990. The 2010 Red List of European butterflies listed 38 of the 482 European species (8%) as threatened and 44 species (10%) as near threatened (note that 47 species were not assessed). A country level analysis indicates that the average Red List rating is highest in central and mid-Western Europe and lowest in the far north of Europe and around the Mediterranean. The causes of the decline of butterflies are thought to be similar in most countries, mainly habitat loss and degradation and chemical pollution. Climate change is allowing many species to spread northward while bringing new threats to susceptible species. We describe examples of possible conservation solutions and a summary of policy changes needed to conserve butterflies and other insects.

Metapopulation dynamics in the butterfly Hipparchia semele changed decades before occupancy declined in The Netherlands.

The survival of many species in human-dominated, fragmented landscapes depends on metapopulation dynamics, i.e., on a dynamic equilibrium of extinctions and colonizations in patches of suitable habitat. To understand and predict distributional changes, knowledge of these dynamics can be essential, and for this, metapopulation studies are preferably based on long-time-series data from many sites. Alas, such data are very scarce. An alternative is to use opportunistic data (i.e., collected without applying standardized field methods), but these data suffer from large variations in field methods and search intensity between sites and years. Dynamic site-occupancy models offer a general approach to adjust for variable survey effort. These models extend classical metapopulation models to account for imperfect detection of species and yield estimates of the probabilities of occupancy, colonization, and survival of species at sites. By accounting for detection, they fully correct for among-year variability in search effort. As an illustration, we fitted a dynamic site-occupancy model to 60 years of presence-absence data (more precisely, detection-nondetection) of the heathland butterfly Hipparchia semele in The Netherlands. Detection records were obtained from a database containing volunteer-based data from 1950-2009, and nondetection records were deduced from database records of other butterfly species. Our model revealed that metapopulation dynamics of Hipparchia had changed decades before the species' distribution began to contract. Colonization probability had already started to decline from 1950 onward, but this was counterbalanced by an increase in the survival of existing populations, the result of which was a stable distribution. Only from 1990 onward was survival not sufficient to compensate for the further decrease of colonization, and occupancy started to decline. Hence, it appears that factors acting many decades ago triggered a change in the metapopulation dynamics of this species, which ultimately led to a severe decline in occupancy that only became apparent much later. Our study emphasizes the importance of knowledge of changes in survival and colonization of species in modern landscapes over a very long time scale. It also demonstrates the power of site-occupancy modeling to obtain important population dynamics information from databases containing opportunistic sighting records.

Declines in common, widespread butterflies in a landscape under intense human use.

Analyses of species' population losses typically show a dichotomy between strongly affected, rare, and localized species and apparently unaffected, common, and widespread species. We analyzed 16 years (1992-2007) of butterfly transect count data from The Netherlands in a reevaluation of the trends of common, widespread species. Fifty-five percent (11 of 20 species) of these species suffered severe declines in distribution and abundance. Overall, cumulative butterfly abundance declined by around 30%. Some of the species in decline used to be omnipresent in gardens and parks, and 2 of the species were previously considered agricultural pests. Based on their declines over the last 16 years, 2 of the 20 species (Lasiommata megera and Gonepteryx rhamni) reached endangered status in The Netherlands under the IUCN (International Union for Conservation of Nature) population-decline criterion, and 2 species (Inachis io and Thymelicus lineola) met vulnerable criterion. Butterflies in farmland, urban, and particularly woodland areas showed the largest decline in species abundance. The abundance of species associated with vegetation types found mainly in nature reserves (dunes, heathland, and, to a lesser extent, seminatural grassland) increased or remained stable. The decline of widespread species requires additional conservation strategies in the wider landscape.

Bias in phenology assessments based on first appearance data of butterflies.

Data on the first appearance of species in the field season are widely used in phenological studies. However, there are probabilistic arguments for bias in estimates of phenological change if sampling methods or population abundances change. We examined the importance of bias in three measures of phenological change: (1) the date of the first X appearances, (2) the date of the first Y% of all first appearances and (3) the date of the first Z% of the individuals observed during the entire flight period. These measures were tested by resampling the data of the Dutch Butterfly Monitoring Scheme and by simulations using artificial data. We compared datasets differing in the number of sampling sites, population abundance and the start of the observation period. The date of the first X appearances proved to be sensitive to the number of sampling sites. Both the date of the first X appearances and the date of the first Y% of all first appearances were sensitive to population trend. No such biases were found for estimates of the first Z% of the flight period, but all three measures were sensitive to changes in the start of the observation period. The conclusions were similar for both the study on butterfly data and the simulation study. Bias in phenology assessments based on first appearance data may be considerable and should no longer be ignored in phenological research.